importance of studying mycology essay

The importance of fungi and mycology for addressing major global challenges

December 2014
IMA Fungus 5(2)
CC BY-NC-ND 3.0

"BioEconomy Research & Advisory" & associated Technical University of Denmark

Abstract and Figures

Discover the world's research

25+ million members
160+ million publication pages
2.3+ billion citations
U.M.M.P.K. Nawarathne

A.D. Bellere

Pragya Tiwari
Kyeung Il Park
Konul F. Bakshaliyeva
Nizami Rza Namazov
Sabia M Jabrailzade

Rashmi Mathur

John Zothanzama

Baiaan H. Alsaadi

Ana Mariana Chirilă

Abdurrahman GÜMÜŞ

Gui H Hofstetter

John W. Taylor

R.A. Shoemaker

Ignacio H. Chapela

Ailsa D. Hocking

L.B.S. Hansen

ENZYME MICROB TECH

Morten Nedergaard Grella

Recruit researchers
Join for free
Login Email Tip: Most researchers use their institutional email address as their ResearchGate login Password Forgot password? Keep me logged in Log in or Continue with Google Welcome back! Please log in. Email · Hint Tip: Most researchers use their institutional email address as their ResearchGate login Password Forgot password? Keep me logged in Log in or Continue with Google No account? Sign up

Mycology What is it? Classification, in Medicine and How to Culture

Definition: what is mycology.

Essentially, mycology is the study of fungi. Here, mycologists directly focus on the taxonomy, genetics, application as well as many other characteristics of this group of organisms.

Currently, over 50,000 species of fungi have been identified in different environments across the globe. While some are free-living and have no impact on human beings (and other animals), some are either beneficial or harmful making it necessary to study and understand them.

Classification

Based on sexual reproduction:.

· Glomeromycota - Members of this phylum form a symbiotic relationship with various plants and trees. As such, they are not necessarily parasites that fully depend on the host. The majority of the species reproduce asexually.

A few species, however, are parasitic in nature. Zygomycetes reproduce asexually and produce zygospores. Although species like R. stolonifer obtain their energy from decaying matter (e.g bread) they cause food spoilage and can cause diseases.

· Basidiomycota - Known as club fungi, this phylum is largely composed of mushrooms, smut fungi, and rust. The majority of basidiomycetes are pathogens of grains and tend to produce sexually. Spores of basidiomycetes are referred to as basidiospores.

· Ascomycota - Also known as sac fungi, members of Ascomycota include mushrooms, yeast, truffles, and morels. A majority are filamentous that either exist as parasites or saprophytes. However, they can also form symbiotic relationships with other organisms. They produce sexually and asexually.

· Chytridiomycota - Members of this group are also known as chytrids and produce spores known as zoospores. In aquatic environments, these organisms move using a flagellum and reproduce asexually.

Mold Fungi (Filamentous/Hyphal)

Common molds include:

For molds, these filamentous structures serve different functions including reproduction and absorbing nutrients among others.

Geographical Classification

According to mycology studies, some groups of fungi have been shown to be restricted to specific geographic regions. It has been shown that while fungi species are distributed across the globe, their diversity in different regions are influenced by such factors as soil chemistry and the climate among others. For this reason, different species of fungi can be associated with certain geographical locations.

Epidemiologic Grouping

Medical mycology, culture of candida albicans.

Requirements

Before starting the actual procedure, it is important to ensure that all material used are clean and sterilized. Although an isolation medium is used for this technique, this is an important step in all culture protocols.

Sample collection and preparation

· Pour the culture medium (SDA ) onto a clean Petri dish

· Using a sterile wire loop of the cotton swab, inoculate the sample on the agar medium using the spiral plating system

· Incubate the culture at 37 degrees Celsius for about one to two days

Observation

Mold fungus: aspergillus flavus.

The fungal sample for this procedure may be obtained from poultry feed samples

See also: Bacteriology , Microbiology , Phycology , Parasitology , Virology , Nematology , Immunology , Protozoology

Return to learning more about Fungi

Return to Mold under the Microscope

Sridhar Rao. (2006). Introduction to Mycology.

MicroscopeMaster.com is a participant in the Amazon Services LLC Associates Program, an affiliate advertising program designed to provide a means to earn fees by linking to Amazon.com and affiliated sites.

Recent Articles

Deltaproteobacteria - examples and characteristics, chemoorganotrophs - definition, and examples, betaproteobacteria – examples, characteristics and function.

9 October 2015

Celebrating the importance of mycological research

Although kew is mostly known for its work on plants, a large part of the research is focused on the diversity and importance of fungi. pepijn kooij explains how mycologists at kew are working to understand the role of fungi in plant diversity..

By Dr Pepijn Kooij , Dr Tuula Niskanen and Dr Laura Martinez-Suz

Estimating numbers

There have been many estimates of the total number of fungi species, reaching up to 6 million. However, the widely accepted range is “at least 1.5, but probably as many as 3 million”, far outnumbering flowering plants (Kew’s estimates go as far as 400,000 species of flowering plants).

As only around 100,000 species of fungi have so far been described worldwide, one of the goals for the mycologists at Kew is to document the missing diversity, but also to investigate their importance for ecosystems as a whole. Key to this task is the Fungarium, containing over 1.25 million specimens of dried fungi, the largest collection of fungal specimens in the world today.

How to investigate fungal diversity

As part of the fungal diversity research at Kew, Tuula Niskanen, one of Kew’s mycologists, studies the diversity and evolution of mushrooms, with a special interest in webcaps (Cortinarius). Webcaps are the most species-rich genus of the Agaricales, the gilled mushrooms, with a worldwide distribution. They are important ectomycorrhizal fungi and play a significant role in the nutrient economy of forest trees. However, they are still very poorly known and many species are not yet described and named. Even in Britain many new species remain to be discovered.

Knowledge of the evolutionary history of fungi provides the means for a better understanding of the current diversity and distribution of species. It also sets a baseline for further studies of diverse evolutionary questions, which cannot advance without the fungal specimens. The vast collections of fungi in Kew and other institutes world-wide provide a significant resource for these studies, which in turn help new expeditions to be targeted to previously unexplored areas.

Plant-fungal symbiosis and environmental change

Globally, plant-fungal partnerships underpin terrestrial ecosystems. "Fungus-roots", or mycorrhizas (myco= fungus, rhiza=root), are ancient, obligate and ubiquitous mutualisms between the vast majority of plants and members of several fungal phyla to exchange carbon derived from photosynthesis for fungal-acquired soil nutrients. We can say that most plants don’t have roots, they have mycorrhizas! For example, tree roots in boreal, temperate and some tropical forests form ectomycorrhizas (ecto=outside), which envelop root tips like gloves, and play crucial ecological roles by determining the nutrient acquisition and drought tolerance of trees. Due to their distinctive ecological niche, mycorrhizal fungi are at particular risk to changes in either their soil environment or host carbon allocation.

Global change is one of the biggest threats to organismal and functional diversity, yet little is known about its potential impacts on plant-fungal interactions. Fungi with different soil exploration types (such as those specialised for long-, medium or short-distance water and nutrient transport) respond strongly to pollution causing eutrophication and acidification in European forests. Kew researchers Laura Martinez-Suz, Martin I. Bidartondo, Sietse van der Linde and William Rimington (Imperial College London) study the evolution, diversity, ecology and distribution of mycorrhizal fungi and their environmental drivers in different ecosystems. Research on this functional guild of fungi is important because, even though they are still largely neglected when it comes to conservation, they are likely to determine the resilience of ecosystems to environmental change.

Fungus-farming ants

Humans have been domesticating crops for approximately 10,000 years. Ants, however, have been growing fungal crops for 50 million years and are considered to be the oldest farmers in the world. Much like humans did, for instance with bananas, the most recently derived group of these ants, the leaf-cutting ants, created a polyploid (with more than two sets of chromosomes) fungal lineage, while maintaining this crop without sexual reproduction or mushroom growth. This polyploidisation may enhance traits the ants benefit from, such as increased nutrition or resistance to disease.

My research focuses on the mechanisms that lie behind the maintenance of the asexuality of these fungi. Even though there doesn’t seem to be any genetic mixing, the fungi cultivated by ants have a high diversity with a still unknown number of species. By investigating fungarium specimens from free-living fungi closely related to the ant crops in the genera Leucoagaricus and Leucocoprinus, it will be possible to find the closest wild relatives from which the ants derived their cultivars. This will in turn help to understand the evolution of this enigmatic mutualism of ants and fungi, and potentially lend insight into our own agricultural symbioses.

References

Kooij, P.W., Aanen, D.K., Schiøtt, M., & Boomsma, J.J. (2015). Evolutionarily advanced ant farmers rear polyploid crops. Journal of Evolutionary Biology, DOI: 10.1111/jeb.12718.

Suz, L.M., Barsoum, N., Benham, S., Dietrich, H.P., Fetzer, K.D., Fischer, R., García, P., Gehrman, J., Kristöfel, F., Manninger, M., Neagu, S., Nicolas, M., Oldenburger, J., Raspe, S., Sánchez, G., Schröck, H.W., Schubert, A., Verheyen, K., Verstraeten, A., & Bidartondo, M.I. (2014). Environmental drivers of ectomycorrhizal communities in Europe's temperate oak forests. Molecular Ecology 23(22): 5628-5644.

Suz, L.M., Barsoum, N., Benham, S., Cheffings, C., Cox, F., Hackett, L., Jones, A.G., Mueller, G.M., Orme, D., Seidling, W., Van der Linde, S., & Bidartondo M.I. (2015). Monitoring ectomycorrhizal fungi at large scales for science, forest management, fungal conservation and environmental policy. Annals of Forest Science. DOI 10.1007/s13595-014-0447-4.

Twitter page Twitter

Learn Mycology

Welcome to the comprehensive guide on Mycology! As a budding mycologist or just someone curious about the diverse world of fungi, you're in for a treat. Fungi are everywhere - from the loaf of bread in your kitchen to the vast forests, playing crucial roles in the environment and our daily lives. This course aims to nurture your curiosity, guiding you through the intricacies of fungi and their manifold interactions with our world.

Course Outline

An introduction to mycology and fungi.

Start your mycological voyage by diving into the very essence of fungi. Explore their roles, unravel myths, and trace the steps of pioneer mycologists. Understand why fungi, often overlooked, are in fact indispensable to life as we know it.

1.1 What is Mycology: Definition and Overview
1.2 The Importance of Fungi
1.3 A Brief History of Mycology: From Discovery to Present Day
1.4 Basics of Mycological Study
1.5 Myths and Misconceptions about Fungi
1.6 Fungi and Biodiversity

The Basics of Fungi

Deconstruct the fungal organism. What makes a fungus, well, a fungus? Discover their structure, their genetic makeup, and the intricate classification system that mycologists use.

2.1 Defining Fungi
2.2 The Structure of Fungi
2.3 Fungal Classification

Fungal Biology

Delve deeper into the life of fungi. From their nutritional habits to their reproductive strategies and the intriguing genetics that drive them. Explore the vibrant and dynamic life of fungi from a biological perspective.

3.1 Fungal Life Cycle
3.2 Fungal Nutrition
3.3 Fungal Genetics

Fungal Ecology

Learn about fungi as global ecological players. Explore their environmental roles, their intricate relationships with other organisms, and their crucial place in biodiversity and ecological successions.

4.1 Fungi in the Environment
4.2 Fungi and Other Organisms
4.3 Fungal Succession and Biodiversity

Human Uses of Fungi

From penicillin to blue cheese, fungi have shaped human history in more ways than one. Dive into the diverse ways humans have harnessed the powers of fungi for food, medicine, and industrial applications.

5.1 Fungi in Food and Drink
5.2 Fungi in Medicine
5.3 Industrial Mycology

Fungal Diseases

While many fungi are beneficial, some can be pathogenic. Explore the world of fungal diseases that affect plants, animals, and even humans.

6.1 Fungal Pathogens of Plants
6.2 Fungal Pathogens of Humans and Animals

Fungal Conservation

In a changing world, fungal diversity is at risk. Delve into the challenges facing fungi and the concerted efforts in place to conserve these essential organisms.

7.1 Threats to Fungal Diversity
7.2 Fungal Conservation Efforts

Advanced Topics in Mycology

For the inquisitive mind, journey into the frontier of mycological research. Understand the evolutionary history of fungi, their behavior, and the cutting-edge research shaping the future of mycology.

8.1 Fungal Evolution
8.2 Fungal Behavior
8.3 Current Research and Future Directions in Mycology
8.4 Fungal Interactions and Chemical Ecology
8.5 Cutting-Edge Research and Future Directions in Mycology

Amazon Storefront

Our Amazon Storefront is a curated collection of products we recommend, hosted on Amazon. By purchasing through our storefront, you not only find quality mycology products but also support our website's growth through commissions we earn, enabling us to continue providing valuable content and recommendations.

Stay connected For any inquiries or assistance, feel free to reach out to us

Spotlight on: Mycology

The Biologist Vol 60(2) p36-37

Mycology is the study of fungi. It is closely associated with plant pathology as fungi cause the majority of plant disease.

Why is mycology important?

Fungi are the primary decomposers of organic material in many ecosystems and so play a crucial part in recycling nutrients and the global carbon cycle. They break down pollutants and the most durable organic materials and have a range of uses such as in medicine and food production. At least 80% of plants rely on mycorrhizal associations – symbiotic relationships between the plant's roots and a fungus that provides the plant with water and nutrients.

What careers are available?

Demand for fungal scientists is quite small but at the same time there is a severe shortage of mycologists, plant pathologists and taxonomists, as all these disciplines are taught less in universities. Still, mycologists can find work in many areas. The importance of fungi in crop growth, plant disease, fermentation and spoilage means there are jobs available in agriculture and the food industry. The unique properties of fungi offer many other industrial applications, such as the bioremediation of polluted land, while medicinal mycology researches potential pharmaceutical uses.

How do I start?

There are no undergraduate courses in mycology in the UK so most mycologists embark on postgraduate research after doing a more general bioscience or microbiology degree. Where mycology is taught as part of a bioscience degree, hospital-based universities tend to concentrate on pathogenic fungi, while others may focus on fungal ecology and plant pathology. Due to a lack of formal training opportunities, academics and employers look for an interest in fungi and a background in plant sciences or microbiology.

Where can I get more information?

The British Mycological Society is a charitable organisation for those working, studying or interested in mycology. There is a strong amateur contribution to the recording, discovery and conservation of fungal species in the UK, with many groups (including the Society's local branches) organising 'fungal forays' into woodland to find interesting or edible specimens. Kew Gardens' fungarium holds 1.25 million specimens and visitors can also see an excellent variety of fungi in its arboretum. UK fungus day is on Sunday 13th October, during Biology Week.

Web resources

www.britmycolsoc.org.uk www.societyofbiology.org/branches www.kew.org/plants-fungi/fungi www.fungitobewith.org

At a glance

Name: Professor Lynne Boddy FSB Profession: Professor of mycology at Cardiff University Qualifications: BSc in biology and mathematical statistics from Exeter; PhD in botany (wood decay) from Queen Mary, London; DSc in ecology of wood decomposition from Exeter Interests: Fungal communities in wood; ecology of cord-forming basidiomycete fungi; climate change effects on fungi; role of fungi in ecosystems; ecology of rare and endangered fungi

What led you to study fungus?

My first encounter with fungi was in student accommodation – I tried to open an old cupboard drawer and found it was stuck shut. I prised it open and there was all this mycelium attaching the drawers together. Then I tried to pull the whole thing out and found it was stuck fast to the wall.

It was the dry rot fungus Serpula lacrimans – its chords can penetrate brick and plaster. It was then I thought, "hey, these are cool".

I studied biology at Exeter University where the famous mycologist John Webster worked, so many think I became hooked because of him, which is probably true. For my PhD I started looking at the process of wood decay, but I instantly realised we needed to know more about the fungi involved.

Describe a typical day?

I spend my days teaching and doing research. I do lots of outreach work to get people to understand why fungi are important and why they are fun to study. Doing hands-on things in the lab is a rare treat – my post- docs do most of it – but when they go out on field work I always try to accompany them. About 85% of our fungal ecology experiments are done in the lab.

What are you working on now?

I study wood-decaying fungi and fungal communities – they're much like communities of plants except they're harder to study as they're hidden. My work is about what all these species are and what affects how they interact.

Fungal species are great fighters. They fight with each other all the time and I liken these interactions to the Premier League in football. You have your Manchester United fungi, who win most of the time, but sometimes one of the less successful ones beats them. Why is that? Certain environmental changes or the presence of invertebrates or bacteria alter these interactions.

I also work with cord-forming fungi. They don't just release spores and hope they land somewhere suitable, they grow mycelium out of the wood and create long foraging structures that search the forest for more dead wood to colonise. Like mycological motorways, these huge structures can shift nutrients around a forest in a matter of minutes, and actually behave a little like animals – when you compare their growth patterns to foraging patterns of ants or termites they're very similar.

What are the potential applications of your work?

It's blue sky research really – I just want to know how these things work but there are potentially lots of applications. Because they are such good fighters, fungi could be used as bio-control agents to prevent the spread of disease. Plus, they are endangered. I want to find out why certain species are so rare and how we can protect them.

Why are there so few mycologists?

There's a real shortage, caused by several problems. Firstly there are only a handful of taxonomists left in institutions, which, when you have 1.5 million species of fungi and only 100,000 have been described, is disastrous.

We have a couple of flourishing mycology research groups but the number elsewhere is dwindling. People retire and don't get replaced. There are more and more biology degrees where fungal biology is not compulsory and some where you don't get taught any at all, which is really worrying considering how important fungi are.

There are two groups of people in mycology: those who specialised in it after doing a biology degree, but also many skilled amateurs and enthusiasts. We are indebted to them and their discoveries, and taxonomy is often in their hands. Their contribution is hugely important.

COVID-19 Tracker
Biochemistry
Anatomy & Physiology
Microbiology
Neuroscience
Animal Kingdom
NGSS High School
Latest News
Editors’ Picks
Weekly Digest
Quotes about Biology

Reviewed by: BD Editors

Mycology Definition

Mycology is the study of fungi, their relationships to each other and other organisms, and the unique biochemistry which sets them apart from other groups. Fungi are eukaryotic organism which belong to their own kingdom. Until advances in DNA technology, it was assumed that fungi were an offshoot of the plant kingdom. DNA and biochemical analysis has revealed that fungi are a separate lineage of eukaryotes, distinguished by their unique cell wall made of chitin and glucans which often surrounds multinucleated cells. Mycology is a necessary branch of biology because fungi is considerably different from both plants and animals.

History of Mycology

Until the 1800’s, it was assumed that fungi were simply a different kind of plant. Mushrooms, the reproductive bodies of fungi, were eaten, used as medicine, and used for their hallucinogenic effects since antiquity. Many classic Greek philosophers and naturalists considered fungi, but still assumed they were more related to plants. By the mid-1800’s the microscope was invented, and scientists began to examine the inner workings of fungi. Microscopes revealed that fungi had distinct features, separate from both plants and animal cells. The term mycology was coined in 1836 in a paper by M.J. Berkeley, when fungi were beginning to be recognized as their own unique kingdom.

However, it was not until the advent of modern biochemistry and DNA analysis that it was fully realized how different fungi were. Instead of a cell wall made of cellulose, the wall in fungi is composed of glucans and chitin, molecules found in plants and insects, respectively. Instead of having a single nucleus, like most plants and animals, fungi are often multinucleated and contain special pores allowing the cytoplasm and nucleus to flow freely between various chambers in the fungal organism. DNA analysis revealed a closer relation to animals than plants. As scientists observed fungal lifecycles further, they realize that the majority of most fungi spends its time as a mold or ooze. This multicellular lifeform moves its way through decaying organic material, utilizing the minerals and organic molecules as it goes. Not only was fungi the major decomposing organism in the world, scientist also determined that certain fungi were responsible for events like fermentation and crop diseases.

With this, the field of mycology exploded. Agricultural mycology focuses on utilizing and controlling fungi in commercial crops. Toxicologists study mushroom and fungi for compounds which adversely affect other organisms. Pharmaceutical companies race to extract useful compounds from mushrooms. Careers in mycology are as diverse and complex as the field itself.

Careers in Mycology

Mycology first became an important science in the agricultural industry, and remains so today. A phytopathologist studies plant diseases, especially those which affect crops. Fungi are a major pest for many crops, but also serve symbiotic roles and allow plants to extract nutrients and water from the soil. Mycology is needed to distinguish between beneficial and harmful fungi, as well as to treat crops and prevent future infections. Further, certain types of fungi are used as pesticides, as they are more natural than synthetic pesticides and can kill targeted insects.

However, mycology has expanded well beyond its origins in agriculture. Once it was realized how broad and diverse the fungi kingdom is, the various roles of fungi in society were better understood. For instance, cheese is produced by various fungi. Mycology can classify and understand these organisms, leading to better and more efficiently produced cheese and dairy products. Yeast is also a form of fungi, and understanding the process of fermentation carried out by yeast is a science in itself. Fermentation science degrees can found from the bachelor level up, and graduates can work in the brewing and distilling industries, creating beer, wines and liquor. Yeast is also used in bread making, and microbiologists are required to maintain the cultures to produce enough yeast for bread production.

A specialized field of mycology is mycotoxicology , or the study of the toxins produced by mushrooms. Typically, a mycotoxicologist has a doctorate degree in biochemistry or organic chemistry, or a medical doctorate with concentrations in mycology and toxins. Fungi produce a variety of chemicals which have toxic effects on all kinds of organisms. Humans have eaten mushrooms since the earliest hunter-gatherers, but many mushrooms remain highly toxic. Other compounds found in mushrooms have potentially beneficial properties which could be used in medicine. Many mycotoxicologists work for pharmaceutical companies, trying to develop new drugs based on these compounds.

Mycology contains still more specializations, and is a continually evolving field. As more research is done, fungi are becoming a large and complex kingdom. Research is expanding and focusing on many special areas, including interesting applications for certain fungi. Some of these applications include radiotrophic fungi which appear to grow in the presence of radioactivity and could possibly alleviate radioactive wastes, and fungi which can break down complex organic substances into carbon dioxide. Many of these applications have tremendous commercial value, and researchers are needed at many institutions to explore these aspects of mycology.

Finally, an ethnomycologist is a scientist who studies the historical uses of fungi. Cultures have used mushroom as food, medicine, hallucinogens, and for a variety of other things. Ethnomycologists study these uses and inform the public and front-line researchers about which fungi have known effects and which are benign. Considering the immense size and diversity of fungi, and the relatively unorganized history of the classification of fungi, ethnomycologists provide a critical function in sorting through the dense but helpful information already gathered by past cultures and societies. The field of mycology is continually expanding as these many professions push the boundaries of knowledge and fill in the missing gaps.

Brusca, R. C., & Brusca, G. J. (2003). Invertebrates . Sunderland, MA: Sinauer Associates, Inc. McMahon, M. J., Kofranek, A. M., & Rubatzky, V. E. (2011). Plant Science: Growth, Development, and Utilization of Cultivated Plants (5th ed.). Boston: Prentince Hall.

Cite This Article

Subscribe to our newsletter, privacy policy, terms of service, scholarship, latest posts, white blood cell, t cell immunity, satellite cells, embryonic stem cells, popular topics, amino acids, hermaphrodite, horticulture, mitochondria, animal cell, endocrine system.

History & Society
Science & Tech
Biographies
Animals & Nature
Geography & Travel
Arts & Culture
Games & Quizzes
On This Day
One Good Fact
New Articles
Lifestyles & Social Issues
Philosophy & Religion
Politics, Law & Government
World History
Health & Medicine
Browse Biographies
Birds, Reptiles & Other Vertebrates
Bugs, Mollusks & Other Invertebrates
Environment
Fossils & Geologic Time
Entertainment & Pop Culture
Sports & Recreation
Visual Arts
Demystified
Image Galleries
Infographics
Top Questions
Britannica Kids
Saving Earth
Space Next 50
Student Center

When did science begin?
Where was science invented?
Why is biology important?
How do fungi obtain nutrition?
What is a fungal spore?

Blackboard inscribed with scientific formulas and calculations in physics and mathematics

Our editors will review what you’ve submitted and determine whether to revise the article.

Mycology Online - Mycology

mycology , the study of fungi , a group that includes the mushrooms and yeasts . Many fungi are useful in medicine and industry . Mycological research has led to the development of such antibiotic drugs as penicillin , streptomycin , and tetracycline , as well as other drugs, including statins (cholesterol-lowering drugs). Mycology also has important applications in the dairy , wine , and baking industries and in the production of dyes and inks. Medical mycology is the study of fungus organisms that cause disease in humans .

An official website of the United States government

The .gov means it’s official. Federal government websites often end in .gov or .mil. Before sharing sensitive information, make sure you’re on a federal government site.

The site is secure. The https:// ensures that you are connecting to the official website and that any information you provide is encrypted and transmitted securely.

Publications
Account settings
My Bibliography
Collections
Citation manager

Save citation to file

Email citation, add to collections.

Create a new collection
Add to an existing collection

Add to My Bibliography

Your saved search, create a file for external citation management software, your rss feed.

Search in PubMed
Search in NLM Catalog
Add to Search

The importance of fungi and of mycology for a global development of the bioeconomy

Affiliation.

1 Institute of Biotechnology and Chemistry, Faculty of Science and Technology, Aalborg University, Denmark.
PMID: 23155503
PMCID: PMC3399105
DOI: 10.5598/imafungus.2012.03.01.09

The vision of the European common research programme for 2014-2020, called Horizon 2020, is to create a smarter, more sustainable and more inclusive society. However, this is a global endeavor, which is important for mycologists all over the world because it includes a special role for fungi and fungal products. After ten years of research on industrial scale conversion of biowaste, the conclusion is that the most efficient and gentle way of converting recalcitrant lignocellulosic materials into high value products for industrial purposes, is through the use of fungal enzymes. Moreover, fungi and fungal products are also instrumental in producing fermented foods, to give storage stability and improved health. Climate change will lead to increasingly severe stress on agricultural production and productivity, and here the solution may very well be that fungi will be brought into use as a new generation of agricultural inoculants to provide more robust, more nutrient efficient, and more drought tolerant crop plants. However, much more knowledge is required in order to be able to fully exploit the potentials of fungi, to deliver what is needed and to address the major global challenges through new biological processes, products, and solutions. This knowledge can be obtained by studying the fungal proteome and metabolome; the biology of fungal RNA and epigenetics; protein expression, homologous as well as heterologous; fungal host/substrate relations; physiology, especially of extremophiles; and, not the least, the extent of global fungal biodiversity. We also need much more knowledge and understanding of how fungi degrade biomass in nature.The projects in our group in Aalborg University are examples of the basic and applied research going on to increase the understanding of the biology of the fungal secretome and to discover new enzymes and new molecular/bioinformatics tools.However, we need to put Mycology higher up on global agendas, e.g. by positioning Mycology as a candidate for an OECD Excellency Program. This could pave the way for increased funding of international collaboration, increased global visibility, and higher priority among decision makers all over the world.

Keywords: biodiversity; biomass conversion; fungal enzymes; global challenges; new biological solutions; secretomics; teaching; training.

PubMed Disclaimer

The Biomass Value Pyramid shows…

The Biomass Value Pyramid shows the entire cascade of value adding products which…

The Peptide Pattern Recognition, (PPR),…

The Peptide Pattern Recognition, (PPR), generated GH13 protein subfamilies which predicted the enzyme…

When grown in swine wastewater,…

When grown in swine wastewater, some duckweed species such as the Spirodela polyrhizza…

Leaf-cutter ant colony established in…

Leaf-cutter ant colony established in the laboratory of JJ Boomsma (University of Copenhagen).…

Related information

Linkout - more resources, full text sources.

Europe PubMed Central
PubMed Central
Citation Manager

NCBI Literature Resources

MeSH PMC Bookshelf Disclaimer

The PubMed wordmark and PubMed logo are registered trademarks of the U.S. Department of Health and Human Services (HHS). Unauthorized use of these marks is strictly prohibited.

Mycology 101

Introduction to Mycology

Introduction to Mycology: Exploring the Fascinating World of Fungi invites learners to embark on a captivating journey into the mysterious and diverse realm of fungi. This in-depth course, designed for enthusiasts and aspiring mycologists alike, offers a comprehensive understanding of the myriad roles that fungi play in our world, from their integral contributions to ecosystems to their profound impacts on human society.

Throughout the 17 robust ( work-at-your-own-pace ) lessons , we dive into topics like the fundamentals of mycology, the importance of fungi in nature, fungal anatomy, life cycles, and classifications, and delve into the unique characteristics of major fungal groups. Discover the intricate world of fungal biodiversity and explore how fungi support ecosystem health, form symbiotic relationships, and facilitate nutrient cycling.

Then, we traverse from the natural to the human realm, discussing practical applications of fungi, from cultivating edible and medicinal mushrooms to industrial uses and ethnomycology. We cap it off with deep dives into medical mycology, fungal toxins, and mycotoxins, and the exciting prospects of fungi as sources of novel pharmaceuticals.

At the 🍄 Mushroom Network , we’re committed to continually nurturing this “ Living Course .” Our promise to you is that we will regularly update and refine the course to incorporate the latest in fungal research and traditional ecological knowledge. What’s more, enrollment in this course is entirely free! This means you can revisit the lessons anytime you wish to refresh your knowledge or explore the exciting new updates.

The best part? Once you’re enrolled in this course, you’ll have lifetime access to these updates. Whether it’s a groundbreaking study, a new method of cultivation, or a novel pharmaceutical use discovered, you’ll be able to benefit from these enhancements as they become available. And did we mention that it’s free? We firmly believe in the open dissemination of knowledge, and we’re excited to have you join us on this ongoing educational adventure.

Note: This Living-Course is designed for Patrons who are curious about mycology and have a basic understanding of biology. No prior knowledge of mycology is required.

The mushroom network, requirements.

Access to a computer or mobile device with a stable internet connection.
Basic understanding of biology and scientific concepts (recommended but not mandatory).
Enthusiasm and curiosity for learning about the fascinating world of fungi.
Comprehensive Curriculum: The course covers a wide range of topics, including fungal biology, taxonomy, ecology, cultivation techniques, medicinal applications, and more. It provides a holistic understanding of mycology.
Engaging Learning Materials: The course offers interactive videos, engaging reading materials, hands-on assignments, and quizzes to ensure an immersive learning experience.
Experienced Instructors: The course is taught by highly knowledgeable and experienced instructors who are passionate about mycology and dedicated to helping Patrons succeed in their learning journey.
Collaborative Learning Community: Patrons have the opportunity to connect with fellow mycology enthusiasts, share insights, and collaborate on projects, fostering a supportive and engaging learning environment.
Practical Applications: The course emphasizes practical applications of mycology, equipping Patrons with skills and knowledge that can be applied to various fields such as agriculture, medicine, environmental conservation, and more.

Target audiences

Mycologists
101 Introductory Courses

View Mycology Courses

[sg_popup id=347]Report a Bug[/sg_popup]

The Mushroom Network is proudly affiliated with: SPORE ZE WORLD | Green Grass Grove | 3G-Camping Retreat | 3G-Prints | Project Aurora .

Modal title.

Open access
Published: 12 December 2014

The importance of fungi and mycology for addressing major global challenges

Lene Lange 1

IMA Fungus volume 5 , pages 463–471 ( 2014 ) Cite this article

14k Accesses

44 Citations

21 Altmetric

Metrics details

In the new bioeconomy, fungi play a very important role in addressing major global challenges, being instrumental for improved resource efficiency, making renewable substitutes for products from fossil resources. upgrading waste streams to valuable food and feed ingredients, counteracting life-style diseases and antibiotic resistance through strengthening the gut biota, making crop plants more robust to survive climate change conditions, and functioning as host organisms for production of new biological drugs.

This range of new uses of fungi all stand on the shoulders of the efforts of mycologists over generations: the scientific discipline mycology has built comprehensive understanding within fungal biodiversity, classification. evolution, genetics, physiology, ecology, pathogenesis, and nutrition. Applied mycology could not make progress without this platform. To unfold the full potentials of what fungi can do for both environment and man we need to strengthen the field of mycology on a global scale.

The current mission statement gives an overview of where we are, what needs to be done, what obstacles to overcome, and which potentials are within reach. It further provides a vision for how mycology can be strengthened

The time is right to make the world aware of the immense importance of fungi and mycology for sustainable global development, where land, water and biological materials are used in a more efficient and more sustainable manner. This is an opportunity for profiling mycology by narrating the role played by fungi in the bioeconomy. Greater awareness and appreciation of the role of fungi can be used to build support for mycology around the world. Support will attract more talent to our field of study, empower mycologists around the world to generate more funds for necessary basic research, and strengthen the global mycology network. The use of fungi for unlocking the full potentials of the bioeconomy relies on such progress. The fungal kingdom can be an inspiration for even more.

Introduction to the Importance of Mycology

Applied mycology has traditionally focused on areas where fungi cause damage. Highly advanced mycological research has generated knowledge, conceptual understanding and insight within fungal plant diseases, human mycoses, indoor climates, and decays in wood, feed and food. There is also extensive knowledge about the added value fungi contribute to beer, bread, wine, spirit, food preservation, and food taste. Much less attention has been given to fungi and fungal products for adding value for industry, agriculture, health, and pharmaceuticals. The last few decades, however, have witnessed very interesting developments regarding use of fungi for new processes, products and solutions of importance for the world, and most importantly for increased conceptual understanding of the kingdom Fungi at large (Hibbett et al. 2007 , James et al. 2006 , Schoch et al. 2006 ).

It is now well documented, even in scaled up practice, that fungi can generate tangible and substantial value through improved resource efficiency, resulting in decreased pollution and greenhouse gas emissions. Such progress stands on the shoulders of the contributions made by mycologists over decades and even centuries to build up mycology as an independent field of science. Importantly, value added use of fungi was founded not only on applied studies but also on basic research aimed at understanding fungal biodiversity, growth, nutrition, physiology, genetics, metabolism, and ecology.

The shift from chemical processes to biological processing, achieved by using fungal (and bacterial) enzymes instead of chemical processes in industries, such as textiles, leather, paper and pulp, has significantly reduced negative impacts on the environment. Use of enzymes in the food and feed industry, such as animal feed, baking, brewing, and wine and juice, has significantly improved what we get out of biological raw materials. Microbial enzymes added to detergents, washing laundry clean even at low temperatures, has significantly reduced CO 2 emissions. The newest chapter in the industrial biotechnology era is to substitute fossil resources with renewable resources (Fig. 1 ).

The value pyramid of biomass conversion: At the bottom, with lowest value, is the bulk use of biomass for combustion, making heat and electricity. Next layer is using biomass for biofuel, a much needed renewable alternative to fossil transport fuel. Further up the biomass value pyramid, is production of specialized and higher value products (materials, chemicals, feed, food and pharma). Such products are not only substituting for fossils but also making use of the complex structures of new biomass, making products not possible to make from fossilized biomass. All uses except for burning, gasification, etc., of the biomass involve use of microbial (primarily fungal) conversion and upgrade. Courtesy of Peter Westermann.

The Role of Fungi and Mycology in Addressing Major Global Challenges

Fungi play an important role in addressing major global challenges. Use of fungal processes and products can lead to increased sustainability through more efficient use of natural resources. Applications range from upgrading bio-waste for value added products to use of renewable plant biomass as a substitute for oil-based products such as biochemicals, plastics, fertilizer, and fuel. Fungal inoculum, introduced into soil together with seed, can promote more robust plant growth through increasing plant uptake of nutrients and water, a robustness of importance for maintaining crop yields under climate change condition. Fungal enzymes can lead to production of food ingredients with prebiotic effects for a healthier human gut biota and hence greater resilience towards life-style diseases. Similarly, use of fungi can be a short cut to healthier animal feed and less use of antibiotics in, for example, meat production, one of the current prime sources of multiple drug resistant bacteria. Fungi are one of nature’s most promising hotspots for finding new drug candidates and antimicrobials. Last but not least, fungi have interesting potential as the new way of manufacturing biological medicines and a wide spectrum of new value added bio-based products.

All such uses of fungi, fungal products and fungal processes reflect the efforts of mycologists over generations. Similar efforts lie behind work to cure and prevent life threatening human mycoses, to control mycotoxin contaminations, and to counteract fungal spoilage of materials and ‘sick building’ syndrome. As a scientific discipline mycology has built comprehensive understanding of the fungal kingdom: fungal biodiversity, physiology, genetics, ecology, pathogenesis, nutrition. Mycology includes understanding at the system, organismal and molecular level. This knowledge and insight constitute the platform that has given rise to uses of fungi in industry, agriculture, food and feed, medicine and health.

Mycology must grow fast beyond where it is today. The potential of fungi for a more sustainable world must be released to address global challenges of climate change, higher demands on natural resources, and the increased burden of lifestyle diseases. Genome sequencing was developed first for the human genome after which bacterial genomes were quickly sequenced. But mycology is catching up. Interestingly, up to now, industry makes use only of a minute portion of the fungal kingdom. Fungal biodiversity is a resource pool for the future. However, fungal diversity is endangered by loss of habitat, causing loss of species and loss of biodiversity in general.

We need to stimulate mycology globally and work more efficiently together to take good care of this diversity and unlock the full potential of the fungal kingdom for future use all over the world. The discipline of mycology needs to be developed to a stage where it can recruit talent for the new generation of mycological researchers and for building the skills needed for the world to change towards the new and more sustainable bioeconomy.

In summary what is needed is: Increased understanding of the fungal kingdom , phylogeny and phylogenomics as a basis for understanding the fungal life-form generally, and for expanding the exploitation of fungal biodiversity for more value added uses; the mycological platform , building mycological know how and skills in all parts of the world. There is a global need for the bioeconomy, for increased resource efficiency and upgrading of biowaste to healthier food and feed ingredients, materials and fuel; a stronger global mycology network , including globally distributed databases (embracing genotypic and phenotypic data), improved opportunities for networking activities, talent recruitment, research education, and for broadening the IMA activities to real global inclusiveness and perspective; Open access mycology , a new concept where cultures and information go together to support knowledge dissemination, enabling distributed uses of fungi for upgrading bio-waste resources to higher value; a concerted effort to build a stronger focus on the role of fungi in nature , how they interact with substrate and other organisms, including global research efforts on the fungal secretome, as a basis for increased collaboration between academia and industry within the field of mycology; revisiting the traditional use of microbial fungi for food , the advanced mycological heritage, revitalized through molecular studies, giving insight in use of microbial consortia for food processing to inspire also the next generation of biobased products; and not the least: excellence in teaching of mycology , to be developed in a global scale to provide a better platform for recruitment of talent, for stimulating a fascination of fungal life, and for building broader skills for the bioeconomy to unfold.

The focus was initially on enzymatic conversion of wheat straw and corn stover to bioenergy. The current trend is to expand the types of biomass exploited, going beyond the use of crop residues alone and including agroindustrial waste (side streams and by-products) to produce higher value products such as bio-chemicals, biomaterials, food and feed ingredients, and circulating the micronutrients back to the soil. In other words, not only converting biomass to lower value bioenergy bulk products. This entire global endeavor builds almost exclusively on fungal enzymes produced on a large scale by fermentation of filamentous fungi, with recombinantly expressed fungal genes. Production hosts of choice were primarily Aspergillus oryzae, A. nidulans and Trichoderma species. Surprisingly the enzyme genes used are still almost all from a rather narrow selection of ascomycete and basidiomycete genera.

In order to take these next steps, we must develop mycological research even more. We need to tell this story and make the world aware of the immense importance of fungi and mycology for sustainable global development, where land, water and biological materials are used in a more efficient and more sustainable manner. This offers us opportunity also for profiling mycology by narrating the role played by fungi and fungal enzymes in the bioeconomy. Greater awareness and appreciation of the role of fungi can be used to build support for mycology around the world. Support will attract more talent to our field of study, empower mycologists around the world to generate more funds for necessary basic research, and strengthen the global mycology network.

The Importance of Mycological Research for Further Biological Solutions to Important Problems

Discovery of novel fungal products for specific industrial needs, builds on basic studies of fungal biodiversity, and on using the experimental molecular mycology tool box for identifying potentially interesting genes and proteins, synthesis pathways, and metabolites. For industrial use it is the secretome of the fungi which is targeted for both metabolite and enzyme discovery Footnote 1 .

Most molecular studies within fungi have focused on what the fungi are and only less on what fungi do and how they interact with other organisms and substrates. The molecular era first focused on elucidating phylogenetic relationships and species identification. Now the time is ripe for intensifying the study of the secretome with regard to composition, development, and evolution. Molecular studies of the secretome (genomics and transcriptomics of secreted proteins) will add new conceptual understanding of the role of the secretome in substrate specialization and organismal speciation. We will learn more about the mechanisms of secretome evolution, going beyond mutation and selection. This evolution was most likely influenced by mechanisms such as series of gene copy/gene loss and retention of the protein with best fit for substrate utilization and growth; or by horizontal transfer, the latter being expected to be found especially in closely interacting ecological niches such as the animal rumen and the phyllosphere.

Discovery of enzymes suitable for industrial processes draws upon mycological research within a wide range of mycological disciplines. These include biodiversity, fungal physiology and ecology, molecular biology, protein chemistry, enzymology and assay technology, and not least experimental mycology, involving yeasts (as screening hosts) and filamentous fungi (as gene donors and protein production organisms). Progress in all such fields has made it possible within a rather few years to find the enzyme cocktail needed to break down recalcitrant lignocellulose to a monomer sugar platform (Carvalheiro et al. 2008 , Kubicek et al. 2014 , Martinez et al. 2009 ).

Until recently, the short cut from enzyme discovery to testing most promising enzyme proteins employed the following steps: optimizing the enzyme discovery by identifying and choosing the conditions (substrate, temperature, aeration) most conducive to enzyme expression, harvesting the induced fungal biomass, extracting total RNA, recovering the mRNA fraction, converting to cDNA, constructing cDNA libraries, and screening the libraries on specific assay plates, allowing for identification of positive transformant yeast colonies, recombinantly expressing a full length gene, resulting in a secreted enzyme, with the activity indicated by the assay plate used (Dalb0ge & Lange 1998, Fowler & Berka 1991 ).

In the last few years, choice of technology for enzyme discovery has moved towards a strategy focusing on genome or transcriptome sequencing instead of (cDNA) library construction and screening (Huang et al. 2014 ). This new sequencing approach allows for higher speed discovery and lower cost experimental work. However, it also leads to the need for additional effort within protein expression. For a particular genome sequence there is much less information about whether it can easily be expressed recombinantly and whether the result of the expression will be a functional enzyme. With the cDNA screening strategy, you have a yeast clone where you know you have a full length gene, that the gene is expressed; and that expression leads to a secreted, functional gene, active in a specific enzyme assay.

Similarly, scaled up production of secreted fungal enzymes rests on experimental results, conceptual understanding and insight achieved through basic mycological research within the fields of genetics, molecular biology and protein expression - embracing also optional expression of basidiomycete, zygomycete and chytridiomycete proteins in ascomycete hosts and production organisms. For improved biological production we need a strengthened basis in fungal phylogeny, progeny trees, protein expression, fermentation technology, and not least more knowledge about the biosynthetic pathways of mycotoxins and mycotoxin occurrence and toxicity (Frisvad & Samson 2004 ). Besides such mycological research, large scale production is also based on knowledge within the field of chemistry and on fungal growth, including modelling studies for optimization of growth conditions.

What is often forgotten when trying to see the big picture is that the success of industrial biotechnology using fungi and fungal products builds primarily on the unique and highly efficient function of the tip cells of filamentous fungi, efficiently secreting proteins produced by the rapidly growing mycelial biomass. Advanced bio-imaging studies have provided valuable insight into this exciting field of biology; more bio-imaging studies across fungal phyla could give insight in the variation found within the fungal kingdom (Harris et al. 2005 ).

Inspiration and Learning from Nature

The need for more types of enzyme blends for efficient decomposition of further kinds of lignocellulosic biomass (not only wheat straw and corn stover) has revitalized the entire field of studies on fungal degradation of lignocellulose, including ascomycetous saprophytes and basidiomycetous white rot and brown rot fungi. This quest has also led to new studies of ecological habitats, where biomass conversion takes place in nature, developed over evolutionary time. The enzymes from the termite larvae gut channel and from the cow rumen, investigated by activity screening of cDNA meta-libraries, were already studied decades ago by some biotechnology companies. Description of the biomass conversion of the ectomycorrhizal Paxillus involutus (Rineau et al. 2012 ), studied by mass spectrometry (MS) for the documentation of bioconversion and Transposon Assisted Signal Trapping (TAST) of the cDNA library for discovery (Hamann & Lange 2006 , Rineau et al. 2012 ), gave new insights into the possible dual function of this ectomycorrhizal fungus.

A most interesting meta-study examined the enzymes produced and secreted in the fungus garden of the leaf cutter ant (Grell et al. 2013 ). This showed that Leucoagaricus gongylophorus , farmed by leaf cutter ants, expressed the entire spectrum of enzymes needed for breaking down the cellulose fibres of the green leafy biomass which the ants bring to the fungal garden in their nest (Fig. 2 ). The ants pretreat by chewing the biomass. The fungus expresses the needed regime of enzyme proteins. The enzymes even survive (in an intact and active form) passage through the gut channel of the ants. Redistribution of the enzymes to the newly harvested leafy biomass is achieved by the ants placing their enzymeholding fecalia on newly harvested leaves on the top of the fungus garden. This entire sophisticated mutualistic system has been comprehensively described (Kooij 2013 ).

Learning from Nature’s “green biorefinery”: Leaf cutter ants carry pieces of green leaves to the fungus garden ( upper ), feeding the fungal symbiont, Leucoagaricus gongylophorus , farmed in the fungal garden in the subterranean ant nest ( lower ; laboratory culture of Jacobus Boomsma). The fungus produces swollen tipped cells, filled with proteins and sugars, the gongylidia, organized in staphylae. The gongylidia are picked by the ants for feeding the ant colony with protein and sugar rich feed. Bottom line is that this successful and complex society, where fungal enzymes convert green leaves into accessible, highly nutritious fungal biomass provide basis for one of the most successful life forms on earth. Photo: Henrik H. De Fine Licht.

In a recent review of the prominent role of fungal enzymes for the success of the leaf cutter ants (Lange & Grell 2014 ), the system is described from a new perspective that sees the fungal partner in the symbiosis not merely as being passively farmed but also as an active partner. In this interpretation the Leucoagaricus fungus is an example of a fungal adaptation path towards developing attractants to recruit insects to disperse the fungal spores (as, for example, flies are attracted to the spore gleba of Phallus impudicus and thus lured into disseminating the fungal spores). The ant-farmed L. gongylophorus benefits from the association by having an assured food supply, by having its colony weeded for intruders, and by being protected from parasite populations (Seifert et al. 1995 ). Lange & Grell ( 2014 ) reviewed the literature and concluded that the mutualism functions on the molecular level due to the fungal enzymes and on the organismal level due to the fungal gongylidia (swollen hyphae filled with proteins and sugars), serving as highly nutritious food for the ants. The ants collect, distribute and chew (pretreat) the leafy biomass, and remove “garbage”. It is the continuous supply of highly nutritious fungal gongylidia organized in bunches, the staphylae, readily harvestable by ants, which provides the basis for the ant colonies to grow to such size and societal complexity. The global success of the ants is based on the fungal enzymes, the nutritional value of the fungal mycelium and the efficiency of the filamentous fungus growth and tip cells.

We can learn from nature about how to construct biorefinery processes. The leaf cutter ant colony including the fungal garden can be seen as the archetype of a green biorefinery. The cow rumen can be seen as a biorefinery for decomposing lignocellulosic straw, the yellow biorefinery. And the termite larvae gut channel can be seen as a biorefinery decomposing lignocellulosic woody materials. Interestingly, we also see that fungi are responsible for most of the lignocellulytic enzymes in most such biomass decomposing habitats in nature. However, bacteria are always found in such biomass conversion niches and most likely are also playing a role in overall biomass conversion.

The New Potential of Fungi and Fungal Products from Biorefineries

There are many new examples in the pipeline of both academic and industrial research for upgrading the value of biomass and waste. One of the most interesting is directly inspired by the leaf cutter ant/fungal garden mutualism. In the fungal garden of the ants, the fungus itself serves as highly nutritious animal feed! Feeding the projected nine billion people on Earth by 2050 (United Nations’ World Population Prospects report) will put enormous pressure globally on available arable land, water, and nutrition. Right now at the global scale we use around 72% of all arable land for the production of animal feed. If more animal feed is produced based on the huge amounts of bio-waste we lose and discard along the chain from crop to food, and from the field to end user, we could release more land for biodiversity conservation and for food production (FAO 2013 ).

The latest research paves the way for making even more efficient use of the potential of biomass. First, cellulose fibres are broken down to sugar monomers by fungal enzymes. Next in the value upgrade is using the sugar platform for growing microbes which produce building blocks for chemicals and for biopolymers, such as bioplastics. The lignin will be developed into binders and materials (still to be developed). And the hemicellulose polymer is processed by fungal enzymes for recovery of C 5 sugar oligosaccharides with prebiotic activity for a more healthy gut microbiota. This means that now we do not just break down nature’s complexity for building up new complexity from the sugar platform. We also recover nature’s complexity, for example proteins and hemicelluloses, to make new and upgraded products, food, feed, and materials with new properties and value adding functionalities.

Fig. 3 shows some of the specific and highly specialized functions of fungal enzymes, currently under evaluation for industrial processing of hemicellulose into high value prebiotic food and feed ingredients. The ambition is to develop food ingredients which can make people more robust against life-style diseases and to develop animal feed for non-ruminant animals such as pigs with the prebiotic effect to improve metabolism. This gives better welfare and less need for antibiotic treatment — and limiting antibiotic use lowers the risk of runaway resistance to antibiotics. All this is achievable by converting waste materials using fungal enzymes.

The hemicellulose plant cell wall polymer, arabinoxylan, is degraded by many different and highly specialized fungal enzymes in nature: 1. endoxylanases; 2. α-L-arabinofuranosidases; 3. glucuronidases; 4. ferulic acid esterases; and 5. acetyl xylan esterases. Ongoing research aims to use such specific fungal enzymes to modify the arabinoxylan, a sub stream from lignocellulose biorefinery, into C 5 sugar oligosaccharides with a prebiotic effect, stimulating the healthy gut fungal and microbial populations of humans and other animals. Modified from Chavez et al. (2006).

The biotech industry has a good track record of using not just plant biomass but also fungal biomass directly as feed. The yeast biomass or “vinasse” left over from large scale production of insulin was used as animal feed for pig production in Denmark for many decades. The pigs loved it! Now a new approach is proposed. Transform sanitized household bio-waste into a fermentation medium, and use it as substrate for fungal growth. From such a system the fungus biomass in itself, as a yeast cream, can be used as animal feed, which through fungal and bacterial biotransformation is one step removed from waste. With the available nitrogen supply the fungus will be able to develop into new nutritious protein-rich animal feed. Focused efforts will have to be made to optimize such a system, including optimizing the waste to fermentation medium process, for example by mixing different complementary and matching waste streams. Research efforts have also been initiated with the objective of strain improvement to achieve an even higher nutritional value of the resulting fungal biomass. Two groups of fungi are being studied for this purpose: yeasts ( Saccharomyces cerevisiae or Candida utilis nonallergenic mutants) and basidiomycetes such as those grown by termites ( Termitomyces , Fig. 4 ) (Aanen et al , 2009 ; Nobre et al. 2010 ), ants ( Leucoagaricus ), or humans ( Pleurotus ). Additional molecular studies can lead to even higher levels of fungal protein content and bio-accessibility. The strain improvement of such “Waste2Value Fungi” can also be expanded to include strains which concentrate micronutrients, such as phosphorus, iron and selenium, further enhancing the nutritional value of the fungal animal feed.

Learning from nature: the specialized basidiomycete Termitomyces titanicus ( Agaricales ) grows in subterranean termite nests. It can grow to form massive and impressive basidiomes, used as a human delicacy (above). The benefit to the termites — even without having developed the sophisticated farming procedure — is accessibility to protein rich feed. The percentages of protein in edible basidiomycetous fungi are high (measured as% of total dry weight; (below); an extraordinarily high protein content has been recorded for Termitomyces species. In future we will be able to make biorefineries by growing fungi on household waste and use the protein rich fungal biomass for animal feed. Photo and table: Duur Aanen.

Another potential way to extend conversion of biowaste for producing food and feed ingredients is to include animal derived materials, such as fish (by-catch and waste) as well as slaughter house waste such as pig bristles and chicken feathers (Fig. 5 ). New discovery efforts are underway to find and develop fungal (and bacterial) enzymes, primarily proteases, for converting such hitherto unexploited protein rich waste streams, for example obtaining otherwise inaccessible protein from the keratin of pig bristles and chicken feathers decomposed by enzymes from Onygena corvina (Huang, Busk & Lange, unpubl.) to produce nutritious food and feed ingredients, without using more land or making further inroads into naturally occurring or farmed fish populations. Another area where we need new enzymes is for the conversion of leftover press cake from production of plant oil from olive, palm tree, sunflower, rape, etc., into bio-accessible protein-rich animal feed. This represents a substantial underexploited source of protein for both animal and human consumption.

Onygena species. Non-pathogenic species of Onygenales , are specialized in breaking down the keratin found in feather, hooves, and horn. The keratin is composed of proteins, bound in a non-bio-accessible form. Among the large number of different proteases produced by O. corvina , we discovered that just three enzymes, belonging to two types of protease families, are sufficient to breakdown both feather and pig bristles. The picture shows O.equina growing on horn, but not on the skull (Northern Ireland, 2012). Photo: Jens H. Petersen.

New Meta-Level Discoveries

The affordable price of genome sequencing has made meta-studies of entire ecological niches doable, for example by metatranscriptomic or metagenomic sequencing, where an entire niche is handled as if it were one organism. This approach enables inclusion of non-culturable organisms as well as all kinds of auxiliary enzymes we still are ignorant of. Another approach under development is to start including more than just ascomycetes and basidiomycetes in the enzyme discovery efforts. Interesting results are being gained right now from chytrids and zygomycetes, notably Entomophthorales (Grell et al. 2011 ) and Mucorales (Huang et al. 2014 ).

New Sequence Analysis Tools, Making Order and Sense Out of Chaos

As outlined above, the need for new enzymes is huge. Building the bioeconomy will include significant efforts in enzyme discovery to facilitate the diversification of substrates to be upgraded and of products to be developed from the biorefinery. Such discovery efforts will simultaneously result in massive accumulation of underutilized genome sequencing data (Murphy et al. 2011 , Grigoriev et al. 2011 ). The vision is straightforward: if we become better at predicting functions from sequences, we could significantly shorten and sharpen the enzyme discovery process. We could go directly from sequence to the subgroup of enzyme genes we would like to screen to identify the one with the highest potential for that specific biomass conversion process. With that specific goal in mind, Busk & Lange ( 2013 ) developed a new sequence analysis approach, Peptide Pattern Recognition (PPR), giving a function-based protein classification by recognition of short, conserved peptide motifs.

PPR, the first non-alignment based sequence analysis methodology, predicts function directly from sequence with an accuracy of 80–97% (Busk & Lange 2013 ). The method was developed further to include a regime by which a large sequence database generated through PPR can be mined for the presence of, for example, glycohydrolases, resulting in a list of genes referred to specific CAZy protein families ( http://www.cazy.org/ ; Cantarel et al. 2009 ). This list can be transformed into one providing an overview of all the functions (illustrated as a list of EC numbers) found in the secretome of that ecological niche. For each function, the protein families that have those functions, often more than one type of protein family for each EC function, can be speciied.

Fungal Inoculum for Stronger and More Robust Plant Growth

Another highly promising field for new microbial and fungal products is the use of inocula for strengthening crop plants, making the plants more robust to abiotic stress and more efficient with regard to water and nutrient utilization. Since the days of the green revolution, plant breeding has taken place almost in isolation (except for breeding for increased tolerance against certain plant diseases and pests). Now, advances in the field of fungal and microbial products for agriculture is making it possible for plant breeders, mycologists, and microbiologists, to work together to find the combination of plants and other organisms which will provide farmers and the world with more robust and resilient agriculture. At a time when climate change challenges agriculture in many parts of the world, and where water and nutrition are at a minimum in many places, the combinations of fungi, microbes, and plants can provide opportunities for significant progress in global food, feed, and biomass production.

It will be interesting to follow developments in this area. Again, a new field is emerging that builds on a platform of knowledge generated through the combined efforts of public and private mycological research, where fungi are seen as having the potential to contribute significantly towards a more sustainable world. This is therefore a highly interesting area for contributions from mycologists specialized in endophytes, mycorrhizas, Penicillium, Aspergillus, Trichoderma, Fusarium , soil fungi, and consortia where bacteria and fungi work together to produce efficient and optimized systems.

One product is already available on the market which increases phosphate availability, “JumpStart” (Novozymes 2012; http://www.bioag.novozymes.com/ ), which is a phosphate inoculant containing the naturally occurring non-GMO soil fungus Penicillium bilaii. The fungus colonizes and grows along plant roots, releasing organic compounds that, in turn, release the “bound” mineral forms of otherwise less available soil and fertilizer phosphate, which thus becomes immediately available for the crop plant.

New Breakthroughs Needed

Next in demand are regimes of enzymes active under low temperature conditions and enzymes active and stable at high temperatures for decomposition of both plant and animal derived biomasses. Here, mycologists specialized in extremophiles can contribute significantly (Zajc et al. 2013 , Pitt & Hocking 2009 , Gleason et al. 2010 ). Expansion of commercialized enzymes from highly extreme ecological niches, such as extremophilic archaea, with enzymes active and stable above 100°C, also calls for new breakthroughs in recombinant expression. Could intensified studies of the archaean splicing mechanisms and protein expression lead to a breakthrough in the development of fungal production hosts for heat stable secreted enzymes from these microorganisms? Improved expression of basidiomycete genes in ascomycetous expression hosts, such as yeasts, also builds on an improved understanding of the variations in protein expression mechanisms within the fungal kingdom. Enzymes to convert the proteins locked into keratin in feathers and bristles into bioavailable protein resources would pave the way for using such animal waste products for highly nutritious animal feed (Huang, Busk & Lange, unpubl.).

There are discoveries to be made of new kinds of antibiotics from fungal hotspots for antibiotics. For example insect pathogenic fungi, which have developed ways to reserve the entire insect carcass for themselves, or cellulose/sugar fungi that protect the dimer and monomer sugars against piggybacking bacteria, preventing them from ‘stealing’ the sugars after the fungi have produced all the enzymes to break down the lignocellulose. Discovery of novel drug candidates with new modes of action and central nervous system active metabolites from fungi, which manipulate insects to adopt behavioral patterns that optimize the chances for dispersal of fungus spores formed on the insect after the insect dies (e.g. ants and flies programmed to climb to the top of grass and cling there until death; or the house-fly which is manipulated to settle on a window glass pain and spread out its wings to maximize the area for fungal spore production and release). Maybe such research efforts can reveal new concepts for neurosignalling, induced in animals, based on stimuli given by fungal metabolites or proteins, but which may also be of relevance for increased understanding of the human central nervous system?

New higher value products can also be developed from fungal “vinasse” biomass left over from large scale biotech production. This is important because there will be a steep increase in amounts of fungal biomass available for upgrade as the technology of biomass conversion in biorefineries develops. In nature, complex patterns and mechanisms of collaboration are expected to occur especially between fungi and bacteria. This is a most interesting area for basic studies, and has the potential to reveal organized consortia we have not even dreamt about so far. A model to use as a starting point for further studies is the enrichment consortium of waste water treatment (Nielsen et al. 2009 , Albertsen et al. 2011 ). In the future, when complex organic substrates are to be broken down to complex products to solve challenging problems, fungal and microbial consortia could provide a short-cut to a solution.

Last, but most importantly, further studies are urgently needed to prevent, control and cure serious human mycoses caused by Coccidioides species (Taylor 2006). The number of fatalities caused by this fungal disease, especially in Africa, approaches that caused by tuberculosis and malaria, but far less effort has been invested in preventing and curing human mycoses. Perhaps basic studies of the secretome (including substances bound to the outer wall structures), and new molecular insights, instruments and technologies could bring us a step further in this very difficult significantly under prioritized area of health and pharmaceutical investment.

How to Strengthen International Mycology for a Biobased Future

It is evident from the above descriptions that investment in strengthening mycology globally is worthwhile. The following efforts are urgently needed to enable this to occur:

Understanding the fungal kingdom, through phylogeny and phylogenomics: support for global participation and global coverage of strains used for resolving the Fungal Tree of Life.

Strengthening the mycological platform for classification, identification and strain collection: support implementation of one name one fungus nomenclature; recollection and sequencing of fungi not included in molecular database, including neo- or epitypification where appropriate; develop principles and systems ready for capturing and categorizing sequence discoveries of novel fungal genes not originating from any known species.

Building a global mycology network: develop globally distributed databases, with global input and access; establish a visiting professor programme and a young investigator international exposure programme; establish an IMA delegate participation programme, allowing all regions to participate equally in IMA ExCo meetings and other organizational IMA activities.

Open access mycology: establish global access/ regional collections of bioeconomically-relevant strains and biological materials, unique in that strains and materials are accompanied by searchable information about uses, recipes/ protocols on safe and fast use for advancement of the bioeconomy, e.g. for upgrading biowaste to new biobased products.

The role of fungi in nature, interactions, and the secretome: provide global assistance to mining sequence databases, faster and more function targeted, for recognizing functional proteins and metabolites; strengthen global discovery and stimulate collaboration between industry and academia in all parts of the world.

Mycological heritage programme: we need to know and tell the fantastic history of how fungi have been used since ancient times, the mycological human heritage revitalized through molecular studies and tests of efficacy of traditional uses of fungi, also the use of fungal and microbial consortia for food processing to inspire or provide leads for the next generation of biobased products?

Excellence in teaching and communication: to share best practice, striving for excellence to stimulate and inspire the next generation of mycologists and to attract talent for further development of mycology, including the production of web-based training aids.

1 Intracellular enzymes are too costly to recover from large scale fermentations for industrial purposes.

Aanen DK, de Fine Licht HH, Debets AJM, Kerstes NAG, Hoekstra RF, Boomsma JJ (2009) High symbiont relatedness stabilizes mutualistic cooperation in fungus-growing termites. Science 326 : 1103–1106.

Article CAS Google Scholar

Albertsen M, Hansen LBS, Saunders AM, Nielsen PH, Nielsen KL (2011) A metagenome of a full-scale microbial community carrying out enhanced biological phosphorus removal. ISME Journal 6 : 1094–1106.

Article Google Scholar

Busk PK, Lange L (2013) Function-based classification of carbohydrate-active enzymes by recognition of short, conserved peptide motifs. Applied and Environmental Microbiology 79 : 3380–3391.

Cantarel BL, Coutinho PM, Rancurel C, Bernard T, Lombard V, Henrissat B (2009) The Carbohydrate-Active EnZymes database (CAZy): an expert resource for glycogenomics. Nucleic Acids Research 37 : D233–D238.

Carvalheiro F, Duarte LC, Gírio FM (2008) Hemicellulose biorefineries: a review on biomass pretreatments. Journal of Scientific & Industrial Research 67 : 849–864.

CAS Google Scholar

Chávez R, Bull P, Eyzaguirre J (2006) The xylanolytic enzyme system from the genus Penicillium . Journal of Biotechnology 123 : 413–433.

Dalbøge H, Lange L (1998) Using molecular techniques to identify new microbial biocatalysts. Trends in Biotechnology 16 : 265–271.

FAO (2013) Food Wastage Footprint: impacts on natural resources. Summary Report . Rome: Food and Agriculture Organizations ( http://www.fao.org /docrep/018/i3347e/i3347e.pdf).

Google Scholar

Fowler T, Berka RM (1991) Gene expression systems forfilamentous fungi. Current Opinion in Biotechnology 2 : 691–697.

Frisvad JC, Samson RA (2004) Polyphasic taxonomy of Penicillium subgenus Penicillium. A guide to identification of the food and air-borne terverticillate penicillia and their mycotoxins. Studies in Mycology 49 : 1–173.

Gleason FH, Schmidt SK, Marano AV (2010) Can zoosporic true fungi grow or survive in extreme or stressful environments? Extremophiles 14 : 417–425.

Grigoriev IV, Cullen D, Goodwin SB, Hibbett D, Jeffries TW, et al. (2011) Fueling the future with fungal genomics. Mycology 2 : 192–209.

Grell MN, Jensen AB, Olsen PB, Eilenberg J, Lange L (2011) Secretome of fungus-infected aphids documents high pathogen activity and weak host response. Fungal Genetics & Biology 48 : 343–352.

Grell MN, Linde T, Nygaard S, Nielsen KL, Boomsma JJ, Lange L (2013) The fungal symbiont of Acromyrmex leaf-cutting ants expresses the full spectrum of enzymes to degrade cellulose and other plant cell wall polysaccharides. BMC Genomics 14 : 928.

Hamann T, Lange L (2006) Discovery, cloning and heterologous expression of secreted potato proteins reveal erroneous pre-mRNA splicing in Aspergillus . Journal of Biotechnology 126 : 265–276.

Harris SD, Read ND, Roberson RW, Shaw B, Seiler S, Piamann M. Momany M (2005) Polarisome meets Spitzenkörper: microscopy genetics, and genomics converge. Eukaryotic Cell 4 : 225–229.

Hibbett DS, Binder M, Bischoff JF, Blackwell M, Cannon PF, et al. (2007) A higher-level phylogenetic classification of the Fungi. Mycological Research 111 : 509–547.

Huang Y, Busk PK, Grell MN, Zhao H, Lange L (2014) Identification of a α-glucosidase from the Mucor circinelloides genome by Peptide Pattern Recognition. Enzyme and Microbial Technology 67 : 47–52.

James TY, Kauff F, Schoch CL, Matheny PB, Hofstetter V, et al. (2006) Reconstructing the early evolution of Fungi using a six-gene phylogeny. Nature 443 : 818–822.

Kooij PW (2013) Fungal adaptations to mutualistic life with ants . PhD thesis, Copenhagen University. ( http://www.pwkooij.com /PWKooij/Publications.html).

Kubicek CP, Starr TL, Glass NL (2014) Plant cell wall-degrading enzymes and their secretion in plant-pathogenic fungi. Annual Review Phytopathology 52 : 427–451.

Lange L, Grell MN (2014) The prominent role of fungi and fungal enzymes in the ant-fungus biomass conversion symbiosis. Applied Microbiology and Biotechnology 98 : 4839–4851

Martinez D, Challacombe J, Morgenstern I, Hibbett D, Schmoll M, et al. (2009) Genome, transcriptome, and secretome analysis of wood decay fungus Postia placenta supports unique mechanisms of lignocellulose conversion. Proceedings of the National Academy of Sciences, USA 106 : 1954–1959.

Murphy C, Powlowski J, Wu M, Butler G, Tsang A (2011) Curation of characterized glycoside hydrolases of fungal origin. Database : bar020 doi:10.1093/database/bar020.

Nielsen PH, Kragelund C, Seviour RJ, Nielsen JL (2009) Identity and ecophysiology of filamentous bacteria in activated sludge. FEMS Microbiology Reviews 33 : 969–998.

Nobre T, Eggleton P, Aanen DK (2010) Vertical transmission as the key to the colonization of Madagascar by fungus-growing termites? Proceedings of the Royal Society of London, Biological Sciences 277 : 359–365.

Pitt JI, Hocking AD (2009) Fungi and Food Spoilage . 3 rd edn. New York: Springer.

Book Google Scholar

Rineau F, Roth D, Shah F, Smits M, Johansson T, et al. (2012) The ectomycorrhizal fungus Paxillus involutus converts organic matter in plant litter using a trimmed brown-rot mechanism involving Fenton chemistry. Environmental Microbiology 14 : 1477–1487.

Schoch CL, Shoemaker RA, Seifert KA, Hambleton S, Spatafora JW, Crous PW (2006) A multigene phylogeny of the Dothideomycetes using four nuclear loci. Mycologia 98 : 1041–1052.

Seifert KA, Samson RA, Chapela IH (1995) Escovopsis aspergilloides , a rediscovered hyphomycete from leaf-cutting ant nests. Mycologia 87 : 407–413.

Taylor JW (2006) Evolution of human-pathogenic fungi: phylogenies and species. In: Molecular Principles of Fungal Pathogenesis (Heitman J, Filler SG, Edwards JE jr, Mitchell AP, eds): 113–132. Washington DC: American Society of Microbiology Press.

Zajc J, Liu Y Dai W, Yang Z, Hu J, Gostincar C, Gunde-Cimerman N (2013) Genome and transcriptome sequencing of the halophilic fungus Wallemia ichthyophaga : haloadaptations present and absent. BMC Genomics 14 : 1–21.

Download references

Author information

Authors and affiliations.

Aalborg University, A.C. Meyers Vænge 15, DK-2450, Copenhagen SV, Denmark

You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Lene Lange .

Additional information

Based on an address presented to the 10 th International Mycological Congress (IMC10), Bangkok, 7 August 2014.

Rights and permissions

This article is distributed under the terms of the Creative Commons Attribution 4.0 International License ( https://creativecommons.org/licenses/by-nc/4.0 ), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.

Reprints and permissions

About this article

Cite this article.

Lange, L. The importance of fungi and mycology for addressing major global challenges. IMA Fungus 5 , 463–471 (2014). https://doi.org/10.5598/imafungus.2014.05.02.10

Download citation

Received : 30 September 2014

Accepted : 01 December 2014

Published : 12 December 2014

Issue Date : December 2014

DOI : https://doi.org/10.5598/imafungus.2014.05.02.10

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

fungal research
fungal products
resource efficiency
global solutions
funding opportunities

ISSN: 2210-6359

Submission enquiries: Access here and click Contact Us
General enquiries: [email protected]

A Beginner's Guide to Mycology

Mycology has impacted your life, I guarantee it. Whether it was that penicillin you took to get better or those cultivated mushrooms you ate for dinner yesterday, you have mycology to thank. Mycologists are scientists who dedicate their careers to all things fungi.

It may be surprising to hear that mycology is not a booming discipline. Most universities don’t offer programs in mycology and this doesn’t seem to be changing. Fungal pathogens infect over 1 billion people worldwide. They also destroy one-third of global food production! Why aren’t there more mycologists working on this stuff? On the other hand, humans benefit from many species of fungi, including those we eat and use as medicine.

Fungi come in all sorts of unusual shapes and size. Photo by Andreas via Pixabay .

How Are Mycology, Mycorrhizae, and Mycelia Different?

Mycology is the study of fungus, which is an entire kingdom of life. Mycology is to fungi as botany is to plants. Most scientists consider mycologists to be a type of microbiologist.

To date, scientists have named over 70,000 fungal species. Researchers believe there are up to 20 times that many fungus species out there. This means there could be about 1.5 million species of fungi on planet Earth. For some context, there are only about 350,000 named species of flowering plants. The fungi kingdom is massive and we know so little about it!

Mycelia is simply the vegetative part of fungi. It is a branching, often invisible substance that is home to many wonders. If you have ever seen a white, webby substance in wet woodchips or in the soil, that is mycelium. Mycelia can form vast networks of fungus. One such network is thought to be the largest organism on the planet.

Mushrooms, in contrast, are the fruiting bodies of these mycelial networks. They are the reproductive structures of fungal organisms. If we compare fungus to a peach tree, the mycelia would be the roots, wood, and leaves of the tree. The mushroom would be the flowers and peaches.

Mycorrhizae are the mycelia that grow in a symbiotic relationship with a plant host (more on this later).

Mycelia growing between pine needles. Photo by Laurel Fan via Flickr .

The Origins of Mycology

Humans have an older relationship with fungi than with agriculture. By analyzing plaque on ancient human teeth, scientists revealed that hunter-gatherers in Europe ate mushrooms as early as 18,000 years ago. Ever since then, our species has found hundreds of uses for this kingdom of life.

In the late 19th and early 20th centuries, groups like the Mycological Society of America and the British Mycological Society began to organize mycologists into a broader community. More recent groups, such as the North American Mycological Association, International Mycological Association, and EUROFUNG have bolstered the mycological community. By publishing ground-breaking research in their scientific journals , these societies have aided the development of mycology around the world.

Trends in Mycology

Fungi remain understudied across the globe. There are various reasons for this. First, fungus is inherently difficult to study. Most species of fungi are impossible to grow in a lab. In nature, fungi have a short ‘fruiting’ period where mushrooms are visible. Without mushrooms, it is really hard to find mycelia.

Few universities offer classes or programs that specialize in mycology. Fungi experts were formerly housed within botany departments (which is funny, since fungi are actually more closely related to animals than plants). However, as botany departments are shrinking , they are less interested in supporting their mycologist colleagues. These department-less mycologists have a hard time fending for themselves in university politics.

While the number of amateur mushroom hunters may have increased in the last decade, the number of people studying fungi hasn’t. Some mycologists think that the lack of coordination, cooperation, and specialization within their scientific field is to blame for the decreasing funding towards mycology at universities.

A picture of the underside of a mushroom. Taxonomists often use the ‘gills’ of a mushroom to identify the species of fungus. Photo by Andreas via Pixabay .

What Makes a Fungus?

All fungi share a handful of common characteristics. In taxonomy, fungi are an entire kingdom of life (although the ‘kingdoms’ are always in dispute ).

At the most basic level , fungi have eukaryotic cells. This means that the cells have a nucleus. This differentiates them from prokaryotic bacteria. Fungi are also multicellular. This separates them from eukaryotic bacteria.

Animals and plants are other groups with eukaryotic cells. Fungi are different from animals because fungi can’t move. They also reproduce by spores rather than sexually. Fungi are distinct from plants because fungi can’t make their own food via photosynthesis. Instead, they absorb food from their surroundings. This process is called decomposition.

Lichens are unusual kinds of fungi. They are a symbiotic, obligate relationship between a species of algae and a species of fungi. Where do lichens fit into the tree of life? Beats me.

By far the largest order of fungi is ascomycetes, which contains the vast majority of named fungi species.

Areas of Mycology

From biotechnology to addiction therapy, mycologists cover a wide variety of topics. Fungi touch many aspects of our lives. They are in our daily food, our backyards, and our medicine. Here is a quick overview of some topics mycologists study.

Edible mushrooms can be big business! These chanterelles can cost up to $30 a pound. Image by Gerhard G. via Pixabay .

Edible Fungi

I’m sure you’ve eaten lots of mushrooms if you’ve made it this far into the article. Mushroom cultivation is a booming business as more people turn to plant-based diets. However, outside of a handful of species of fungi, most edible fungi species must be wild-harvested. These species include true delicacies , like chanterelles , truffles , and morels . Mushrooms don’t only taste delicious, they are super healthy, too. A recent study determined that daily mushroom consumption is linked to a 45% decrease in cancer! Some emerging health-food trends include the arctic fungus superfood, Chaga .

Mychorrizae and the Environment

Fungi perform crucial roles in our environment. Every ecosystem needs decomposers. Decomposers are organisms that break down material. Without decomposers, logs and dead animals would never rot into the Earth. Fungi are essential decomposers in the ecology of many habitats.

Fungi also aid plant growth. With the exception of the pea family, plants can’t make their own nitrogen. Nitrogen is an absolutely critical element plants need to grow. Mycorrhizae can convert nitrogen present in our atmosphere to the kind of nitrogen useable by plants. The plants use this nitrogen and in return give the mycorrhizae energy. These symbiotic relationships increase the biodiversity and resiliency of ecosystems on our planet.

Medical Mycology

We should all be thankful for fungi because they are directly responsible for the increase in the human lifespan. The discovery of penicillin in 1928 had profound consequences on the human species. Penicillin works by distracting the enzyme in bacteria that produce cell walls. This distraction results in dead, disease-causing bacteria. Penicillin led to the antibiotic revolution. These drugs were a large part of increasing the human lifespan by nearly 30 years in places such as the U.S.

A fungal disease of bananas causing cigar end-rot. Photo by Scot Nelson via Flickr .

Pathogenic Fungi

Fungi are the largest source of plant disease. These fungi cause diseases in many ways. They can enter a plant through leaves, bark, or roots. Some economically devastating fungi can destroy entire harvests of the world’s most important crops. These crops include rice, corn, cereals, bananas, tomato, cotton, and so many more. These fungi are serious business. It’s no wonder that many of the employed mycologists focus on plant pathology.

Entheogenic (a.k.a. Magic) Mushrooms

Yes, those kinds of mushrooms. Psilocybin is a hallucinogenic compound found in certain species of fungi. This group of mushrooms is found all around the world. The first evidence of human use of these mushrooms dates back 7000-9000 years ago. This evidence is based on trippy prehistoric rock art in northern Africa.

Today there is a research boom in these powerful mushrooms. There is strong, emerging evidence that psilocybin can drastically help people with addiction and depression . In a few decades, we may see a much broader acceptance of these medicinal mushrooms.

Why Aren’t There More Mycologists?

Short answer: I don’t know. It really seems like we need more people studying these species. Fungi can be wildly beneficial to humans and also incredibly destructive. Maybe they are hard to research. Maybe research into fungi just isn’t sexy enough for government or private funding. Whatever the case, it seems like an increase in people studying fungal biology would help the human species as a whole.

Featured image by Steve Buissinne via Pixabay

Check us out on EarthSnap , a free app brought to you by Eric Ralls and Earth.com.

An official website of the United States government

The .gov means it’s official. Federal government websites often end in .gov or .mil. Before sharing sensitive information, make sure you’re on a federal government site.

The site is secure. The https:// ensures that you are connecting to the official website and that any information you provide is encrypted and transmitted securely.

Publications
Account settings

Preview improvements coming to the PMC website in October 2024. Learn More or Try it out now .

Advanced Search
Journal List
v.3(1); 2012 Jun

The importance of fungi and of mycology for a global development of the bioeconomy

Institute of Biotechnology and Chemistry, Faculty of Science and Technology, Aalborg University, Denmark

Peter K. Busk

Morten n. grell, yuhong huang, mette lange, bo pilgaard, xiaoxue tong.

The vision of the European common research programme for 2014–2020, called Horizon 2020 , is to create a smarter, more sustainable and more inclusive society. However, this is a global endeavor, which is important for mycologists all over the world because it includes a special role for fungi and fungal products. After ten years of research on industrial scale conversion of biowaste, the conclusion is that the most efficient and gentle way of converting recalcitrant lignocellulosic materials into high value products for industrial purposes, is through the use of fungal enzymes. Moreover, fungi and fungal products are also instrumental in producing fermented foods, to give storage stability and improved health. Climate change will lead to increasingly severe stress on agricultural production and productivity, and here the solution may very well be that fungi will be brought into use as a new generation of agricultural inoculants to provide more robust, more nutrient efficient, and more drought tolerant crop plants. However, much more knowledge is required in order to be able to fully exploit the potentials of fungi, to deliver what is needed and to address the major global challenges through new biological processes, products, and solutions. This knowledge can be obtained by studying the fungal proteome and metabolome; the biology of fungal RNA and epigenetics; protein expression, homologous as well as heterologous; fungal host/substrate relations; physiology, especially of extremophiles; and, not the least, the extent of global fungal biodiversity. We also need much more knowledge and understanding of how fungi degrade biomass in nature.

The projects in our group in Aalborg University are examples of the basic and applied research going on to increase the understanding of the biology of the fungal secretome and to discover new enzymes and new molecular/bioinformatics tools.

However, we need to put Mycology higher up on global agendas, e.g. by positioning Mycology as a candidate for an OECD Excellency Program. This could pave the way for increased funding of international collaboration, increased global visibility, and higher priority among decision makers all over the world.

INTRODUCTION

Horizon 2020 is a visionary document for the European common research programme 2014–2020 ( http://ec.europa.eu/research/horizon2020 ). The vision is to create a smarter, more sustainable and more inclusive society. However, such endeavor is not only European. It is global. Most importantly, for the members of the International Mycological Association (IMA), it includes a special role for fungi and fungal products. Therefore, it is an agenda of special relevance for mycologists all over the world.

The Horizon 2020 document emphasizes that the most important goals and objectives for common research programmes are to address the major global challenges. Among the challenges of priority is climate change, the need for increased efficiency in resource utilization, and the urgency of developing renewable substitutes for fossils; and not least to provide for improved human health – combating life- style diseases and ensuring food security for a rapidly growing population. Essential for overcoming much of these challenges is improved use of natural resources; especially biological resources, plant nutrients and water. Regarding the efficient use of bioresources, we can do much better: After harvesting the food and feed, crop residues beyond what is needed to sustain a productive and healthy soil, are left to rot or burned. Further, the potentials of side streams and waste streams from agroindustries often remain unexploited. Also, the organic part of municipality waste is deposited in landfills, burned, or used as combustion feed stock in power plants. However, biomaterials are much too precious for such low value uses. We need more upgraded use of bioresources to both feed the growing population and as a substitute for what we now get from fossil resources.

THE POTENTIAL ROLE OF FUNGI

In nature, the breakdown of plant materials is primarily by fungi, by the means of secreted fungal enzymes. Driven by the urge for non-food based bioenergy, industrial scale conversion of biowaste has been researched and developed over the last ten years. After all this research, the conclusion is that the most efficient and gentle way of converting recalcitrant lignocellulosic materials for industrial purposes, is through the use of fungal enzymes ( Lange 2010 ). Through such conversions, the building blocks of the organic materials are kept intact, ready to use in the value cascade ( Fig. 1 ). The enzymatic conversion of biowaste and -sidestreams will provide the basis for an entirely new way for the more efficient use of natural resources, paving the way for a larger bioeconomy sector in a more biobased society:

An external file that holds a picture, illustration, etc.
Object name is ima-3-87-g001.jpg

The Biomass Value Pyramid shows the entire cascade of value adding products which can be produced from agricultural crop residues and other left over bio materials. The lowest value is achieved by burning the biomass and converting it into heat and electricity. Higher value products can be achieved by converting the biomass through treatment with fungal enzymes!

Plant materials, obtained as crop residues, municipality waste, or from agroindustrial waste streams, will increasingly substitute for fossil carbon from crude oil. Not just substituting fossil energy with bioenergy, but more importantly also substituting the higher value fossil-based materials, such as plastics and chemical building blocks, with biomaterials made from the sugar molecules of the plant cell wall polymers. Thus, the conversion of biowaste is primarily the conversion of plant cell wall materials into higher value products; achieved primarily by a process based on the refined use of fungi and fungal enzymes.

But fungi are providing even more of the solutions for meeting and addressing the various global challenges: Fungi and fungal products are also instrumental in producing fermented foods, to give storage stability and improved health; and it is a fungus (baker’s yeast) that is the production organism of choice for producing insulin for the global population of diabetics. Also, cholesterol lowering drugs (the statins), major immunosuppressant drugs (cyclosporins), the cancer drug Taxol, and penicillins are fungal products.

Climate change will lead to increasingly severe stress on agricultural production and productivity, and here the solution may very well be that fungi will be brought in use as a new generation of agricultural inoculants (e.g. mycorrhizas, endophytes, biocontrol agents) to provide more robust, more nutrient efficient, and more drought-tolerant crop plants.

POSITIONING MYCOLOGY IN THE WORLD

Mycologists have over time delivered so much knowledge about fungi (taxonomy, physiology, genetics, host/substrate relations (including plant pathology and studies of biotrophic interactions), molecular biology, metabolites, enzymes and protein expression) that biological products, biological processes, and biological solutions to important problems are already widespread within many industrial sectors. To mention a few: fungal enzymes are instrumental in laundry detergents at lower temperature and in the less polluting production of both paper and textiles, by replacing chemical processing. Thus, our knowledge and insight into fungal growth and fungal products (proteins as well as metabolites) have made biological processes competitive against chemical processes because they have been developed to be both highly efficient and safe. However, much more knowledge is required in order to be able to fully exploit the potentials of fungi, to deliver what is needed, and to address the major global challenges through new biological processes, products, and solutions.

Additional basic knowledge about fungi is required across an entire spectrum of research fields: The fungal proteome and metabolome; the biology of fungal RNA and epigenetics; protein expression, homologous as well as heterologous; fungal host/substrate relations; physiology, especially of extremophiles; and not least the extent of global fungal biodiversity. Indeed, many of the new applications of fungi and fungal products will be made possible through “unlocking the magic” of fungi we have not yet discovered – let alone described, characterized, or classified.

We also need much more knowledge and understanding of how fungi degrade biomass in nature, and especially on how they interact with each other and with microorganisms, especially bacteria. In order to achieve all this, we need to train next generation of mycologists to be experts in their fields, mastering both the new and the classical methods. Besides researchers, we also need to train the skilled workers in how to handle biological production at the industrial scale. Last but not least, we need skilled and enthusiastic teachers at all levels who can teach about the fascinating world of the fungi, both the friends and the foes, from kindergartens to graduate schools.

THE WAY FORWARD

As a first step forward, we propose a specific global learning loop for knowledge sharing of relevance to speeding up the application of mycology in addressing issues of global concern.

Most importantly, we see that we need to start to change our mindset as mycologists, taking the importance of fungi and fungal products seriously in our personal research agendas. Not with the objective of making all of us to work in applied mycological research in a traditional sense, but recognizing that we also need blue-sky, curiosity-, biodiversity- and exploration- driven research within mycology – perhaps more than ever before, in order to realize the huge potential.

To this end, mycologists in our research group in Aalborg University, Denmark, located on the AAU Copenhagen Campus, now orient our research projects to have a double focus, to: (1) forward the scientific field in which we are working, by increased understanding of the biology of the fungal secretome (regulation, composition and function); and (2) discover new enzymes and new molecular/bioinformatics tools, thereby contributing to the development of new biological products, biological processes, and biological solutions to important problems. Examples of activities with such a double focus, both basic and applied are:

The phylogeny of a fungal cellulase

A comparative study of an endoglucanase belonging to protein family GH45, gave surprising results, which lead to a new enzyme discovery approach: A phylogenetic analysis of the GH45 proteins, from all parts of the fungal kingdom, asco-, basidio-, zygo-, and chytridiomycetes ( Kauppinen et al . 1999 ), indicated that distantly related fungi, such as the basidiomycete Fomes fomentarius and the ascomycete Xylaria hypoxylon , had GH45 cellulases in their secretome with an extremely high similarity in the amino acid composition of their active site. Strikingly, both these fungi inhabit and decompose very similar substrates (hard wood). A similar pattern can be seen amongst straw decomposing fungi, for example the basidiomycete Crinipellis scabella and the chytrid Rhizophlyctis rosea . These two fungi are from two very different parts of the fungal kingdom. Anyway, their GH45 cellulase proteins have an almost identical amino acid composition of their active sites. These observations can tentatively be explained by the following molecular mechanism: Evolution of the fungal GH45 is impacted by gene copying and subsequent gene loss, maintaining the version of the gene which is most suitable for breaking down the cellulose of the substrate of the fungus. This conclusion provided the basis for a new screening approach: select a relevant ecological niche in nature with regard to type of substrate, temperature, and pH; construct a meta-library of the entire microbial (fungal and bacterial) community at such a site; and screen this library for the best enzyme candidates for industrial applications. It also inspired the following hypothesis: evolution of the fungal secretome composition may be interpreted as taking place at the molecular level rather than at the organismal level.

Peptide pattern recognition (PPR)

A new method has been invented for the improved prediction of protein function from protein sequences. It is unique in being non-alignment-based, and permits the comparison of a vast number of sequences with even very low sequence identities. PPR analysis is potent for revealing new protein subfamily groupings, where the subgrouping is correlated with a specific function ( Fig. 2 ). Such new understanding can again be used to understand the biological role of the secreted proteins, interactions between organisms, and interactions between the organism and the substrate. A new subfamily can be described by a list of peptides that is specific to just that subfamily. PPR analysis, moreover, opens the possibility of finding more of a given type of functional proteins belonging to a single subfamily. This can be done by using the conserved peptides for discovering new subfamily members, either by following a bioinformatics approach or by screening biological materials with degenerated primers, constructed based on the list of the identified most conserved peptides ( Busk & Lange 2011 , 2012).

An external file that holds a picture, illustration, etc.
Object name is ima-3-87-g002.jpg

The Peptide Pattern Recognition, (PPR), generated GH13 protein subfamilies which predicted the enzyme function correctly with 78–100 % accuracy; except for one enzyme class (3.2.1.133) where for so far unknown reasons the PPR subgrouping did not match the function annotations found in the CAZy database.

We analyzed 8138 GH13 proteins represented in the B. Henrissat CAZy database with PPR to generate subfamilies. The subfamily-specific peptide lists were used to predict the function of 541 functionally characterized GH13 proteins. Overall, the function of 85 % of the proteins was correctly predicted ( Fig. 2 ). The figure shows the percentage correct prediction of the enzymatic functions for each of the enzyme classes (new data; P.K. Busk & L. Lange, unpubl.).

Fungal decomposition of specific substrates

Understanding enzymatic degradation of plant cell wall materials is improved by studying in parallel both the plant cell wall composition (by the CoMPP technology, Moller et al . 2007 ) and the fungal secretome enzymes of the fungus responsible for the degradation. The materials under study in a Chinese/Danish research project are duckweed ( Cheng & Stomp 2009 ; Fig. 3 ) and industrial pulp of non-food uses of basic rhizomes such as sweet potato ( Zhang et al . 2011 ), cassava, and Canna edulis . Using next generation sequencing, the transcriptomes of tropical fungal species, isolated from relevant substrates, are analyzed and novel enzymes are expected to be identified. The secretomes will be further characterized to compare the phylogenetic relationships of the secretome proteins as compared to the phylogenetic relationships of the organisms (L. Bech, Y. Huang, Z. Hai, P.K. Busk, W.G.T. Willats, M.N. Grell, and L. Lange, unpubl.).

An external file that holds a picture, illustration, etc.
Object name is ima-3-87-g003.jpg

When grown in swine wastewater, some duckweed species such as the Spirodela polyrhizza contains up to 40 % protein, which makes it a valuable animal feed source. Picture by courtesy of Armando Asuncion Salmean.

Accessory proteins

In 2011 it was discovered that proteins of family GH61 act directly on crystalline cellulose, partially degrading and loosening the structure of the microfibrils, thereby increasing the substrate accessibility for other types of cellulases ( Beeson et al. 2012 , Langston et al. 2011 , Quinlan et al . 2011 , Westereng et al . 2011 ). The PPR analysis of all publicly available GH61 sequences resulted in a tentative subgrouping in 16 new subfamilies. We are now studying the possible correlation of such subfamily groupings with the function of the given GH61 proteins, attempting to answer the following biological questions: What is the function and role of the high number of very different GH61 genes, as is so commonly seen among plant cell wall degrading fungi? We wish to increase understanding of the biological role of these non-hydrolytic accessory proteins in nature; and to provide a basis for choosing which GH61 subfamily proteins should be incorporated into new and improved industrial enzyme blends for conversion of lignocellulosic biomasses into free sugars (M. Lange, P.K. Busk, and L. Lange , unpubl.).

Enzymes from thermophilic fungi

Are the enzymes of thermophilic fungi more thermotolerant than those of mesophilic fungi? We are attempting to answer this fundamental physiological question, and at the same time provide a basis for developing a new type of biomass conversion process which can function at high temperatures, in order to improve the efficiency of the added enzymes and to speed up the biomass conversion ( Busk & Lange 2011 ).

A molecular analysis of biomass conversion in the leaf-cutter ant fungal garden

The fungal garden of leaf-cutter ants constitutes a natural biomass conversion system ( Fig. 4 ). Mediated by fungal secreted enzymes, leaf fragments brought into the nest by the ants are converted to food for the ant larvae as well as serve as substrate for fungal growth. In this study, we investigated which enzymes are produced and their relative expression level along the decomposition gradient of the garden structure ( Fig. 4 ), using the DeepSAGE method. DeepSAGE is a global digital transcript-profiling technology, facilitating measurement of rare transcripts ( Nielsen et al. 2006 ). The results of the study have given us interesting new molecular insights into a social insect-fungus symbiosis that relies on conversion of a fresh leaves biomass, recalcitrant to degradation (M.N. Grell, K.L. Nielsen, T. Linde, J.J. Boomsma, and L. Lange, unpubl.). Now the question arises: what can we learn from the type of biomass degradation that the fungus growing leaf cutter ants have developed so successfully?

An external file that holds a picture, illustration, etc.
Object name is ima-3-87-g004.jpg

Leaf-cutter ant colony established in the laboratory of JJ Boomsma (University of Copenhagen). The ants have built three fungal gardens under plastic beakers. The beaker has been removed from the garden to the upper right. The gradient of biomass decomposition, from top to bottom, is indicated by the green arrow. The dark material on the surface of the garden is newly incorporated leaf fragments. Non-degraded material is removed by the ants from the bottom of the garden and placed in the refuse dump (upright beaker to the lower right) (photo, Morten N. Grell).

The subgrouping of esterases and their possible function in biomass conversion in nature

At present we focus on a study of additional and so far almost neglected types of enzymes needed for full biomass conversion, more specifically, on the esterases, especially the ferulic acid esterases (X. Tong, P.K. Busk, M.N. Grell, and L. Lange, unpubl.). A feature of plant cell wall polysaccharides is that they are able to cross-link, and that such cross-links can include phenolic groups represented by ferulic acid (feruloyl). The ferulic acid units can be oxidatively cross-linked by cell wall peroxidases into other polysaccharides, proteins and lignin. This cross-linking increases plant resistance to microbial degradation. The enzymes responsible for cleaving the ester-link between the polysaccharide main chain of xylans and either monomeric or dimeric feruloyl are the ferulic acid esterases (EC 3.1.1.73). The breakage of one or both ester bonds from dehydrodimer cross-links between plant cell wall polymers is essential for optimal action of carbohydrases on substrates such as cellulosic biomass. Subfamily groupings within the field of lipases and esterases are still disputed and unresolved. We attempt to use the PPR method also within these types of enzymes, to provide increased insight in the fungal secretome by achieving function-related subgroupings also of this class of enzymes; and to elucidate further the role also of esterases in biomass conversion.

Studies of secreted enzymes from edible wood-decaying fungi

These studies aim at providing a basis for onsite production of enzyme blends for biomass conversion. Edible basidiomycetes, such as Pleurotus ostreatus , are chosen because they do not produce mycotoxins which would prohibit their use as production organisms; and because they have been shown to have the potential to secrete sufficient biomass degrading enzymes, to significantly lower the need for commercial enzyme blends in the production of second generation biofuels. Some even produces secondary metabolites with potential for use in other industries. The combination of these attributes can provide a significant cost reduction of the final products and, most importantly, open for decentralized low-investment use of biorefinery technologies for the production of animal feed, fertilizer, and fuel from crop residues (B. Pilgaard, L. Bech, M. Lange, and L. Lange, unpubl.).

The evolution of obligate insect pathogens, elucidated by studies of their secreted enzymes

In an earlier secretome study of field-collected grain aphids ( Sitobion avenae ) infected with fungi of the order Entomophthorales (subphylum Entomophthoromycotina), we identified a number of pathogenesis-related, secreted enzymes ( Grell et al. 2011 ). Among these were cuticle degrading serine proteases and chitinases, involved in fungal penetration of the aphid cuticle, and a number of lipases most likely involved in nutrient acquisition. In a continuation of this study, we are investigating the distribution and variation of selected enzyme-encoding genes within the genera Entomophthora and Pandora , using fungal genomic DNA originating from field-collected, infected insect host species of dipteran (flies, mosquitoes) or hemipteran (aphid) origin. We anticipate that this study will shed new light on this highly specialized group of enthomophthoralean insect pathogenic fungi and their secreted enzymes (M.N. Grell, A.B. Jensen, J. Eilenberg, and L. Lange, unpubl.).

Evidence for a new biomass conversion role of ectomycorrhizal fungi and their use of a chemical mechanism for biomass conversion

The ectomycorrhizal fungus Paxillus involutus converts organic matter in plant litter using a trimmed brown-rot mechanism involving both enzymatic activities and Fenton chemistry (Rineau et al . 2012). These results could serve as a model for future industrial biomass conversion, combining chemistry and biology to achieve more efficient biomass conversion.

Studies of the cellulases of the aerobic soil chytrids

Pilgaard et al. (2011) have provided insight in the roots and origin of the fungal cellulases by studying the cellulases of aerobic soil inhabiting chytrids; and we are also attempting to further elucidate the aerobic chytrid secretome potentials for industrial exploitation of this unique group of fungi, so far almost totally neglected.

In order to achieve the goal of more mycological knowledge brought into use, for a more sustainable world of tomorrow, where the bioeconomy becomes an important pillar for our global society, we need fungi to be recognized with heightened visibility. They need to be higher up on global agendas. One way towards that goal could be to position Mycology as a candidate for an OECD Excellency Program. This could pave the way both for increased (national and international) funding of international collaboration, increased global visibility, and hopefully higher priority among decision makers all over the world. We hope you as mycologists, and the IMA as a global institution, will work together towards realizing this vision.

Acknowledgments

Funding for the above mentioned projects have been supplied by: The Danish Council for Strategic Research (BioRef, Bio4Bio; FunSecProt) and Novozymes. We further thank Peter Westermann, Associate Professor at Aalborg University, for letting us use his figure on the biomass cascade, and Armando Asuncion Salmean Ph. D. student at University of Copenhagen, for his picture of duckweed.

Beeson WT, Phillips CM, Cate JH, Marletta MA. (2012) Oxidative cleavage of cellulose by fungal copper-dependent polysaccharide monooxygenases . Journal of the American Chemical Society 134 : 890–892 [ PubMed ] [ Google Scholar ]
Busk PK, Lange L. (2011) A novel method of providing a library of n-mers or biopolymers . Patent application EP11152232.2 [ Google Scholar ]
Busk PK, Lange L. (2011) Novel glycoside hydrolases from thermophilic fungi . Patent Application EP11152252.0. [ Google Scholar ]
Cheng JJ, Stomp A-M. (2009) Growing duckweed to recover nutrients from wastewaters and for production of fuel ethanol and animal feed . CLEAN - Soil, Air, Water 37 : 17–26 [ Google Scholar ]
Grell MN, Jensen AB, Olsen PB, Eilenberg J, Lange L. (2011) Secretome of fungus-infected aphids documents high pathogen activity and weak host response . Fungal Genetics and Biology 48 : 343–352 [ PubMed ] [ Google Scholar ]
Kauppinen S, Schülein M, Schnorr K, Lassen S.F, Andersen KV, Urs SK, Katila P, Lange L. (1999) Comparative analysis of a cellulase gene (Cel45) found in all major fungal groups . 22 nd Fungal Genetics Conference at Asilomar : Abstract. [ Google Scholar ]
Lange L. (2010) The importance of fungi for a more sustainable future on our planet . Fungal Biology Reviews 24 : 90–92 [ Google Scholar ]
Langston JA, Shaghasi T, Abbate E, Xu F, Vlasenko E, Sweeney MD. (2011) Oxidoreductive cellulose depolymerization by the enzymes cellobiose dehydrogenase and glycoside hydrolase 61 . Applied and Environmental Microbiology 77 : 7007–7015 [ PMC free article ] [ PubMed ] [ Google Scholar ]
Moller I, Sørensen I, Bernal AJ, Blaukopf C, Lee K, Øbro J, Pettolino F, Roberts A, Mikkelsen JD, Knox JP, Bacic A, Willats WGT. (2007) High-throughput mapping of cell-wall polymers within and between plants using novel microarrays . The Plant Journal : for cell and molecular biology 50 : 1118–1128 [ PubMed ] [ Google Scholar ]
Nielsen KL, Hogh AL, Emmersen J. (2006) DeepSAGE - digital transcriptomics with high sensitivity, simple experimental protocol and multiplexing of samples . Nucleic Acids Research 34 : e133 [ PMC free article ] [ PubMed ] [ Google Scholar ]
Pilgaard B, Gleason F, Lilje O, Lange L. (2011) Lignocellulosic enzymes in three species of zoosporic fungi from NSW soils . 2011 Annual Conference of the Ecological Society of Australia : Abstract. [ Google Scholar ]
Quinlan RJ, Sweeney MD, Leggio LL, Otten H, Poulsen J-CN, Johansen KS, Krogh KBRM, Jørgensen CI, Tovborg M, Anthonsen A, Tryfona T, Walter CP, Dupree P, Xu F, Davies GJ, Walton PH. (2011) Insights into the oxidative degradation of cellulose by a copper metalloenzyme that exploits biomass components . Proceedings of the National Academy of Sciences, USA 108 : 15079–15084 [ PMC free article ] [ PubMed ] [ Google Scholar ]
Westereng B, Ishida T, Vaaje-Kolstad G, Wu M, Eijsink VGH, Igarashi K, Samejima M, Ståhlberg J, Horn SJ, Sandgren M. (2011) The putative endoglucanase PcGH61D from Phanerochaete chrysosporium is a metal-dependent oxidative enzyme that cleaves cellulose . PLoS ONE 6 : e27807 [ PMC free article ] [ PubMed ] [ Google Scholar ]
Zhang L, Zhao H, Gan MZ, Jin YL, Gao XF, Chen Q, Guan JF, Wang ZY. (2011) Application of simultaneous saccharification and fermentation (SSF) from viscosity reducing of raw sweet potato for bioethanol production at laboratory, pilot and industrial scales . Bioresource Technology 102 : 4573–4579 [ PubMed ] [ Google Scholar ]

Biology Article

Mycology is the study of fungi and their unique relationships with other organisms and the environment. It deals with the genetic and biochemical properties of fungi and their importance in human lives.

Important Topics in Mycology

Fungi are eukaryotic organisms that belong to their own kingdom. They were initially included under kingdom Plantae . But recent researches have revealed that fungi are a separate lineage of eukaryotes which are distinguished by their cell wall structure.

Mycology is an important science in the agricultural industry. Fungi act as major pests for many crops but also live in symbiotic association with many plants and provide them with nutrition and water. Mycology helps to differentiate between useful and harmful fungi and how the crops with fungal infections can be treated.

Fungi produce various toxins that are harmful to other organisms. Most of the mushrooms produce toxins. While a few have medicinal properties. Yeat is another form of fungi which is used in brewing, distilling and bread making.

The study of fungi has focussed on various other applications of fungi including their property of alleviating radioactive waste. Fungi can also breakdown complex substances to produce carbon dioxide.

Fungi have been used as food, medicine, and a variety of other things. The field of mycology is really vast and can be understood further under the list of articles mentioned above.

BIOLOGY Related Links

Register with BYJU'S & Download Free PDFs

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

View all journals
Explore content
About the journal
Publish with us
Sign up for alerts
Review Article
Open access
Published: 12 September 2022

The importance of antimicrobial resistance in medical mycology

Neil A. R. Gow ORCID: orcid.org/0000-0002-2776-5850 1 ,
Carolyn Johnson ORCID: orcid.org/0000-0002-0494-8136 2 ,
Judith Berman 3 ,
Alix T. Coste ORCID: orcid.org/0000-0001-9481-9778 4 ,
Christina A. Cuomo ORCID: orcid.org/0000-0002-5778-960X 5 ,
David S. Perlin ORCID: orcid.org/0000-0002-1268-5524 6 ,
Tihana Bicanic ORCID: orcid.org/0000-0002-2676-838X 7 , 8 ,
Thomas S. Harrison 1 , 7 , 8 ,
Nathan Wiederhold ORCID: orcid.org/0000-0002-2225-5122 9 ,
Mike Bromley ORCID: orcid.org/0000-0002-7611-0201 10 ,
Tom Chiller 11 &
Keegan Edgar 11

Nature Communications volume 13 , Article number: 5352 ( 2022 ) Cite this article

19k Accesses

95 Citations

122 Altmetric

Metrics details

Antifungal agents
Antimicrobial resistance
Clinical microbiology

Prior to the SARS-CoV-2 pandemic, antibiotic resistance was listed as the major global health care priority. Some analyses, including the O’Neill report, have predicted that deaths due to drug-resistant bacterial infections may eclipse the total number of cancer deaths by 2050. Although fungal infections remain in the shadow of public awareness, total attributable annual deaths are similar to, or exceeds, global mortalities due to malaria, tuberculosis or HIV. The impact of fungal infections has been exacerbated by the steady rise of antifungal drug resistant strains and species which reflects the widespread use of antifungals for prophylaxis and therapy, and in the case of azole resistance in Aspergillus , has been linked to the widespread agricultural use of antifungals. This review, based on a workshop hosted by the Medical Research Council and the University of Exeter, illuminates the problem of antifungal resistance and suggests how this growing threat might be mitigated.

Tackling the emerging threat of antifungal resistance to human health

The rapid emergence of antifungal-resistant human-pathogenic fungi

Molecular mechanisms governing antifungal drug resistance

Introduction and perspective.

The age of antibiotics, spanning only 80 years, is now entering a period of progressive and widespread emergence of drug-resistant organisms that threaten to bring this era to an end 1 , 2 , 3 , 4 . Microbial pathogens, including fungi, tend to have short generation times, plastic genomes, and the ability to adapt to natural environments that contain many potentially toxic compounds, which exert strong selective pressures. The eukaryotic biochemistry of fungi makes them particularly pernicious pathogens because of a more limited number of selective drug targets against which inhibitors can be designed that are non-toxic for human, animal, and plant hosts. Furthermore, no therapeutic vaccines or adjunct immunotherapies are available to support human health care; this necessitates reliance on a limited armoury of antifungal drug classes to treat a rising tide of fungal infections. These challenges are exacerbated by the emergence of drug resistant, tolerant, or insensitive organisms and increasing numbers of susceptible hosts. Resistant strains of fungi have been identified shortly after the introduction of new antifungal drugs and despite new antifungals in the pipeline 5 , once they are introduced clinically, we should anticipate that resistance will ultimately emerge unless mitigating strategies are deployed. Resistance is the result of genetic mutations and induced protective mechanisms. Rapid plasmid-mediated spread of resistance has not been detected in fungi (as opposed to bacteria). However, antifungal resistance and tolerance can be acquired rapidly, often by the induction of protective stress response pathways, sometimes including the acquisition of aneuploidy or other forms of copy number variation. Thus, the emergence of fungal strains and species with single or multiple drug resistance profiles poses significant challenges in the treatment of medical, veterinary and agricultural hosts 6 , 7 , 8 , 9 , 10 .

Of the estimated five million species of fungi, less than 100 species are frequent agents of human disease, and most deaths are due to organisms within the genera Candida , Aspergillus and Cryptococcus 3 . However, a cadre of new emerging pathogens are rising in clinical importance, and these include some highly drug-resistant species, including Scopulariopsis and Lomentospora 11 . Antifungal resistance. This can be a consequence of the response to patient antifungal treatment, but many human pathogenic fungi also have an environmental phase where resistance can emerge 12 . For example, antifungal resistance in Aspergillus fumigatus is clearly associated with environmental selection of resistance as a consequence of exposure to agricultural azoles used in crop protection 13 . Indeed, estimates suggest that one in 20 culturable isolates of this fungus isolated from the air are tebuconazole resistant 14 . Some strains of Candida glabrata , Candida krusei , most strains of Scedosporium and the Mucorales, and the recent emergent species Candida auris display reduced susceptibility to commonly used antifungals. The problems of antifungal resistance are compounded by problems of late diagnosis and consequently treatment delays. Very high levels of morbidity and mortality 1 are associated with comorbidities, (e.g., haematological malignancies, solid organ transplantation, ICU stays, HIV, SARS-CoV-2, and influenza), rising numbers of susceptible hosts, host immune status, drug accessibility, drug tolerance, treatment with biologics and the formation of fungal biofilms. Life-threatening fungal infections also tend to be prevalent in resource-limited areas of the world with fewer health care options, including access to antifungal diagnostics and drugs. Low- and middle-income countries face additional challenges, including indiscriminate use of antifungal drugs, and limited stewardship 15 , 16 . Cumulatively, these factors result in hundreds of millions of serious fungal infections and between 1 and 1.5 million attributed fungal infection-related deaths per year 1 , 2 .

This review summarizes the conclusions of a workshop hosted by the Medical Research Council and the University of Exeter in May 2021. The workshop brought together a group of medical mycologists with diverse research interests (Supplementary Table 1 ) to outline the scale of the threat and the opportunities to mitigate the consequences of antifungal resistance.

Mechanisms of antifungal resistance and tolerance

The number of fungal infections has continued to increase over the past 20 years, due, in part, to improved enumeration and identification of fungal infections by international organizations (e.g., GAFFI and SENTRY). In addition, the rate of antifungal resistance in yeasts continues to rise globally 17 , 18 , alongside the increased emergence of non- albicans Candida species 18 . The recent emergence of multidrug-resistant yeasts 19 , such as C. auris and C. glabrata is reminiscent of the situation with bacteria. Antifungal resistance in filamentous fungi, notably A. fumigatus 20 , has been linked to the increased use of antifungal agents, particularly azoles, both in the environment and in the clinic. Over the past 20 years, terbinafine-resistant strains of Trichophyton spp. have emerged in India, and 13% of these isolates are also resistant to azoles 21 , 22 .

Sensu-stricto , antifungal drug resistance, like antibacterial drug resistance, is the ability of a fungal isolate to grow well in the presence of drug concentrations that inhibit the growth of most isolates of that species. To formalize and quantify susceptibility for clinical microbiology labs, two major consortia, the Clinical & Laboratory Standards Institute (CLSI) and the European Committee on Antimicrobial Susceptibility Testing (EUCAST) have defined breakpoints as the minimal inhibitory concentration (MIC) of a drug, above which an isolate is considered resistant to clinical treatment, as well as epidemiologic cut-off values (ECVs or ECOFFs) that define the upper limit of the wild type susceptible population when breakpoints are unavailable 23 . Drug-resistant isolates are more likely to fail treatment and to cause breakthrough infections 24 .

Antifungal drug resistance is usually due to stable and heritable point mutations or insertions/deletions that directly affect the interaction of the drug with its target (Fig. 1 ) 20 , 25 . In addition to antifungal drug resistance, several more subtle drug responses that may have clinical significance have been studied primarily in yeasts. These include tolerance, heteroresistance, biofilm formation, aneuploidy, and persistence (reviewed in ref. 26 ).

Antifungal drug resistance (left side) is detected as elevated MIC due to direct effects on drug (orange circle) or drug target (blue star), via reduced binding affinity of the target for the drug, increased levels of the target that dilute the drug effect, or by reducing the intracellular drug concentration via drug efflux or blocked drug uptake. Antifungal drug tolerance (right side) is a physiological response to drug stress involving pathways that buffer the stress, such that some cells are able to grow, albeit slowly, in the presence of drug concentrations that are inhibitory to other cells in the population. This involves physiological shifts in: the cell wall or membrane integrity pathways (including pathways regulated by Hsp90, calcineurin, and the Crz1 transcription factor, and pathways affecting membrane lipid composition); protein translation machinery including the TOR pathway; and modifications of mitochondrial function. Loss of mitochondrial DNA in tolerant species (e.g., C. glabrata and Saccharomyces cerevisiae) , also leads to high drug efflux via Pdr1 and drug resistance, but cellular fitness is highly compromised in these ‘petite’ isolates, which are therefore not thought to be clinically relevant. Heteroresistance (across top) is a semi-stable mechanism, often due to whole chromosome aneuploidy, that can confer either resistance (increased MIC), via increased expression of a target or of efflux pumps, or tolerance (susceptible MIC but increased growth in drug) via altered stress response pathways. Biofilms (bottom) are a sessile physiological state that grows slowly and exhibits drug resistance and/or tolerance due to multiple mechanisms, including sequestration of the drug by large amounts of extracellular matrix. Aneuploidy, gene amplification, copy number variation and loss of heterozygosity (LOH) can confer resistance or tolerance, depending on the specific genes and combinations of genes that are involved.

Antifungal drug tolerance, often termed ‘trailing growth’ in clinical studies, appears as partial growth after >24 h in susceptibility assays, because tolerance is due to the slow growth of some cells in the population that eventually grow in inhibitory drug concentrations 27 . When the growing cells are re-tested, again only some progeny cells grow, implying that tolerance is a physiological or epigenetic phenomenon or that it is transient. Aneuploidy can confer resistance or tolerance as well as cross-tolerance and appears in response to a range of drugs and pathogenic yeast species 28 , 29 , 30 , 31 , 32 , 33 and, like copy number variation, is maintained primarily under drug pressure. Among the specific genes that affect tolerance are genes encoding transcription factors Czf1 (ref. 34 ) and Gzf3 (ref. 35 ), an iron acquisition factor Iro1 (ref. 36 ) and sphingolipid biosynthesis 37 . Tolerance involves a broad range of stress response pathways, such as the cell wall and membrane integrity pathways (Hsp90, calcineurin, PKC), the TOR pathway that responds to and regulates protein translation, as well as pathways that bypass or alleviate the drug stress indirectly, such as membrane lipid biosynthesis and central metabolic pathways, where their contribution to tolerance remains to be understood (Fig. 1 ).

Heteroresistance to fluconazole, which has been detected in Cryptococcus neoformans 38 and C. glabrata 39 , refers to the presence of a small subpopulation, usually <1% of the total, with intrinsic antifungal resistance, which can be selected for and become the dominant population on treatment. For example, in C. neoformans , a common heteroresistance mechanism is the acquisition of aneuploid chromosomes that carry genes encoding the drug target and/or efflux pump genes 38 , 40 , although aneuploidy does not explain all instances of heteroresistance 38 , 40 (Fig. 1 ).

Biofilms are a physiological adaptation to surface attachment that enables survival in the face of antifungal drugs, via multiple mechanisms, including sequestration of the drug in the extracellular matrix material that is secreted in extracellular vesicles 28 , 41 . The physiological changes that accompany biofilm formation are transient, being lost when cells exit the biofilm state and grow as yeast. Biofilms are influenced by genetic background 29 and can exhibit increased drug resistance and drug tolerance, although the degree of overlap between these processes remains to be explored. Finally, persistence is a concept seen in bacteria treated with bactericidal drugs, where rare (>0.1% of the population for most commonly used antibiotics 42 ), metabolically quiescent cells survive by not metabolizing the cidal drug. Antifungal persistence was associated with biofilms in one study, but it has been more difficult to detect (reviewed in ref. 43 ) and its relevance remains controversial 44 .

Genetic background plays a major role in antifungal tolerance, with the degree of tolerance much higher in some clinical isolates than others, such as fluconazole tolerance in Candida albicans 35 , 36 . In addition, tolerance is more evident with fungistatic drugs like azoles, yet is seen with fungicidal drugs such as echinocandins in some species. It appears that C. auris (Box 1 ) is highly resistant to azoles and also exhibits high levels of azole tolerance 45 , 46 , 47 . In C. glabrata , mitochondria play a role in the appearance of tolerance to echinocandins 48 .

Tolerance or trailing growth is not quantified in diagnostic assays. However, tolerance can be measured via minor modifications of current susceptibility assays 35 , 36 , 49 . Several small-scale studies suggested that higher tolerance of invasive C. albicans strains contribute to treatment failures and increased patient mortality 36 , 50 , 51 . Larger clinical studies are needed to determine the degree to which tolerance plays a role in treatment failures. In addition, understanding how the complex circuitry that allows cells (or only some cells) to grow under stress conditions is an important challenge currently being explored with a range of approaches.

One major approach to studying the acquisition of resistance and tolerance is experimental evolution in the presence of inhibitory or sub-inhibitory drug concentrations. Inhibitory drug concentrations select for the rare resistant isolates, while sub-inhibitory concentrations often enable the appearance of tolerant cells 52 . The effects of variables such as the genetic background of the starting isolate and/or differences in culture conditions (in vitro and in animal models) can be evaluated by their effect on the rate of resistance mutation appearance. The evolved progeny can be analyzed using selective screens that either sequence only specific genes known to be involved in resistance (e.g., direct targets of azoles ( ERG11/CYP51A ) or echinocandins ( FKS1/FKS2 ), or that use genome-wide sequencing to identify potential new resistance and tolerance mechanisms by comparing them to the progenitor strain sequence 53 , 54 , 55 ). Mutations that affect levels of drug transporters, can also be found in highly resistant isolates 55 and mutations in genes that affect stress response pathways are expected in tolerant isolates.

A second complementary approach is either to collect time series of isolates from single patients during a course of antifungal therapy, or to collect single isolates from large sets of patients 47 , 56 . Time series can be analyzed similarly to experimental evolution experiments and identified mutations can be correlated with clinical data, including changes in the application of antifungal therapies. Evaluating isolates for mutations known to confer drug resistance can establish the prevalence of specific mutations, however unstable changes such as aneuploidy, heteroresistance and physiological adaptations such as cell wall compensation changes can be missed unless drug selection is maintained 40 , 57 . Examining how these mutations have spread through the population by mapping them to a phylogeny of the isolates, can determine the level of stratification and number of independent resistance emergence of events 47 .

Another caution in all such studies is that when only a single isolate from a sample is examined it is likely that only the most frequent genotype will be identified. A more comprehensive view comes from analysis of many isolates from the same patient sample and repeated sampling over time 58 , 59 . This can reveal the variation within a host and the frequency with which mutations are maintained or lost, such as when the drug treatment is altered. Because the frequency of cells that carry a mutation conferring drug resistance may change over time, such studies can evaluate the degree of variation between the isolates collected from a single patient sample.

Genome-wide sequencing of large sample sizes allows the evaluation of mutation frequencies that correlate with resistance. Examining the sequence of genes linked to resistance in clinical isolates that exhibit resistance requires an understanding of how the mutation may affect cellular properties linked to drug sensitivity. Identifying variants systematically associated with resistance or tolerance, such as by genome-wide association studies, has the potential to identify new mechanisms of resistance. Genome sequencing can also detect correlations across studies, between the types of genome changes that arise in vitro versus those that arise in clinical samples 55 , which can strengthen confidence in the clinical relevance of those gene alleles, copy number variations, or aneuploidies.

Evaluating the genes essential for growth in the presence of drugs can identify new mechanisms important for resistance. Several screens have identified mutants that cannot grow in the presence of an antifungal drug—these include screens of large-scale gene deletions 60 , conditionally repressed strains 61 and in vivo transposon libraries 62 . Recent studies have sought to infer gene essentiality comprehensively, under any condition or in the presence of drug, by combining data from in vivo screening, genetic interactions, and gene expression using machine learning 62 or neural network algorithms 61 . Carrying out such experiments with large-scale mutant collections can provide a comprehensive catalogue of resistance mutations and an estimate of the rate at which resistance arises.

Once candidate mutations are identified, either via evolution in vitro, in animal models, or in clinical isolates, reverse genetic functional tests can introduce the specific change into a sensitive isolate and/or correct the change in the resistant isolate, and then analyze the relevant drug responses 55 . For copy number and aneuploid mutants, deleting or overexpressing those genes hypothesized to be causative can support or refute the hypothesis.

Multi-disciplinary approaches are needed to underpin development of clinical strategies to mitigate antifungal resistance. These include using experimental evolution in vitro and in more clinically relevant infection models to study ex vivo micro-evolution in serial clinical isolates from relevant infection sites. These studies would be further enhanced by incorporating other factors contributing to clinical failure such as drug exposure and treatment response biomarkers.

Box 1 The urgent threat of Candida auris drug resistance

Candida auris represents a major threat to global health as resistance to multiple classes of antifungal drugs is common. The pathogen has a unique ability to colonize human skin and mucosa and persist on surfaces in hospital and nursing home environments, including hands of healthcare workers, bed rails, medical equipment, and other surfaces, causing difficult-to-eradicate outbreaks 74 . Whilst a rare cause of candidaemia in most centres in Europe and the USA, a much higher prevalence has been reported from other parts of the world. For example, C. auris was reported as the leading cause of candidaemia (40%) at a North Indian tertiary hospital ICU, and accounted for 14% of all candidaemias reported in South Africa in 2016-2017, and 38% of all candidaemias at a single centre in Kenya in 2010–2016 (refs. 128 , 129 ). Risk factors for C. auris candidaemia include prolonged hospital and ICU stay, prior antifungal treatment, older age and having a central venous catheter in situ 130 , 131 , 132 .

Most C. auris isolates are resistant to azoles, roughly half also display lower sensitivity to the polyene amphotericin B 76 , and 41% of isolates studied are multidrug resistant. Resistance or tolerance to echinocandins, the preferred treatment, can emerge on treatment, and pan-resistance to all three drug classes has been reported. C. auris is genetically diverged from more commonly observed Candida species, and the closest related species in the Candida haemulonii complex also display high rates of resistance. Studies of the genetic basis of resistance have identified high frequency mutations in drug targets ( ERG11 and FKS1 ) as well as in the Tac1B and Mrr1 transcription factors 16 that control drug transporter expression that increase resistance to azoles and echinocandins, which vary in type and frequency between the four major genetic clades of C. auris 47 , 48 . By contrast, the mechanism of amphotericin resistance is largely unknown, aside from a report linking unusually high resistance levels to a loss of function mutation in the ERG6 protein involved in ergosterol biosynthesis 129 . C. auris is also highly adaptive to a wide variety of stressors including drug pressure and displays genetic diversity (manifest by a variety of karyotypes) and ability to develop drug tolerance during therapy, which pave the way for development of resistance 132 , 133 . New drugs such as those in late-stage clinical development that represent novel targets are needed to thwart this daunting multidrug-resistant pathogen.

Clinical consequences of antifungal resistance

Aspergillus , Cryptococcus and Candida spp. are the dominant human fungal pathogens globally, causing invasive infections of the lung, brain, and bloodstream, respectively. Repeated use of antifungals in at-risk groups, empiric, or targeted therapy of mucosal or invasive fungal infections, as well as the widespread use of azoles in agriculture, have altered the landscape of fungal species displaying resistance to one or more classes of antifungals. Most concerning is the triazole-resistant mould A. fumigatus and multidrug-resistant yeast species such as C. glabrata and C. auris 63 . Antifungal resistance threatens the limited antifungal armamentarium and affects clinical outcomes by delaying mycological clearance, and increasing breakthrough infections, relapse, and excess mortality. Intrinsic or acquired antifungal resistance are factors contributing to clinical failure in human infection (Fig. 2 ). Resistance is also potentiated by a number of factors such as: host immunosuppression (resulting in persistence or delayed clearance of infection); suboptimal antifungal pharmacokinetics (due to low oral bioavailability, lack of therapeutic drug monitoring, poor long-term treatment adherence together with inadequate antifungal drug dose, duration and/or penetration to the site of infection); and lack of source control with fungal persistence in difficult-to-reach niches such as deep-seated abscesses and device-associated biofilms 64 , 65 , 66 (Fig. 2 ).

All of the factors contributing to clinical failure in invasive fungal infection are also drivers of antifungal resistance.

Aspergillus spp. triazole resistance has been described in both environmental and clinical isolates from patients with pulmonary aspergillosis. Agricultural fungicides and long-term triazole treatment in individuals with chronic lung disease select for triazole resistance. Resistance prevalence varies by geographic region and patient population, with reported ranges between 1 and 10%, with some ICU cohorts from the Netherlands having >25% resistant isolates 67 , 68 . A 2011–2015 Dutch retrospective cohort study on cultured Aspergillus isolates from ICU and non-ICU patients reported a 19% frequency of azole resistance, with higher 6-week mortality for triazole-resistant invasive aspergillosis compared to triazole-susceptible infection treated with the first-line agent voriconazole 69 . European guidelines advocate using liposomal amphotericin B or voriconazole-echinocandin combination therapy where rates of triazole resistance exceed 10% 70 : however, routine surveillance is hampered by challenges in obtaining respiratory specimens from vulnerable patient groups and limited access to phenotypic (MIC) or genotypic ( Cyp51A ) azole susceptibility testing in hospital laboratories. Increasing exposure, diagnostic dilemmas and resultant azole exposure for aspergillosis associated with influenza and COVID-19 infections represent further drivers for resistance emergence 71 and call for adequately powered trials on the efficacy of combination therapy against invasive aspergillosis in a broad range of patient populations.

In HIV-associated cryptococcal meningitis, intrinsic heteroresistance to fluconazole in clinical isolates of C. neoformans and Cryptococcus gattii is associated with reduced fungal clearance and relapse due to secondary fluconazole resistance, even when used at the high currently recommended doses of 1200 mg/d 72 , 73 . Combination therapy of fluconazole with flucytosine eliminates the emergence of resistant subpopulations, improving fungal clearance compared to fluconazole alone 40 .

The predominant cause of mucosal and invasive candidiasis, C. albicans , is intrinsically sensitive to antifungals: however, acquired resistance can evolve with prolonged or repeated exposures to antifungals (e.g., recurrent oral, oesophageal, or vulvovaginal candidiasis). Due to factors described in Fig. 2 , a robust correlation between fluconazole MIC and clinical success in candidiasis is challenging to establish. A review of 1295 patient-episode-isolate events (692 mucosal and 603 invasive candidiasis) from 12 published clinical studies demonstrated an overall success rate of 85% for those episodes in which the fluconazole MIC was ≤8 μg/ml (sensitive), 67% for with MIC 16 to 32 μg/ml (sensitive, dose-dependent), and 42% for with resistant isolates (MIC ≥ 64 μg/ml) 74 . C. glabrata is another prominent cause of mucosal and bloodstream infections. This species has intrinsic heteroresistance to azoles and evolves stable resistance to both azoles and echinocandins following drug exposure, generating MDR isolates refractory to conventional therapy 75 . The SENTRY Antifungal Surveillance Programme reported an increase in worldwide prevalence of fluconazole-resistant C. glabrata from 8.6% to 10.1% from 1997–2014 and echinocandin resistance ranging between 1.7−3.5%; of concern, 5.5–7.6% C. glabrata isolates were resistant to both echinocandins and azoles 76 . In some tertiary care centres in the US, echinocandin resistance exceeds 13%, with elevated echinocandin MICs and the presence of FKS mutations predicted by prior echinocandin exposure and associated with clinical failure and 30-day mortality 77 . The gastrointestinal tract is largely considered the main reservoir for selection of drug resistant C. glabrata . These major fungal pathogens are now joined by C. auris as a major AMR concern. This is a newly emerged fungal pathogen classified as an urgent global threat by public health agencies due to its high transmissibility and multidrug resistance to azoles, polyenes, and sometimes echinocandins 78 , 79 , 80 (Box 1 ).

Translational pipeline and strategies to reduce and mitigate antifungal resistance

Stewardship.

Historically, antifungal drugs have been used in many patients without fungal infections through prophylactic and empiric treatment strategies. This problem is exacerbated by the poor sensitivity of traditional culture-based diagnostics, and the potentially fatal consequences of treatment delay in vulnerable patient groups, such as those with haematological malignancies 64 , 65 . Such broad use has inevitably increased selection for secondary drug resistance, and breakthrough infections by resistant species. For fungi, notably Candida spp. that can be transmitted from patient to patient, population level resistance may also rise and spread 81 , 82 . Further development and wider adoption of stewardship programmes is needed to ensure that prescribing follows evidence-based guidelines, and future research may be guided by the identification of biomarkers of drug resistance 83 , 84 . A body of literature attests to the fact that stewardship programmes can reduce inappropriate prescribing, and thus reduce selective pressure, without adversely affecting clinical outcomes – although such studies may often be insufficiently powered to detect changes in clinical endpoints compared with changes in drug use or expenditure 85 , 86 . Reporting systems and target setting have been used to monitor and promote best prescribing practice for antibacterials and could be adapted for antifungals to improve infection control including improving hospital hygiene, contact precautions based on screening for patients colonised with drug resistant organisms, and interventions to restrict the overuse antimicrobials 87 .

Antifungals that are currently in phase 2 or 3 clinical trials for the treatment or prophylaxis of fungal infections. The antifungal names as well as other identifiers are provided, along with the clinical trial number and phase, and the types of fungal infections for enrolment. Information was obtained from ClinicalTrials.gov, a database of publicly and privately funded clinical studies (accessed June 27, 2022).

Improved diagnostics

Fortunately, advances in diagnostics have enabled a shift towards more targeted pre-emptive treatment. PCR and immunoassay-based diagnostics for fungal invasive disease have become the mainstay for most well-resourced clinical diagnostics laboratories, and can deliver on-site results in under 12 h either at point of care or from minimally processed clinical samples 88 , 89 , 90 , with comparable or improved sensitivity compared with culture 91 , 92 . Pan-fungal β-D glucan assays are widely used to screen for fungal infections from clinical specimens, alongside more species-specific diagnostics such as the Aspergillus Platelia assay (Galactomannan; Bio-Rad) and several PCR-based assays. Unfortunately, these assays are often only available in reference or specialist centres, which can extend turnaround times leading to delayed treatment. Greater diagnostic mycology laboratory capacity is needed, as well as near-bedside tests such as the cryptococcal and Aspergillus lateral flow assays 93 , 94 .

Few current commercial assays can specifically identify intrinsically resistant species or detect strains that have acquired resistance 95 . Thus, MIC determination following culture continues to be the gold standard for resistance detection, although this time-consuming diagnosis is usually obtained too late to influence clinical outcome 96 .

A recently described pyrosequencing-based diagnostic directly screens respiratory samples for mutations associated with azole resistance in CYP51A of A. fumigatus . This assay has the advantage of rapidly detecting resistance even where culturing has not been possible 96 , allowing a rapid switching of therapy, when a signal is detected. However, because only 50% of azole-resistant clinical isolates have SNPs in CYP51A , the negative predictive value of this test is nominal.

The implementation of next-generation sequencing technologies in fungal diagnostics has the potential to provide further diagnostic granularity and to enable the detection and differentiation of multiple fungal species from a single sample. DNA metabarcoding using genomic targets such as ITS1 (ref. 95 ) can identify atypical pathogens within 12 h of acquiring a sample. While the cost and technical expertise required for metabarcoding diagnostics is not prohibitive for labs with molecular diagnostic experience, several technical hurdles remain to be overcome. These include identifying short genomic targets with the diagnostic potential to distinguish sub-species of pathogenic fungi, and the availability of DNA databases with suitable diversity accurate curation.

Metagenomic diagnostics, which involve sequencing all DNA from a sample without the need for amplification, is revolutionising resistance detection for tuberculosis 97 . However, implementing metagenomic diagnostics for invasive fungal diseases is currently limited by the increased cost of sequencing large fungal genomes, low coverage of the fungal genome that may limit precision resistance diagnostic assessments to be made, and by the degree to which we understand the molecular mechanisms that contribute to antifungal resistance.

Further work, and strategic trials are needed to develop and integrate new molecular diagnostics that include detection of resistance into novel management pathways. Such pathways could enable rapid targeted therapy and improved clinical outcomes for patients with fungal infections, as well as the safe discontinuation of antifungal treatment for patients without evidence of fungal infections. In resource-limited settings, laboratory mycology based on low-cost culture-based assays and near-bedside tests is paramount.

New antifungal drugs

Several new antifungals are currently in either pre-clinical or clinical development (Fig. 3 ). Some new agents are within established drug classes that may offer advantages to currently available agents. These include: (1) rezafungin, an echinocandin with a long half-life that may allow for less frequent intravenous administration; (2) encochleated amphotericin B, which is administered orally; (3) oteseconazole, a tetrazole whose structure may be more specific for fungal lanosterol 14α-demethylase than human cytochrome P450 enzymes, thus leading to fewer drug-drug interactions, and (4) opelconazole, a triazole specifically designed for inhaled delivery 98 , 99 , 100 .

There are also several therapeutic candidates in development that represent new antifungal classes with novel mechanisms of action. Ibrexafungerp, which received U.S. FDA approval in 2021, is the first member of the triterpenoid class, which, like echinocandins, inhibits the synthesis of 1,3-β-D-glucans, and can be administered orally 98 , 99 , 100 . Two other candidates that are currently in clinical trials are olorofim and fosmanogepix. Olorofim is the first member of a novel class of antifungals, the orotomides, which target fungal pyrimidine synthesis through inhibition of the enzyme dihydroorotate dehydrogenase, thus limiting the formation of uridine-5'-monophosphate (UMP) a key precursor of DNA and RNA synthesis 101 . Olorofim is unique in that it has activity against many pathogenic moulds, including those that have reduced susceptibility to other antifungals (e.g., Scedosporium spp., Microascus/Scopulariopsis ) or are pan-resistant (e.g., Lomentospora prolificans ). However, olorofim lacks activity against yeasts, as well as the Mucorales. Manogepix is the active component of fosmanogepix, a prodrug that is rapidly converted to the active moiety by systemic phosphatases following administration 102 . Manogepix targets glycosylphosphatidylinositol (GPI)-anchored protein maturation by inhibiting the fungal inositol acyltransferase enzyme GWT1, which is responsible for trafficking and anchoring mannoproteins to the fungal cell membrane and cell wall 103 . Manogepix has broad-spectrum activity against yeasts and moulds, including strains with acquired resistance to different antifungals, including azole-resistant A. fumigatus , Fusarium sp., Scedosporium sp., Lomentospora sp., C. glabrata and C. auris 102 , 104 . Thus, both olorofim and manogepix may offer hope against resistant pathogens. Nonetheless, it must be noted that resistance has been observed in vitro with exposure to each of these therapeutic candidates 105 , 106 , 107 .

Combination therapy

The extension of our antifungal armamentarium could open the door to combination therapy strategies analogous to those that have proven so successful in the treatment of a wide variety of bacterial, viral, and parasitic infections. The clearest rationale is to suppress the development of resistance to a single agent, especially when the genetic barrier to resistance is low, the infectious organism load high, and treatment duration long. In addition, combinations may act additively or synergistically to increase microbial killing, potentially allowing dose reductions of one or other agent if toxicity is dose-limiting. In cryptococcal meningitis, the addition of flucytosine to fluconazole has been demonstrated to prevent the selection of hetero-resistant colonies that otherwise leads to treatment failure 40 , and combinations of flucytosine with fluconazole and with amphotericin B accelerate the rate of clearance of infection and reduce mortality 108 , 109 , 110 , 111 This is an additional benefit to the original rationale for the use of this combination being therapy which was to decrease toxicity by lowering the drug dosage 108 .

Further work is needed to efficiently test combinations of existing, repurposed, and new agents against other systemic and chronic fungal infections, including candidemia, invasive aspergillosis and chronic pulmonary aspergillosis, given increasing resistance to and high attributable mortalities despite currently recommended monotherapies. Studies may be clinical and post-licensing as with cryptococcal combinations, where phase-II early fungicidal activity studies were crucial to efficiently select combinations for phase III trials 108 , 110 , 111 . Alternatively, studies may be initiated earlier in new drug development, based on careful PK-PD studies in animal models, and driven by industry or academia. Care should be taken that combinations are not used without good evidence of efficacy - especially given that some combinations maybe antagonistic, at least in vitro 112 . Within industry, the priority must be to obtain licensure, usually with use in monotherapy, although in tuberculosis and HIV there is strong precedent for the licensing of treatments in combination 113 , which provides a possible additional pathway for new drugs including any selected from the start for synergies with current agents. Development of penicillin/penicillinase inhibitor combinations for bacterial infections provides a specific example of such an approach that has proved to be of enduring value in the clinic 114 .

International collaboration across multiple sites, co-funding mechanisms, and, where possible, simplification of trial procedures and data collection, could facilitate adequately powered combination studies. The importance of this point can be exemplified by the results of a previous clinical trial of an azole-echinocandin combination for invasive aspergillosis: the trial was underpowered (70% power to detect a 60% reduction in mortality), so that although mortality was 30% lower with the combination, the benefit did not reach the conventional level of statistical significance 115 .

Policy, communications, and advocacy

Similar to other drug-resistant diseases, the scope of potential policy work for antifungal resistance is large. There are many diverse systems that impact the development and proliferation of antifungal-resistant pathogens, including agriculture, health care, surveillance, diagnostic testing, and drug development. Each of these systems, including those discussed earlier, affect resistant fungi in unique ways, as outlined in Table 1 .

This diversity presents both challenges and advantages to policy development and communication. Developing clear and convincing evidence-supported messages to encourage action in each of these systems requires significant time, effort, and relationship-building. However, this also represents a great opportunity for action. There are numerous pathways by which effective policy can make an impact, even if there is not universal concurrence; the development of new drugs will reduce the burden of disease, even if surveillance efforts remain underfunded. As a result, effective communications employ strategies that appeal to disparate groups and capitalize on existing communication and policy channels targeted toward these groups. For example, the US Environmental Protection Agency recently added a list of cleaning products effective against C. auris , adding to existing lists of cleaning products designed to reduce healthcare-acquired infections 116 .

Fungal disease is not typically considered a top priority when considering funding, research, and health policy—indeed typically only 3% of infectious disease research budgets support medical mycology 3 . This may change as more data are collected demonstrating the burden of these diseases. However, encouraging the inclusion of fungi in high-profile issues may be critical to spreading awareness. For example, antibiotic resistance is an issue of concern, and many policymakers may be more likely to consider the issue of antifungal resistance when the issues are packaged together, as in the US Centers for Disease Control and Prevention’s (CDC) ‘Antibiotic Resistance Threats in the United States’ report 117 .

For example, the US CDC included C. auris as an “urgent threat” and Aspergillus fumigatus on the “watch list” of antibiotic-resistant threats. This led to these pathogens inclusion in policy groups, such as the Presidential Advisory Council for Combating Antibiotic-Resistant Bacteria (despite being fungal pathogens). In turn, both pathogens were included in the Antibiotic-Resistant Lab Network and received dedicated funding because of their classification. While these pathogens might not have garnered the same interest when communicated separately, they earned more awareness and resources because they were presented with other serious threats. Similar examples globally can be seen in the inclusion of fungal disease in initiatives such as the WHO’s Global Antimicrobial Resistance Surveillance System (GLASS) 118 .

Unlike other pathogens for which the resistance pathways, risk factors, epidemiology, diagnostic practices, and treatment are well-defined and well-known, fungal pathogens pose unique challenges to communicators and advocates. Recently the WHO has extended an invitation to participate in a survey to create a priority list of fungal pathogens 119 . Policymakers may understandably feel some hesitancy in implementing policies given this uncertainty. When communicating policy concerns and recommendations, acknowledging what remains unknown and focusing effort where there is convincing evidence that policy changes will improve health is critical. This is especially true where there are potentially significant costs associated with a policy, such as changes in the commercial use of azole fungicides, which is increasingly implicated in the development of resistant A spergillus 120 , 121 .

Data do not tell the whole story, however, and patient or patient advocacy groups can help target audiences humanize the impact of fungal diseases. While data are useful to understand the burden across populations and create policies to reduce that burden, the true impact of these diseases can be lost in the numbers and among competing priorities. Patient stories can help engage policymakers, the public, and researchers alike. These stories are also able to be used in many formats, as verbal testimony, written letter, or even on social media 122 .

As progress is being made within countries, we cannot forget the importance of international collaboration. Many countries do not have dedicated public health staff to address fungal disease, but nearly every country is, or will soon be, impacted in some way by antifungal-resistant pathogens as spread continues. Drug-resistant fungal infections are becoming more common across Europe 123 . C. auris has rapidly spread throughout the globe, including in many low- and middle-income countries without existing resources to combat these threats. Moreover, many cases emerging in previously naïve countries are linked to travel, including the sentinel case in the western US 124 .

Despite the caveats inherent to health communication of any variety, it is crucial to remember that prevention works. Prevention is the most cost-effective solution we have to combat resistant fungal infections, and policy and communications are key tools to improve prevention activities. Given limited funding opportunities, public health efforts supported by science are going to be a beneficial investment; it is just a matter of teaching others that they will be as well.

Conclusions

The impact of fungal disease, potentiated by drug resistant infections, has become an urgent health priority, but innovation and progress have been limited by capacity in both discovery and translational research sectors. Without enhanced visibility of mycology to all stakeholders, including funders, researchers, industry, patients, and the public, it will be difficult to incentivize the development of capacity in this area and to catalyse interdisciplinary working to encourage step changes in therapeutic and diagnostic opportunities for the treatment of fungal infections. Dispersed specialist communities can achieve greater impact and effective advocacy through global coordination and integration of their work with allied fields of public health and infection biology to make inroads into public and private sector investment.

It is clear that the value of development of new broad-spectrum therapeutic options that are already in the pipeline would be augmented by improved diagnostics and greater understanding of the conditions and mechanisms that promote resistance and tolerance.

This position paper outlines significant progress in these respects, yet the global burden of serious fungal infection remains high, and trends continue upwards. Investment now is needed to reverse these trends and to adopt an integrated One Health approach encompassing environmental, clinical, agricultural, and social perspectives that is reviewed by GAFFI 125 . Without this investment it is possible or probable that drug resistant fungal infections will increasingly compromise successful treatment of mycotic disease.

Woolhouse, M. & Farrar, J. Policy: an intergovernmental panel on antimicrobial resistance. Nature 509 , 555–557 (2014).

Article PubMed Google Scholar

O’Neill, J. Tackling Drug-Resistant Infections Globally: Final Report and Recommendations . https://amr-review.org/sites/default/files/160518_Final%20paper_with%20cover.pdf (2022).

Brown, G. D. et al. Hidden Killers: Human fungal infections. Sci. Transl. Med. https://doi.org/10.1126/scitranslmed.3004404 (2012).

Bongomin, F., Gago, S., Oladele, R. O. & Denning, D. W. Global and multi-national prevalence of fungal diseases-estimate precision. J. Fungi (Basel) 3 , 57 (2017).

Article Google Scholar

Denning, D. W. & Bromley, M. J. How to bolster the antifungal pipeline. Science 347 , 1414–1416 (2015).

Article ADS CAS PubMed Google Scholar

Fairlamb, A. H., Gow, N. A., Matthews, K. R., Waters, A. P. Drug resistance in eukaryotic microorganisms. Nat. Microbiol . https://doi.org/10.1038/nmicrobiol.2016.92 (2016).

Fisher, M. C., Hawkins, N. J., Sanglard, D. & Gurr, S. J. Worldwide emergence of resistance to antifungal drugs challenges human health and food security. Science 360 , 739–742 (2018).

Article CAS PubMed Google Scholar

CDC. Antibiotic Resistance Threats in the United States, 2019 . www.cdc.gov/DrugResistance/Biggest-Threats.html (2019).

JPIAMR. JPIAMR Strategic Research and Innovation Agenda on Antimicrobial Resistance. https://www.jpiamr.eu/wp-content/uploads/2021/04/JPIAMR_SRIA_2021.pdf (2022).

Fisher, M. C. et al. Tackling the emerging threat of antifungal resistance to human health. Nat. Rev. Microbiol https://doi.org/10.1038/s41579-022-00720-1 (2022).

Article PubMed PubMed Central Google Scholar

Wiederhold, N. Review of the novel investigational antifungal olorofim. J. Fungi 6 , 122 (2020).

Article CAS Google Scholar

Burks C., Darby A., Go´mez Londoño L., Momany M. & Brewer M. T. Azole-resistant Aspergillus fumigatus in the environment: Identifying key reservoirs and hotspots of antifungal resistance. PLoS Pathog . https://doi.org/10.1371/journal.ppat.1009711 (2021).

Castelo-Branco, D. et al. Collateral consequences of agricultural fungicides on pathogenic yeasts: a one health perspective to tackle azole resistance. Mycoses 65 , 303–311 (2022). 2022.

Shelton, J. M. G. et al. Citizen science surveillance of triazole-resistant Aspergillus fumigatus in United Kingdom residential garden soils. Appl. Environ. Microbiol . https://doi.org/10.1128/AEM.02061-21 (2022).

Stott K. E. et al. Cryptococcal meningoencephalitis: time for action. Lancet Infect. Dis . https://doi.org/10.1016/S1473-3099(20)30771-4 (2021).

van Schalkwyk E. et al. Epidemiologic shift in candidemia driven by Candida auris, South Africa, 2016–2017 Emerg. Infect. Dis . https://doi.org/10.3201/eid2509.190040 (2019).

Pfaller, M. et al. Antifungal susceptibilities of Candida, Cryptococcus neoformans and Aspergillus fumigatus from the Asia and Western Pacific region: data from the SENTRY antifungal surveillance program (2010–2012). J. Antibiot. 68 , 556–561 (2015).

Lamoth, F., Lockhart, S. R., Berkow, E. L. & Calandra, T. Changes in the epidemiological landscape of invasive candidiasis. J. Antimicrob. Chemother. 73 , i4–i13 (2018).

Arendrup, M. C. & Patterson, T. F. Multidrug-resistant Candida : epidemiology, molecular mechanisms, and treatment. J. Infect. Dis. 216 , S445–S451 (2017).

Berger, S., El Chazli, Y., Babu, A. F. & Coste, A. T. Azole resistance in Aspergillus fumigatus : a consequence of antifungal use in agriculture? Front. Microbiol. 8 , 1024 (2017).

Sacheli, R. & Hayette, M.-P. Antifungal resistance in dermatophytes: genetic considerations, clinical presentations and alternative therapies. J. Fungi 7 , 983 (2021).

Ebert, A. et al. Alarming India-wide phenomenon of antifungal resistance in dermatophytes: a multicentre study. Mycoses 63 , 717–728 (2020).

Espinel-Ingroff, A. & Turnidge, J. The role of epidemiological cutoff values (ECVs/ECOFFs) in antifungal susceptibility testing and interpretation for uncommon yeasts and moulds. Rev. Iberoam. Micol. 33 , 63–75 (2016).

Cornely, O. A. et al. Defining breakthrough invasive fungal infection-position paper of the mycoses study group education and research consortium and the European Confederation of Medical Mycology. Mycoses 62 , 716–729 (2019).

PubMed PubMed Central Google Scholar

Lee, Y., Puumala, E., Robbins, N. & Cowen, L. E. Antifungal drug resistance: molecular mechanisms in Candida albicans and beyond. Chem. Rev. 121 , 3390–3411 (2021).

Berman, J. & Krysan, D. J. Drug resistance and tolerance in fungi. Nat. Rev. Microbiol. 18 , 319–331 (2020).

Article CAS PubMed PubMed Central Google Scholar

Delarze, E. & Sanglard, D. Defining the frontiers between antifungal resistance, tolerance and the concept of persistence. Drug Resist. Updat. Rev. 23 , 12–19 (2015).

Zarnowski, R. et al. Coordination of fungal biofilm development by extracellular vesicle cargo. Nat. Commun. 12 , 6235 (2021).

Article ADS CAS PubMed PubMed Central Google Scholar

Huang, M. Y., Woolford, C. A., May, G., McManus, C. J. & Mitchell, A. P. Circuit diversification in a biofilm regulatory network. PLoS Pathog . https://doi.org/10.1371/journal.ppat.1007787 (2019).

Maisonneuve, E. & Gerdes, K. Molecular mechanisms underlying bacterial persisters. Cell 157 , 539–548 (2014).

Selmecki, A., Forche, A. & Berman, J. Aneuploidy and isochromosome formation in drug-resistant Candida albicans. Science 313 , 367–370 (2006).

Yang, F. et al. Aneuploidy enables cross-adaptation to unrelated drugs. Mol. Biol. Evol. 36 , 1768–1782 (2019).

Yang, F. et al. Aneuploidy underlies tolerance and cross-tolerance to drugs in Candida parapsilosis. Microbiol. Spectr . https://doi.org/10.1128/Spectrum.00508-21 (2021).

Hayes, B. M. E., Anderson, M. A., Traven, A., van der Weerden, N. L. & Bleackley, M. R. Activation of stress signalling pathways enhances tolerance of fungi to chemical fungicides and antifungal proteins. Cell. Mol. Life Sci. 71 , 2651–2666 (2014).

Delarze, E. et al. Identification and characterization of mediators of fluconazole tolerance in Candida albicans . Front. Microbiol . https://doi.org/10.3389/fmicb.2020.591140 (2020).

Rosenberg, A. et al. Antifungal tolerance is a subpopulation effect distinct from resistance and is associated with persistent candidemia. Nat. Commun. 9 , 2470 (2018).

Article ADS PubMed PubMed Central CAS Google Scholar

Gao, J. et al. Candida albicans gains azole resistance by altering sphingolipid composition. Nat. Commun. 9 , 4495 (2018).

Sionov, E., Chang, Y. C., Garraffo, H. M. & Kwon-Chung, K. J. Heteroresistance to fluconazole in Cryptococcus neoformans is intrinsic and associated with virulence. Antimicrob. Agents Chemother. 53 , 2804–2815 (2009).

Ben-Ami, R. et al. Heteroresistance to fluconazole is a continuously distributed phenotype among Candida glabrata clinical strains associated with in vivo persistence. mBio https://doi.org/10.1128/mBio.00655-16 (2016).

Stone, N. R. H. et al. Dynamic ploidy changes drive fluconazole resistance in human cryptococcal meningitis. J. Clin. Invest. 129 , 999–1014 (2019).

Zarnowski, R. et al. Candida albicans biofilm–induced vesicles confer drug resistance through matrix biogenesis. PLoS Biol . https://doi.org/10.1371/journal.pbio.2006872 (2018).

Salcedo-Sora, J. E. & Kell, D. B. A quantitative survey of bacterial persistence in the presence of antibiotics: towards antipersister antimicrobial discovery. Antibiotics 9 , 508 (2020).

Article CAS PubMed Central Google Scholar

Wuyts, J., Van Dijck, P. & Holtappels, M. Fungal persister cells: the basis for recalcitrant infections? PLoS Pathog . https://doi.org/10.1371/journal.ppat.1007301 (2018).

Denega, I., d’Enfert, C. & Bachellier-Bassi, S. Candida albicans biofilms are generally devoid of persister cells. Antimicrob. Agents Chemother . https://doi.org/10.1128/AAC.01979-18 (2019).

Kim, S. H. et al. Genetic analysis of Candida auris implicates Hsp90 in morphogenesis and azole tolerance and Cdr1 in azole resistance. mBio https://doi.org/10.1128/mBio.02529-18 (2019).

Bhattacharya, S., Holowka, T., Orner, E. P. & Fries, B. C. Gene duplication associated with increased fluconazole tolerance in Candida auris cells of advanced generational age. Sci. Rep . https://doi.org/10.1038/s41598-019-41513-6 (2019).

Chow, N. A. et al. Tracing the evolutionary history and global expansion of Candida auris using population genomic analyses. mBio https://doi.org/10.1128/mBio.03364-19 (2020).

Garcia-Rubio, R. et al. Multifactorial role of mitochondria in echinocandin tolerance revealed by transcriptome analysis of drug-tolerant cells. mBio https://doi.org/10.1128/mBio.01959-21 (2021).

Gerstein, A. C., Rosenberg, A., Hecht, I. & Berman, J. diskImageR: quantification of resistance and tolerance to antimicrobial drugs using disk diffusion assays. Microbiol 162 , 1059–1068 (2016).

Astvad, K. M. T., Sanglard, D., Delarze, E., Hare, R. K. & Arendrup, M. C. Implications of the EUCAST trailing phenomenon in candida tropicalis for the in vivo susceptibility in invertebrate and murine models. Antimicrob. Agents Chemother . https://doi.org/10.1128/AAC.01624-18 (2018).

Levinson, T. et al. Impact of tolerance to fluconazole on treatment response in Candida albicans bloodstream infection. Mycoses 64 , 78–85 (2021).

Gerstein, A. C. & Berman, J. Candida albicans genetic background influences mean and heterogeneity of drug responses and genome stability during evolution in fluconazole. mSphere https://doi.org/10.1128/mSphere.00480-20 (2020).

Todd, R. T. & Selmecki, A. Expandable and reversible copy number amplification drives rapid adaptation to antifungal drugs. eLife https://doi.org/10.7554/eLife.58349 (2020).

Yang, F. et al. Tunicamycin potentiates antifungal drug tolerance via aneuploidy in Candida albicans . mBio https://doi.org/10.1128/mBio.02272-21 (2021).

Rybak, J. M. et al. Mutations in TAC1B: a novel genetic determinant of clinical fluconazole resistance in Candida auris . mBio https://doi.org/10.1128/mBio.00365-20 (2020).

Ford, C. B. et al. The evolution of drug resistance in clinical isolates of Candida albicans . eLife https://doi.org/10.7554/eLife.00662 (2015).

da Silva Dantas, A. et al. Crosstalk between the Calcineurin and cell wall integrity pathways prevents chitin overexpression in Candida albicans . J. Cell Sci . https://doi.org/10.1242/jcs.258889 (2021).

Summerbell, R. C., Cooper, E., Bunn, U., Jamieson, F. & Gupta, A. K. Onychomycosis: a critical study of techniques and criteria for confirming the etiologic significance of nondermatophytes. Med Mycol. 43 , 39–59 (2005).

Jousset, A. B. et al. Concomitant carriage of KPC-producing and non-KPC-producing Klebsiella pneumoniae ST512 within a single patient. J. Antimicrob. Chemother. 75 , 2087–2092 (2020).

CAS PubMed Google Scholar

Furukawa, T. et al. The negative cofactor 2 complex is a key regulator of drug resistance in Aspergillus fumigatus . Nat. Commun. 11 , 427 (2020).

Fu, C. et al. Leveraging machine learning essentiality predictions and chemogenomic interactions to identify antifungal targets. Nat. Commun. 12 , 6497 (2021).

Segal, E. S. et al. Gene Essentiality Analyzed by In Vivo Transposon Mutagenesis and Machine Learning in a Stable Haploid Isolate of Candida albicans . mBio https://doi.org/10.1128/mBio.02048-18 (2018).

Arastehfar, A. et al. Drug-resistant fungi: an emerging challenge threatening our limited antifungal armamentarium. Antibiotics 9 , 877 (2020).

Perlin, D. S., Rautemaa-Richardson, R. & Alastruey-Izquierdo, A. The global problem of antifungal resistance: prevalence, mechanisms, and management. Lancet Infect. Dis. 17 , e383–e392 (2017).

Von Eiff, M. et al. Pulmonary aspergillosis: early diagnosis improves survival. Respiration 65 , 341–347 (1995).

Morrell, M., Fraser, V. J. & Kollef, M. H. Delaying the empiric treatment of Candida bloodstream infection until positive blood culture results are obtained: a potential risk factor for hospital mortality. Antimicrob. Agents Chemother. 49 , 3640–3645 (2005).

Lestrade, P. P. A., Meis, J. F., Melchers, W. J. G. & Verweij, P. E. Triazole resistance in Aspergillus fumigatus : recent insights and challenges for patient management. Clin. Microbiol. Infect . 7 , https://doi.org/10.1016/j.cmi.2018.11.027 (2019).

Rybak, J. M., Fortwendel, J. R. & Rogers, P. D. Emerging threat of triazole-resistant Aspergillus fumigatus . J. Antimicrob. Chemother . https://doi.org/10.1093/jac/dky517 (2019).

Lestrade, P. P. A. et al. Voriconazole resistance and mortality in invasive aspergillosis: a multicenter retrospective cohort study. Clin Infect Dis . https://doi.org/10.1093/cid/ciy859 (2019).

Ullmann, A. J. et al. Diagnosis and management of Aspergillus diseases: executive summary of the 2017 ESCMID-ECMM-ERS guideline. Clin Microbiol Infect . https://doi.org/10.1016/j.cmi.2018.01.002 (2018).

Meijer, E. F. J., Dofferhoff, A. S. M., Hoiting, O., Buil, J. B. & Meis, J. F. Azole-resistant COVID-19-associated pulmonary aspergillosis in an immunocompetent host: a case report. J. Fungi . (Basel). https://doi.org/10.3390/jof6020079 (2020).

Rajasingham, R. et al. Global burden of disease of HIV-associated cryptococcal meningitis: an updated analysis. Lancet Infect. Dis. 17 , 873–881 (2017).

Temfack, E. et al. Cryptococcal antigen in serum and cerebrospinal fluid for detecting Cryptococcal Meningitis in adults living with human immunodeficiency virus: systematic review and meta-analysis of diagnostic test accuracy studies. Clin. Infect. Dis. 72 , 1268–1278 (2021).

Pfaller, M. A., Diekema, D. J. & Sheehan, D. J. Interpretive breakpoints for fluconazole and Candida revisited: a blueprint for the future of antifungal susceptibility testing. Clin Microbiol Rev . https://doi.org/10.1128/CMR.19.2.435-447.2006 (2006).

Healey, K. R. & Perlin, D. S. Fungal resistance to echinocandins and the MDR phenomenon in Candida glabrata . J. Fungi . https://doi.org/10.3390/jof4030105 (2018).

Pfaller, M. A., Diekema, D. J., Turnidge, J. D., Castanheira, M. & Jones R. N. Twenty years of the SENTRY antifungal surveillance program: results for Candida species from 1997–2016. Open Forum. Infect. Dis . https://doi.org/10.1093/ofid/ofy358 (2019).

Alexander, B. D. et al. Increasing echinocandin resistance in Candida glabrata : clinical failure correlates with presence of FKS mutations and elevated minimum inhibitory concentrations. Clin Infect Dis . https://doi.org/10.1093/cid/cit136 (2013).

Lockhart, S. R. Candida auris and multidrug resistance: defining the new normal. Fungal. Genet. Biol . https://doi.org/10.1016/j.fgb.2019.103243 (2019).

Lockhart, S. R. et al. Simultaneous emergence of multidrug-resistant Candida auris on 3 continents confirmed by whole-genome sequencing and epidemiological analyses. Clin Infect Dis . https://doi.org/10.1093/cid/ciw691 (2017).

Eyre, D. W. et al. A Candida auris outbreak and its control in an intensive care setting. N. Engl. J. Med . https://doi.org/10.1056/NEJMoa1714373 (2018).

Ksiezopolska, E. & Gabaldón, T. Evolutionary emergence of drug resistance in Candida opportunistic pathogens. Genes 9 , 461 (2018).

Article PubMed Central CAS Google Scholar

Arastehfar, A. et al. Clonal Candidemia outbreak by Candida parapsilosis Carrying Y132F in Turkey: evolution of a persisting challenge. Front. Cell. Infect. Microbiol . https://doi.org/10.3389/fcimb.2021.676177 (2021).

Honarvar, B., Bagheri Lankarani, K., Taghavi, M., Vahedi, G. & Mortaz, E. Biomarker-guided antifungal stewardship policies for patients with invasive candidiasis. Curr. Med Mycol. 4 , 37–44 (2018).

de Cesare, G. B., Hafez, A., Stead, D., Llorens, C. & Munro, C. Biomarkers of caspofungin resistance in Candida albicans isolates: a proteomic approach. Virulence 13 , 1005–1018 (2020).

Bienvenu, A. L. et al. A systematic review of interventions and performance measures for antifungal stewardship programmes. J. Antimicrob. Chemother. 73 , 297–305 (2018).

Hart E., Nguyen M., Allen M., Clark C. M. & Jacobs D. M. A systematic review of the impact of antifungal stewardship interventions in the United States. Ann. Clin. Microbiol. Antimicrob . https://doi.org/10.1186/s12941-019-0323-z (2019).

Sharland, M. et al. Encouraging AWaRe-ness and discouraging inappropriate antibiotic use-the new 2019 Essential Medicines List becomes a global antibiotic stewardship tool. Lancet Infect. Dis. 19 , 1278–1280 (2019).

Sanguinetti, M. et al. Diagnosis and treatment of invasive fungal infections: looking ahead. J. Antimicrob. Chemother . https://doi.org/10.1093/jac/dkz041 (2019).

Terrero-Salcedo, D. & Powers-Fletcher, M. V. Updates in laboratory diagnostics for invasive fungal infections. J. Clin. Microbiol . https://doi.org/10.1128/JCM.01487-19 (2020).

Wickes, B. L. & Wiederhold, N. P. Molecular diagnostics in medical mycology. Nat. Commun. 9 , 5135 (2018).

Monday, L. M., Parraga Acosta, T. & Alangaden, G. T2Candida for the diagnosis and management of invasive Candida infections. J. Fungi . https://doi.org/10.3390/jof7030178 . (2021).

Vergidis, P. et al. High-volume culture and quantitative real-time PCR for the detection of Aspergillus in sputum. Clin. Microbiol Infect. 26 , 935–940 (2019).

Boulware, D. R. et al. Multisite validation of cryptococcal antigen lateral flow assay and quantification by laser thermal contrast. Emerg. Infect. Dis. 20 , 45–53 (2014).

Heldt, S. & Hoenigl, M. Lateral flow assays for the diagnosis of invasive Aspergillosis : current status. Curr. Fungal Infect. Rep. 11 , 45–51 (2017).

Kidd, S. E., Chen, S. C., Meyer, W. & Halliday, C. L. A new age in molecular diagnostics for invasive fungal disease: are we ready. Front Microbiol 10 , 2903 (2020).

Novak-Frazer, L. et al. Deciphering Aspergillus fumigatus cyp51A-mediated triazole resistance by pyrosequencing of respiratory specimens. J. Antimicrob. Chemother. 75 , 3501–3509 (2020).

Yu, G., Zhao, W., Shen, Y., Zhu, P. & Zheng, H. Metagenomic next generation sequencing for the diagnosis of tuberculosis meningitis: a systematic review and meta-analysis. PLoS One . https://doi.org/10.1371/journal.pone.0243161 (2020).

Rauseo, A. M., Coler-Reilly, A., Larson, L. & Spec, A. Hope on the horizon: novel fungal treatments in development. Open Forum. Infect. Dis . https://doi.org/10.1093/ofid/ofaa016 (2020).

Hoenigl, M. et al. The antifungal pipeline: Fosmanogepix, Ibrexafungerp, Olorofim, Opelconazole, and Rezafungin. Drugs 81 , 1703–1729 (2021).

Van Daele, R. et al. Antifungal drugs: what brings the future? Med Mycol . https://doi.org/10.1093/mmy/myz012 (2019).

Oliver, J. D. et al. F901318 represents a novel class of antifungal drug that inhibits dihydroorotate dehydrogenase. Proc. Natl Acad. Sci. USA 113 , 12809–12814 (2016).

Shaw, K. J. & Ibrahim, A. S. Fosmanogepix: a review of the first-in-class broad spectrum agent for the treatment of invasive fungal infections. J. Fungi . https://doi.org/10.3390/jof6040239 (2020).

Watanabe, N. A. et al. E1210, a new broad-spectrum antifungal, suppresses Candida albicans hyphal growth through inhibition of glycosylphosphatidylinositol biosynthesis. Antimicrob. Agents Chemother. 56 , 960–971 (2012).

Miyazaki, M. et al. In vitro activity of E1210, a novel antifungal, against clinically important yeasts and molds. Antimicrob. Agents Chemother. 55 , 4652–4658 (2011).

Kapoor, M., Moloney, M., Soltow, Q. A., Pillar, C. M. & Shaw, K. J. Evaluation of Resistance Development to the Gwt1 Inhibitor Manogepix (APX001A) in Candida Species. Antimicrob Agents Chemother . https://doi.org/10.1128/AAC.01387-19 (2019).

Liston, S. D., Whitesell, L., Kapoor, M., Shaw, K. J. & Cowen, L. E. Enhanced efflux pump expression in Candida mutants results in decreased manogepix susceptibility. Antimicrob Agents Chemother . https://doi.org/10.1128/AAC.00261-20 (2020).

Buil, J. B. et al. Resistance profiling of Aspergillus fumigatus to olorofim indicates absence of intrinsic resistance and unveils the molecular mechanisms of acquired olorofim resistance. Emerg. Microbes Infect. 11 , 703–714 (2022).

Nussbaum, J. C. et al. Combination flucytosine and high-dose fluconazole compared with fluconazole monotherapy for the treatment of cryptococcal meningitis: a randomized trial in Malawi. Clin. Infect. Dis. 50 , 338–344 (2010).

Jackson, A. T. et al. A phase II randomized controlled trial adding oral flucytosine to high-dose fluconazole, with short-course amphotericin B, for cryptococcal meningitis. AIDS 26 , 1363–1370 (2012).

Molloy, S. F. et al. Antifungal combinations for treatment of cryptococcal meningitis in Africa. N. Engl. J. Med . 378 , 1004–1017 (2018). ACTA Trial Study Team.

Jarvis, J. N. et al. Single-dose liposomal amphotericin B treatment for cryptococcal meningitis. N. Engl. J. Med . 386 , 1109–1120 (2022).

Van Rhijn, R., Hemmings, S., Valero, C., Amich, J. & Bromley, M. J. Olorofim and the azoles are antagonistic in A. fumigatus and functional genomic screens reveal mechanisms of cross resistance. Preprint at https://www.biorxiv.org/content/10.1101/2021.11.18.469075v1 (2021).

TB Alliance. FDA Approval of BPaL Regimen an Important Breakthrough in TB Control. https://www.kncvtbc.org/en/2019/08/14/fda-approval-of-bpal-regimen-an-important-breakthrough-in-tb-control/ (2022)

Tooke, C. L. et al. β-Lactamases and β-Lactamase Inhibitors in the 21st century. J. Mol. Biol. 431 , 3472–3500 (2019).

Marr, K. A. et al. Combination antifungal therapy for invasive aspergillosis: a randomized trial. Ann. Intern. Med. 162 , 81–89 (2015).

United States Environmental Protection Agency. List P: Antimicrobial Products Registered with EPA for Claims Against Candida Auris. https://www.epa.gov/pesticide-registration/list-p-antimicrobial-products-registered-epa-claims-against-candida-auris#:~:text=List%20P%3A%20Antimicrobial%20Products%20Registered%20with%20EPA%20for,%20%201%20%206%20more%20rows%20 (2022).

United States Centre for Disease Control. 2019 AR threats Report . https://www.cdc.gov/drugresistance/pdf/threats-report/2019-ar-threats-report-508.pdf (2022).

World Health Organisation. Global Antimicrobial Resistance use and Surveillance System (GLASS). https://www.who.int/initiatives/glass#:~:text=GLASS-FUNGI%20focuses%20on%20the%20surveillance%20of%20invasive%20fungal,men%20with%20urethral%20discharge%20and%20suspected%20urogenital%20infections (2022).

World Health Organisation. First WHO Fungal Priority Pathogens List . https://www.who.int/news-room/articles-detail/invitation-to-participate-in-survey-to-establish-the-first-who-fungal-priority-pathogens-list-(fppl) (2022).

Toda, M., Beer, K. D., Kuivila, K. M., Chiller, T. M. & Jackson, B. R. Trends in agricultural triazole fungicide use in the United States, 1992–2016 and possible implications for antifungal-resistant fungi in human disease. Environ. Health Perspect . https://doi.org/10.1289/EHP7484 (2021).

Beer, K. D. et al. Multidrug-resistant Aspergillus fumigatus carrying mutations linked to environmental fungicide exposure—three states, 2010–2017. Morb. Mortal. Wkly Rep. 67 , 1064–1067 (2018).

CDC. Center for Disease Control: Personal Stories of Fungal Disease . https://www.cdc.gov/fungal/personalstories/index.html#Cryptococcus (2022).

Snelders, E. et al. Emergence of azole resistance in Aspergillus fumigatus and spread of a single resistance mechanism. PLoS Med . https://doi.org/10.1371/journal.pmed.0050219 (2008).

Woodworth, M. H. et al. Sentinel case of Candida auris in the Western United states following prolonged occult colonization in a returned traveller from India. Micro. Drug Resist. 25 , 677–680 (2019).

Gaffi. Global Action for Fungal Infections . https://gaffi.org/ (2022).

Loyse, A. et al. Leave no one behind: response to new evidence and guidelines for the management of cryptococcal meningitis in low-income and middle-income countries. Lancet Infect. Dis. https://doi.org/10.1016/S1473-3099(18)30493-6 (2019).

Unitaid. A plan to Slash HIV Deaths. https://unitaid.org/advanced-hiv-disease/#en (2022).

Shastri, P. S. et al. Candida auris candidaemia in an intensive care unit—prospective observational study to evaluate epidemiology, risk factors, and outcome. J. Crit. Care . https://doi.org/10.1016/j.jcrc.2020.01.004 (2020).

Adam, R. D. et al. Analysis of Candida auris fungemia at a single facility in Kenya. Int J. Infect. Dis. 85 , 182–187 (2019).

Pandya, N. et al. International multicentre study of Candida auris Infections. J. Fungi (Basel) 7 , 878 (2021).

Shor, E. & Perlin, D. S. Coping with stress and the emergence of multidrug resistance in fungi. PLoS Pathog . https://doi.org/10.1371/journal.ppat.1004668 (2015).

Li, J. et al. Novel ERG11 and TAC1b mutations associated with azole resistance in Candida auris. Antimicrob. Agents Chemother . https://doi.org/10.1128/AAC.02663-20 (2021).

Rybak, J. M. et al. In vivo emergence of high-level resistance during treatment reveals the first identified mechanism of amphotericin B resistance in Candida auris. Clin. Microbiol Infect. https://doi.org/10.1016/j.cmi.2021.11.024 (2021).

Download references

Acknowledgements

N.A.R.G. acknowledges support of Wellcome Senior Investigator (101873/Z/13/Z), and Wellcome Collaborative awards (200208/A/15/Z, 215599/Z/19/Z) and the MRC Centre for Medical Mycology [MR/N006364/2]. J.B. acknowledges funding from the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement No 951475, ERC SYN FungalTolerance). D.S.P. is supported by NIH grant AI109025. T.B. is the recipient of the Emerging leaders award in Antimicrobial resistance from the Medical research foundation (MRF-160-0009-ELP-BICA-C0802). N.W. has received funding (to UT Health San Antonio) from Astellas, bioMerieux, F2G, Maxwell Biosciences, Sfunga, and the U.S. NIAID/NIH. M.B. receives Wellcome Trust support from collaborative and biomedical resource grants (219551/Z/19/Z, 208396/Z/17/Z), the NIHR Manchester Biomedical Research Centre and F2G. C.A.C. is a fellow in the CIFAR Fungal Kingdom program.

Author information

Authors and affiliations.

MRC Centre for Medical Mycology, School of Biosciences, University of Exeter, Geoffrey Pope Building, Exeter, EX4 4QD, UK

Neil A. R. Gow & Thomas S. Harrison

Medical Research Council, Polaris House, Swindon, SN2 1FL, UK

Carolyn Johnson

Shmunis School of Biomedical and Cancer Research, George S. Wise Faculty of Life Sciences, Tel Aviv University, 418 Britannia Building, Ramat Aviv, 69978, Israel

Judith Berman

Microbiology Institute, University Hospital Lausanne, rue du Bugnon 48, 1011, Lausanne, Switzerland

Alix T. Coste

(CAC) Infectious Disease and Microbiome Program, Broad Institute of MIT and Harvard, Cambridge, MA, 02142, USA

Christina A. Cuomo

Center for Discovery and Innovation, Hackensack Meridian health, Nutley, NJ, 07110, USA

David S. Perlin

Institute of Infection and Immunity, St George’s University of London, London, SW17 0RE, UK

Tihana Bicanic & Thomas S. Harrison

Clinical Academic Group in Infection, St George’s University Hospitals NHS Foundation Trust, London, SW17 0QT, UK

Department of Pathology and Laboratory Medicine, University of Texas Health Science Center at San Antonio, San Antonio, TX, 78229, USA

Nathan Wiederhold

Manchester Fungal Infection Group, Division of Evolution, Infection, and Genomics, Faculty of Biology, Medicine and Health, University of Manchester, CTF Building, 46 Grafton Street, Manchester, M13 9NT, UK

Mike Bromley

Center for Disease Control and Prevention Mycotic Disease Branch 1600 Clifton Rd, MSC-09, Atlanta, 30333, GA, USA

Tom Chiller & Keegan Edgar

You can also search for this author in PubMed Google Scholar

Contributions

All authors contributed equally to the writing and revision of the manuscript. The order of authors reflects the order of the contributions in the review. A consortium of authors contributed to the Exeter-MRC workshop on fungal AMR on May 4th 2021 that provided analysis for this review. Attendees of the workshop are acknowledged in Supplementary Table 1 .

Corresponding authors

Correspondence to Neil A. R. Gow or Carolyn Johnson .

Ethics declarations

Competing interests.

The authors declare no competing interests.

Peer review

Peer review information.

Nature Communications thanks Florent Morio and the other, anonymous, reviewers for their contribution to the peer review of this work.

Additional information

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Supplementary information

Supplementary information, rights and permissions.

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/ .

Reprints and permissions

About this article

Cite this article.

Gow, N.A.R., Johnson, C., Berman, J. et al. The importance of antimicrobial resistance in medical mycology. Nat Commun 13 , 5352 (2022). https://doi.org/10.1038/s41467-022-32249-5

Download citation

Received : 31 March 2022

Accepted : 22 July 2022

Published : 12 September 2022

DOI : https://doi.org/10.1038/s41467-022-32249-5

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

This article is cited by

Synergistic activity of gold nanoparticles with amphotericin b on persister cells of candida tropicalis biofilms.

M. A. Dasilva
K. F. Crespo Andrada
María Gabriela Paraje

Journal of Nanobiotechnology (2024)

Vitamin B5 metabolism is essential for vacuolar and mitochondrial functions and drug detoxification in fungi

Jae-Yeon Choi
Shalev Gihaz
Choukri Ben Mamoun

Communications Biology (2024)

Nanotechnology’s frontier in combatting infectious and inflammatory diseases: prevention and treatment

Yujing Huang
Xiaohan Guo

Signal Transduction and Targeted Therapy (2024)

In vivo femtosecond laser nanosurgery of the cell wall enabling patch-clamp measurements on filamentous fungi

Tanja Pajić
Katarina Stevanović
Mihailo D. Rabasović

Microsystems & Nanoengineering (2024)

Synthesis and molecular docking simulations of novel azepines based on quinazolinone moiety as prospective antimicrobial and antitumor hedgehog signaling inhibitors

Ahmed A. Noser
A. A. El-Barbary
Mohamed shahien

Scientific Reports (2024)

By submitting a comment you agree to abide by our Terms and Community Guidelines . If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.

Quick links

Explore articles by subject
Guide to authors
Editorial policies

Sign up for the Nature Briefing: Microbiology newsletter — what matters in microbiology research, free to your inbox weekly.

Buy 1 supplement receive 3 for free. Use Code SUMMER24 -->

Item added to your cart

What is mycology a rapidly growing field on the study of mushrooms, what is mycology mycology is the study of mushrooms, but there is so much more to know about this fascinating and extremely important field.

Mycology is the study of mushrooms, and it is a rapidly growing field with many intersections with other topics. From the classification and evolution of fungal species, to the use of magic mushrooms, to using mushroom mycelium for packaging material the field of mycology is diversifying. Historically mycology has primarily focused on fungal pathogens found in the agricultural world. Fungal pathogens have huge impacts on the total productivity of agriculture. Fungi can account for over 20% of total crop loss throughout the world. Many of the mycology programs in universities focus on identifying fungi or reducing fungal pathogens on agricultural crops. So what is mycology beyond the world of identification and pathology?

What is mycology in the 21st century: Mycology is the study of mushrooms, but we also have a new definition

Mycology is turning into a new science as cultivation and the use of fungi to support humans is increasing. What is mycology in the radical sense? Mycology is the study of mushrooms and the planet and methods of allying with fungi to improve ourselves, the planet, and our relationships. Mycology is shifting and hopefully will continue to shift towards the positive aspects of mushrooms. What are some of the positive aspects of mushrooms, let's explore new avenues for mycology.

what is mycology? Mycology is the study of mushrooms

Artwork by Martin Bridge. thebridgebrothers.com

What is Mycology: Mushroom Cultivation

Mushroom cultivation is a huge positive field of mycology. In Asia there are over 10 research institutes and hundreds of scientists dedicated to the science and research of mushroom cultivation, here in the US, where we are still in the mushroom dark ages, there is probably 1 department in the entire country, at the University of Pennsylvania that looks at mushroom cultivation. With a minuscule 15 or so species of mushrooms cultivated in the US of the 1000's of edible mushrooms there is plenty of room to grow. Per capita consumption of mushroom is increasing in the US and will likely only expand in the coming years. Research and education looking at and promoting new methods of cultivation, new species of cultivation, increased consumption, methods of preparation and use, will be important as mycology continues to grow.

What is Mycology: Mushroom Supplements

Mushroom supplements are a rapidly expanding field. With growth expected to go from 20 billion in 2018 to 33 billion in 2024 much more research can go into the health benefits mushrooms offe r and the difference between supplements on the market. The pharmaceutical and health care industry is MASSIVE in the United States, Mycology could more and more play a big part in prevention and treatment of diseases in the United States including cancer, neural diseases, anxiety and depression, and mental health. From psilocybin to shiitakes, many species of mushrooms are being studied for different potential health effects.

Mycology is the study of mushrooms that could radically support the evolution of health care in the United States by studying how to use and incorporate mushrooms into creating a healthy body. At the same time there are many questions of what anatomical part of the mushroom is best to use for health benefits. Mycelium on grain is commonly used in US created supplements, but the efficacy is questionable, with up to 60% of the biomass remaining as grain and only 30-40% being converted to fungal tissue. Studying the difference in efficacy between these products is critical for the health of both the supplement industry and the consumer. Finding ways to viably convert mushroom fruiting bodies into extracted supplements is an important role mycology could pick up

What is Mycology: Mycoremediation

Everyone knows we humans have a waste problem. Not only quantity but quality of our waste is extremely bad. Here come the fungi! Mycology has an AMAZING opportunity to work with the issues of massive amounts of inorganic waste material along with massive spills of toxic materials. The potential of fungi to digest plastic is incredible and developing this into a viable large scale solution to plastic waste is a cause worth fighting for! What is more is that fungi have the unique capacity to break down petrochemicals which are spilled into the environment.

Mycology could be a leading field in remediation of the planet and response to catastrophes like oil spills. By developing methods of collecting petrochemicals and inoculating them with massive amounts of mushroom material mycology could have massive positive impacts on the world. It is time for Mycology to evolve beyond identification and fungal diseases. Mycology needs to embrace some of the newer areas of application and issues the world is facing. Instead of removing mycology departments from universities and cutting budgets mycology as a field should be expanded as an inter-disciplinary field impacting everything from human health to planet health.

Hopefully you have a better grasp on what is mycology. Overall, mycology is the study of mushrooms, but it goes so much deeper than that. How do you want to see mycology applied in the 21st century? Are you working on amazing projects transforming waste into healthy soils with fungi? Let us know about it as there might be a way we can help lend support!

Choosing a selection results in a full page refresh.
Opens in a new window.

The contribution of fungi to the global economy

Open access
Published: 12 July 2023
Volume 121 , pages 95–137, ( 2023 )

Cite this article

You have full access to this open access article

Allen Grace T. Niego 1 , 2 , 3 ,
Christopher Lambert 4 , 5 ,
Peter Mortimer 6 ,
Naritsada Thongklang 1 , 2 ,
Sylvie Rapior 7 , 8 ,
Miriam Grosse 4 , 5 ,
Hedda Schrey 4 , 5 ,
Esteban Charria-Girón 4 , 5 ,
Arttapon Walker 1 , 2 ,
Kevin D. Hyde 1 , 2 , 9 , 10 &
Marc Stadler ORCID: orcid.org/0000-0002-7284-8671 4 , 5

22k Accesses

39 Citations

122 Altmetric

10 Mentions

Explore all metrics

Fungi provide ecological and environmental services to humans, as well as health and nutritional benefits, and are vital to numerous industries. Fermented food and beverage products from fungi are circulating in the market, generating billions of USD. However, the highest potential monetary value of fungi is their role in blue carbon trading because of their ability to sequester large amounts of carbon in the soil. There are no conclusive estimates available on the global monetary value of fungi, primarily because there are limited data for extrapolation. This study outlines the contribution of fungi to the global economy and provides a first attempt at quantifying the global monetary value of fungi. Our estimate of USD 54.57 trillion provides a starting point that can be analysed and improved, highlighting the significance of fungi and providing an appreciation of their value. This paper identifies the different economically valuable products and services provided by fungi. By giving a monetary value to all important fungal products, services, and industrial applications underscores their significance in biodiversity and conservation. Furthermore, if the value of fungi is well established, they will be considered in future policies for effective ecosystem management.

The importance of fungi and mycology for addressing major global challenges

Amazing Potential and the Future of Fungi: Applications and Economic Importance

Fungi: Essential Elements in the Ecosystems

Avoid common mistakes on your manuscript.

Introduction.

Methods used to give monetary value to fungi

Data gathering

Market-based approach

Value Transfer Method (VTM)

Total Economic Value (TEV)

Real value of products and services from fungi

Value of carbon stocks being traded from forests

Industrial uses of fungi, medicines and pharmaceuticals.

Antibacterials

Penicillins

Cephalosporins

Fusidic acid

Pleuromutilin

Antimycotics

Echinocandins

Enfumafungin

Griseofulvin

Cardiovascular drugs

Immunosuppressive and immunomodulatory agents

Cyclosporin

Mycophenolic acid (mycophenolate mofetil)

Nematicides

Traditional Chinese medicines

Ophiocordyceps/Cordyceps

Food and beverages

Functional food and nutraceuticals

Agaricus bisporus

Hericium spp.

Lentinula edodes

Monascus purpureus

Ophiocordyceps sinensis

Fermented food

Baked goods

Alcoholic beverages and spirits

Non-alcoholic fermented beverages

Fungal food additives: organic acids

Citric acid

Fumaric acid

Gluconic acid

Itaconic acid

Lactic acid

Fungi based proteins

Food enhancers

Indonesian tempeh

Food colorants/pigments

Astaxanthin

Carotenoids

Monascus pigments

Commodities

Ergothioneine

Mycopesticides

Strobilurin fungicides

Mycorrhiza-based biofertilizers

Fungal enzymes

Galactosidase

Value of wild and cultivated mushrooms being traded for food

Emerging myco-based material industry, recreational fungal foraging, other potential biotechnological applications of fungi, bioremediation, monetary valuation of fungi, future perspectives/implication to policy makers, acknowledgements.

It is irrefutable that fungi play a pivotal role in any ecosystem. Detrital pathways are necessary for carbon and nutrient cycling (Liu et al. 2020a ). Fungi have very essential functions in addressing major global challenges. Wild mushrooms, especially if they can be cultivated, constitute a significant non-timber forest resource that provides various benefits, especially food and income for local communities in many parts of the world (Mortimer et al. 2012 ). They can be instrumental in improving resource efficiency as renewable substitutes for fossil resources. Moreover, they can be valuable sources of food, nutraceuticals, cosmeceuticals, amino acids, proteins, carbohydrates, vitamins, fats, and minerals (Ślusarczyk et al. 2021 ; Badalyan et al. 2022 ). They produce manifold bioactive compounds with medicinal, pharmaceutical, and industrial applications (Hyde et al. 2019 ; Niego et al. 2021 ; Galappaththi et al. 2022 ).

These numerous benefits of fungi are often overlooked, and the general population is often unaware of their value. Applied research has unfolded the role and potential of fungi in environmental services, such as bioremediation, mitigation of pollution, and as a resource for the development of drugs to combat various human diseases (Hyde et al. 2019 ). To fully utilize the potential of fungi, we need to understand their services rendered in ecosystems on a global scale. With the move towards a biobased circular economy (Meyer et al. 2020 ), we now consider it appropriate to make the world aware of the immense value of fungi for sustainable global development. Economic valuation is essential to create transparency with respect to the services rendered by fungi in ecosystems compared to other economic assets. This will provide clarity in political discussions, often dominated by monetary worth since ecosystem conversion appears to be more profitable than conservation. Economic criteria represent as well key elements during decision-making processes regarding conservation policies (Martin-Lopez et al. 2008 ; Takala et al. 2021 ).

The current review paper narrates the contribution of fungi in bio-economies. A greater understanding of the roles and value of fungi can be used to build global support for mycological appreciation. It will, in particular, raise policymakers' awareness of the importance of fungi, leading to future research and conservation efforts.

Methods used to give monetary value to fungal services

Data sources.

The available data were collected and extrapolated from published studies in scientific journals, databases, online market research, and reliable news and articles. Scientific journals were the major sources of reports for this paper. The published academic papers considered are from reliable journals, not listed in Beal’s list of potential predatory journals and publishers ( https://beallslist.net/ ). Key databases, which are listed below, provided reliable data for chemical/drug information (ChEBI 2022 ; Drugbank Online 2022a ; National Center for Biotechnology Information 2022a ; Special Chem 2022 ), industry reports and international trade reports (DEFRA 2006 ; OEC 2022a ; Tridge 2022a ), agricultural production (Food Data Central 2019 ; UNData 2022 ) and information on the populations (World Population Review 2022 ). Online market research reports are the main source for the market value of fungal products used in our study (Table 1 ). They provided reliable sources of data since they are based on literature analysis, annual reports, technical papers, news releases, white papers, conference papers, government publications, trade data and other literature for market studies. News on TV and social media, as well as articles in newspapers and magazines featuring economy news and market reviews can also provide the needed knowledge for fungal-marketed products, since many marketed values are not published in scientific journals (Table 1 ). However, we had experienced that some of the provider companies requested substantial fees when we asked them for details. This is why we were not able to add data on some of the included product lines.

Identification of economically important products and services of fungi

The first and most relevant method for this article is to identify and list the major individual products and services of fungi that contribute to the global economy. The marketable products and services are categorized into (i) wild and cultivated mushrooms traded for food, (ii) industrial uses of fungi (e.g. medicines and pharmaceuticals, food and beverages, and commodities), (iii) emerging biomaterial industries (e.g. myco-based materials, biofuels), (iv) marketable ecosystem services (carbon stock, fungal foraging). Only the top products, considered as those with a minimum global market value of at least a million dollars, were included. The valuation of the included products was taken from 2017 to 2022 or the latest available data point, respectively.

Literature reviews

Whenever possible, the monetary value of each product or service was identified based on reliable data sources, which were listed above. However, when there were not enough direct data available in the literature, monetary values were determined by using the value transfer method (Johnston and Wainger 2015 ).

Value transfer method

The value transfer or benefit transfer method refers to applying quantitative estimates of ecosystem service values from existing studies to another context. The values given in the literature are monetary estimates of benefits or costs. Data have been extrapolated from a study site and applied with adjustments to other policy sites. Value transfer is a continuum of approaches depending on the information available (Johnston and Wainger 2015 ). This method is applicable in giving monetary value to fungal traded services such as carbon stocking and recreational foraging.

Summation of values

Market values are categorized as definite value, product value and traded value, respectively. Definite value is the monetary value of a fungus itself (e.g., Agaricus bisporus and “ Cordyceps ”—i.e., Ophiocordyceps ; edible fungi: wild and cultivated fungi). Product value is the market value of the products derived from fungi or with the involvement of fungi in the production process (e.g., medicines and pharmaceuticals, fermented food and beverages, commodities and mycobased-material). Traded value is the market value from the services of fungi (e.g., carbon stock, recreational foraging). All these marketed values are combined to give an estimate of the monetary value of the fungal contribution to the global economy. All monetary values are given in US dollars (USD). For numerous industries that rely on fungi for the production processes, we have assumed the total industry value as the value of fungi, as it is impossible to disentangle the values of individual ingredients, and these products would not exist without the fungi involved.

Scope and limitation of the study

This study focusses on consumptive and marketed products and services rendered by fungi with direct effect on human well-being. Only those with a large impact on the global economy (minimum of USD 1 million) are included in the summation of values. Other ecosystem services and important roles in ecosystem functioning by fungi listed in Niego et al. ( 2023 ) with an indirect effect on humans will not be considered for valuation in this study, but future research is recommended to give them monetary value.

According to the Kyoto protocol, the Clean Development Mechanism (CDM) promotes afforestation and reforestation as carbon sequestration activities. It was expected to be the most effective and easily measurable method for impounding carbon as biomass above and below the ground. During the negotiation of the Kyoto Protocol under the United Nations Framework Convention on Climate Change, 38 countries were required to reduce greenhouse gas (GHG) emissions by at least 5.2% below the 1990 levels. This goal was achieved by reducing greenhouse gas emissions or increasing greenhouse gas sinks (Tulpule et al. 1998 ; Grubb et al. 1999 ). A carbon price is an instrument used to capture the cost of greenhouse gas emissions, such as the damage caused to the environment. Governments and business sectors increasingly agree that carbon pricing plays a vital role in the transition to a decarbonized economy. In addition to charging emitters for carbon dioxide (CO 2 ) emissions, carbon pricing also provides incentives to individuals who emit less (Barron et al. 2019). Global carbon trading was valued at €760 billion (USD 817 billion) in 2021 with EU emission allowances above €80/t (USD 86/t) (Refinitiv 2022 ).

Carbon stock in forests can be found as 31% biomass and 69% in the soil (IPCC 2005 ). In the study by Clemmensen et al. ( 2013 ), it was stated that 50–70% of sequestrated carbon in the soil was derived from roots and root-associated microorganisms such as mycorrhizal fungi. Approximately 85% of the terrestrial plants form mycorrhizal associations with fungi. The ectomycorrhizal and ericoid mycorrhizal plants associates are the majority of trees (92%) and shrubs (85%), while arbuscular mycorrhizal plant associates are mostly herbaceous (50%) (Soudzilovskaia et al. 2020 ).

Different types of mycorrhizal association present in forest ecosystems can affect the global distribution of carbon stock. Soudzilovskaia et al. ( 2019 ) estimated the storage of carbon in arbuscular (AM), ectomycorrhizal (EcM) and ericoid (ER) mycorrhizal vegetation in above-ground biomass as 241 ± 15, 100 ± 17, and 7 ± 1.8 billion metric tons, respectively. We assumed that the EcM and ER vegetations are in the forest, since these groups of fungi are mostly associated with trees. Based on the recent study of Edy et al. ( 2022 ) AM fungi are also present in 75% of tropical forests. The monetary values of carbon sequestered by different mycorrhizal vegetation were estimated by multiplying the values by the carbon emission allowance of USD 86/t of carbon (Refinitiv 2022 ) (Table 2 ). The total monetary value of the carbon stock from above-ground biomass by mycorrhizal vegetation worldwide is USD 24,768 billion, equivalent to 31% of the carbon stock in the forest (IPCC 2005 ). From this value, the total value of carbon stock in the forest is estimated to be USD 79,897 billion, of which USD 55,129 billion is contributed by soil carbon of 69% (IPCC 2005 ). Since 50–70% of the sequestered carbon in the soil is associated with root-associated microorganisms, we can roughly assume that the value of the carbon stock in the soil related to mycorrhizae is at least USD 27,565 billion. Thus, the total market value of the carbon stock associated with mycorrhizae is USD 52,333 billion .

Fungi and the welfare obtained from them have been instrumental in the development of essential industrial processes worldwide. Among others, the secondary metabolites produced by these organisms represent a valuable resource in the development of medicines and pharmaceuticals. Fungal-based processes and fungal enzymes also play a key role in the production of food and beverages, as well as different commodities. The market value of these products steadily increases as the years go by. This chapter will discuss the different products and their market values, based on the available data.

Fungi have been explored for their medicinal value over thousands of years (Zhang et al. 2015 ). Aside from nutritional benefits, their ability to produce secondary metabolites makes them an important source for innovative chemistry, which led to the development of several important pharmaceuticals that have been beneficial for the advance of human civilization over time. While natural products and secondary metabolites in general have resulted in many drugs for the treatment of various diseases (Newman and Cragg 2020 ), fungal metabolites have been only developed for part of these indications, as amply summarized by Bills and Gloer ( 2016 ). For instance, contrary to what is stated in many reviews that have been published on the subject, there are no ethical anticancer drugs from fungi on the market. Taxol, for instance, is exclusively produced using plants as sources and the recent classification of the fungal “endophyte” that was reported to be able to produce the metabolite as a member of the Basidiomycota (Cheng et al. 2022 ) raises serious concerns as to whether fungi can produce this metabolite at all. In any case, fungal metabolites have given rise to the development of several drugs that gained a rather high commercial value in other indications, above all as anti-infectives, but even cardiovascular and immunomodulatory agents. However, even the discovery of some blockbuster drugs like cyclosporin has been attributed to their ability to exert antibiosis, and their true potential was discovered later when they were tested in more sophisticated bioassays. Growth inhibitory or toxic properties of fungal secondary metabolites have regularly been linked with ensuring their survival in competitive natural environments (Keller 2019 ). Of the fungal natural product—derived drugs found on the market, the majority were originally isolated from species of the Ascomycota, while a few are derived from Basidiomycota (Bills and Gloer 2016 ; Sandargo et al. 2019 ). Compared to the vast number of described compounds, only a handful have been followed up upon to be further developed into marketable drugs (Mapook et al. 2022 ) (Table 3 ). Nevertheless, the secondary metabolite-inspired drugs that are on the market have been proven indispensable for modern medicine and managing diseases in general, displaying important sources of economic revenue.

Penicillins are the first β-lactam antibiotics explored for their medicinal value and are still one of the most widely used antibiotic agents to fight bacterial infectious diseases. They were first described from Penicillium rubens (Houbraken et al. 2011 ) but also reported from other species such as P. chrysogenum and P. griseofulvum (Laich et al. 2002 ). Penicillins are still in use to treat a wide range of bacterial infections (Lobanovska and Pilla 2017 ). Aside from the formulated naturally occurring derivatives penicillin G and V, other semi-synthetic derivatives among the vast diversity of analogues are Amoxicillin, Ampicillin, Piperacillin, Dicloxacillin, Nafcillin and Oxacillin (Wright 1999 ). The global Penicillin market size is estimated to be worth USD 1.96 billion in 2022 and is forecasted to increase to USD 2.4 billion by 2028 with a compound annual growth rate of 3.8% (Market Watch 2022a ).

Cephalosporins are another class of β-lactam antibiotics originally isolated from strains of the genus Acremonium. Following its first report by Newton and Abraham ( 1955 ), a semi-synthetic derivative of cephalosporin C named cephalothin reached the market in 1964 (Asbel and Levison 2000 ). Since then, in total five generations of cephalosporin-derived antibiotics were developed, further improving upon its selectivity against drug resistant bacteria (Lin and Kück 2022 ). Among the indications are skin infections, with and without multi drug resistant bacteria and meningitis, to name a few (Lin and Kück 2022 ). The global cephalosporin market size was valued at USD 18.7 billion in 2022 , and it is expected to reach USD 22.3 billion by 2028, exhibiting a CAGR of 2.83% during 2023–2028 (Imarc 2022a ).

First generation cephalosporins are commonly used to manage skin and soft tissue infections caused by susceptible strains of Staphylococcus aureus and group A Streptococcus . Those include cefazolin, a parenteral formulation, and cephalexin, an oral equivalent, among others (Lin and Kück 2022 ).

Second generation cephalosporins have a better performance against Gram-negative bacterial infections than the first-generation derivatives and are used to treat urinary and tract infections, and skin and soft tissue infections. A variety of orally administered second-generation agents (cefaclor, cefprozil, loracarbef, cefpodoxime) are commonly used in the outpatient management of sinopulmonary infections and otitis media (Lin and Kück 2022 ).

Third generation cephalosporins are typically used for serious pediatric infections, including meningitis and sepsis (Lin and Kück 2022 ).

Fourth generation cephalosporins include, e.g. cefepime, which is active against Pseudomonas aeruginosa and retains good activity against methicillin-susceptible staphylococcal infections (Lin and Kück 2022 ).

An example for Fifth generation cephalosporins is ceftaroline, the active metabolite of the prodrug ceftaroline fosamil. This is a broad-spectrum cephalosporin with bactericidal activity against resistant Gram-positive bacteria, including Methicillin-resistant Staphylococcus aureus (MRSA), and common Gram-negative pathogens (Lin and Kück 2022 ). The development of cephalosporins is a prime example of how a basic structure derived from natural sources can be gradually improved via semisynthesis and medicinal chemistry approaches, finally leading to broad spectrum antibiotics. However, this development has taken several decades of hard work.

In this context, it is also worthwhile to mention the carbapenems , which are also β-lactam antibiotics but were discovered from Streptomyces species, which are Gram-positive Actinobacteria. Olivanic acids were the first isolated carbapenems from S. clavuligerus in a screening campaign for β-lactamase inhibitors (Brown et al. 1976 ), but the compound family only went into clinical relevance with the isolation of thienamycin from S. cattleya , which exhibited superior physicochemical properties (Kahan et al. 1979 ; Papp-Wallace et al. 2011 ). Carbapenems combine two activities as being able to bind to the same target as other β-lactams, in addition to their inherent property to act as a β-lactamase inhibitor. The first carbapenem to enter the market was the semi-synthetic analogue imipenem (Miyadera et al. 1983 ) in 1985, which was followed by other synthetic analogues like meropenem (Sunagawa et al. 1990 ), biapenem (Ubukata et al. 1990 ), ertapenem (Sundelof et al. 1997 ), doripenem (Tsuji et al. 1998 ), and panipenem (Neu et al. 1986 ). These compounds are used to treat a broad band of bacterial infections, including complicated nosocomial bacteriosis, or infections caused by multi-drug resistant staphylococci and Pseudomonas (Papp-Wallace et al. 2011 ). Due to their inability to pass the gastrointestinal tract, they are commonly administered intravenously. However, the new drug tebipenem is currently in clinical trials as the first orally available carbapenem (Jain et al. 2018 ). The global carbapenem market was valued at USD 3.9 billion in 2021 and is expected to exhibit a growth rate (CAGR) of 4.5% during 2022–2030 (Polaris Market Research 2022 ).

Fusidic acid is a bacteriostatic triterpene derived from the fungus Ramularia coccinea (Godtfredsen et al. 1962 ; as Fusidium coccineum ) and used as a topical medication for the treatment of skin infections with Gram-positive bacteria (Fernandes 2016 ). It acts as a bacterial protein synthesis inhibitor by binding to the elongation factor G (EF-G) bound to ribosomes. This effectively prevents the translocation of the newly assembled polypeptide chain during protein synthesis. Recycling of ribosomal subunits is thus inhibited after completion of transcription (Gao et al. 2009 ) The global fusidic acid market value was estimated at USD 171.89 million in 2021 with projection to reach up to USD 271.84 million by 2028, exhibiting a CAGR of 6.77% during the forecast period (Market Watch 2022b ).

Pleuromutilin and its derivatives are antibacterials produced by Basidiomycota that inhibit protein synthesis in bacteria by inhibiting peptide bond formation through binding to the 50S subunit of ribosomes (Paukner and Riedl 2017 ). For details we refer to the recent review by (Mapook et al. 2022 ). The pleuromutilin derived compounds tiamulin and valnemulin covered 2.5% of total sales of antimicrobials used for food producing animals (European Medicines Agency 2013). Lefamulin, a new pleuromutilin antibiotic with the brand name XENLETA™, which is used for the treatment of community-acquired pneumonia (CAP), is expected to peak sales of USD 267 million by the end of 2025 (Joseph 2019 ). As this drug is fairly new to the Pharma market and has a molecular target that is not addressed by the conventional antibiotics, a substantial increase in the sales is to be expected in the coming years (Fig. 1 ).

Chemical structures of antibacterial drugs

Echinocandins are a group of cyclic non-ribosomal lipopeptides reported from Ascomycota of the classes Leotiomycetes and Eurotiomycetes (Hüttel 2021 ). Currently, there are three semi-synthetically modified drugs, derived from echinocandin B (producer: Aspergillus delacroxii ; Benz et al. 1974 ), pneumocandin A 0 ( Glarea lozoyensis ; Schwartz et al. 1989 ) and FR901379 ( Coleophoma empetri ; Iwamoto et al. 1994 ) on the market. All three drugs, anidulafungin , caspofungin and micafungin are used to treat and prevent invasive fungal infections including candidemia, abscesses, esophageal candidiasis, and certain other invasive Candida infections (Hüttel 2021 ). Furthermore, these pharmaceutics are used as prophylaxis for Candida infections during stem cell transplantation. Antifungal effect relies on the inhibition of the synthesis of β-(1,3)- d -glucan, an integral component of the fungal cell wall (Sawistowska-Schröder et al. 1984 ). The global market size for echinocandins was valued at USD 515.20 million in 2021 and is expected to reach USD 796.69 million by 2029, with a CAGR of 5.6% (Data Bridge Market Research 2022 ).

Enfumafungin is a triterpene glycoside and hemiacetal derived from the endophytic fungus Hormonema sp. (Peláez et al. 2000 ). It is a potent antimycotic which, similarly to the echinocandins, inhibits the biosynthesis of β-(1,3)- d -glucan (Jallow and Govender 2021 ). Its semi synthetic derivative Ibrexafungerp was the first-in-class triterpenoid antifungal drug with broad spectrum antifungal activity (Ghannoum et al. 2020 ) that has been approved for the treatment of vulvovaginal candidiasis (VVC) and only recently entered the market (Espinel-Ingroff and Dannaoui 2021 ). It is sold under the brand name Brexafemme® approved by the FDA in June 2021 with net revenues of USD 0.7, 1.3, and 1.16 million in the first three quarters of 2022 respectively (GlobeNewswire 2022 ; Scynexis 2022 ).

Griseofulvin is a secondary metabolite produced by Penicillium spp. and was first discovered from P. griseofulvum (Oxford et al. 1939 ). It has been developed as an orally available, fungistatic drug used to treat superficial fungal skin infections such as tinea capitis and pedis (Gupta et al. 2017 ). The onychomycosis treatment market value exceeded USD 4.2 billion in 2020 and expected to reach USD 6.2 billion in 2027 (Global Market Insights 2022a ). Although there is no market value reported in the literature for griseofulvin, it is worth mentioning this important product because it is one of the most valuable treatments for onychomycosis on the market (Fig. 2 ).

Chemical structures of antimycotic drugs

Statins are prescribed for their cholesterol-lowering capacity to prevent mortality linked towards cardiovascular diseases, such as coronary artery disease and stroke (Endo 2008 ). Among the marketed statins are pravastatin , the semi-synthetic product of compactin (or mevastatin) isolated from Penicillium citrinum (Endo et al. 1976 ), lovastatin and its semi-synthetic derivative simvastatin (Pedersen and Tobert 2004 ). In addition, some of the synthetic statins inspired by nature introduced to the market are fluvastatin, atorvastatin, rosuvastatin and pitavastatin (Endo 2010 ). Lovastatin was concurrently isolated from Aspergillus terreus and Monascus ruber (reported as Monacolin K, Endo 1979 ) and was one of the first statins to be extensively used for its cholesterol-lowering ability (Steinberg 2006 ) and even discussed as auxiliary drugs in treatment regiments against cancer (Jiang et al. 2021 ). The global statins market size was valued at USD 14.3 billion in 2021 , and was expected to reach USD 17.5 billion by 2027, with a CAGR of 3.4% during 2022–2027 (Imarc 2022b ) (Fig. 3 ).

Chemical structures of cardiovascular drugs

Cyclosporin A represents the first calcineurin inhibitor, which was reported from the hypocrealean soil fungus Tolypocladium inflatum (originally identified as “ Trichoderma polysporum ” by Rüegger et al. 1976 ) and entered the market in 1983 (Survase et al. 2011 ). This substance was shown to exhibit a better bioactivity profile in terms of its immunosuppressive activities as compared to other synthetic drugs on the market, which turned out to be a game changer in the field of transplantation medicine (Demain and Sanchez 2009 ; Survase et al. 2011 ). The market sales of cyclosporin A totaled to USD 1.5 billion in 2009. Recently, cyclosporine was valued at USD 1.99 billion in 2021 , and it is projected to reach the value of USD 6.08 billion in 2030 (Verified Market Research 2022a ).

Fingolimod is an immunomodulating drug developed from myriocin. It was first reported from Melanocarpus albomyces (previous name: Myriococcum albomyces ; cf. Kluepfel et al. 1972 ). The semi-synthetic derivative was shown to modulate the sphingosine 1-phosphate receptor, which is exploited to treat patients with a relapsing–remitting form of multiple sclerosis (MS, Volpi et al. 2019 ). It is sold since 2010 (Thomas et al. 2017 ) under the brand name Gilenya (Drugbank Online 2022b ). Gilenya has a market share of USD 79,411 per 1000 Medicare beneficiaries in 2014 (San-Juan-Rodriguez et al. 2019 ) with annual sales worth USD 3.34 billion in 2018 (Miller 2019 ).

Mycophenolic acid is derived from Penicillium brevicompactum and originally reported by Gosio ( 1893 ) as the first fungal antibiotic. However, it was only much later that systemic studies and semi-synthetic derivatization improved its toxic side effects to enable its usage as an immunosuppressant for organ transplantations as of starting from 1995 (Lipsky 1996 ). The compound itself, however, is toxic and commonly prescribed as its formulated prodrug, mycophenolate mofetil. Upon release of its active component (mycophenolic acid), it inhibits the inosine monophosphate dehydrogenase, which is required for the de novo synthesis of guanosine (Lenihan and Tan 2020 ). The global market value of mycophenolate mofetil was USD 1581.34 million in 2022 (Market Watch 2023a ). Mycophenolate sodium was later introduced to the market as an enteric-coated slow-release formulation of mycophenolic acid to decrease the gastrointestinal side effects of mycophenolate mofetil (Lenihan and Tan 2020 ). This drug has been in discussion for its application in other medical conditions associated with an apparently hyperactive immune systems as well, such as psoriasis (Bentley 2000 ).

Mizoribine is an immuno-suppressive sold in Japan, Korea, and China and commonly used for treatment of renal diseases and prevention of renal transplants rejection (Kawasaki 2009 ). The compound was first isolated from Penicillium brefeldianum (Mizuno et al. 1974 ) and came on the market in 1984, however, limitations concerning its efficacy allegedly limits its use worldwide (Kawasaki 2009 ). Mizoribine is estimated to be worth several million USD in 2021, however, the exact value was not available to us (Fig. 4 ).

Chemical structures of immunosuppressive and immunomodulatory agents

Emodepside has been the first drug on the market that is derived from a secondary metabolite produced by a fungal endophyte (Sasaki et al. 1992 ; Scherkenbeck et al. 2002 ; Helaly et al. 2018 ). Molecular Data in patents already pointed towards this fungus, which was isolated from a tea plant in Japan, being a member of the genus Rosellinia (Harder et al. 2011 ). However, it was only shown recently that ascospore-derived isolates of the latter xylariaceous genus can indeed be linked with the production of the nematicidal PF1022 derivatives (Wittstein et al. 2020 ). The semi-synthetic derivative is marketed under Emodepside and used as an anti-helminthic drug in veterinary medicine (Willson et al. 2003 ). Currently, attempts are ongoing to develop the compound as a remedy for human pathogenic nematode infections. In this sense, emodepside appears as a promising candidate with a novel mode of action against human river blindness (onchocerciasis), a disease caused by the filarial worm Onchocerca volvulus , and which treatment relies on the natural product ivermectin (Krücken et al. 2021 ). Recently, it has been validated that emodepside exhibits good in vitro efficacy against microfilarial, third larval, fourth larval, and adult stages of various filarial genera and species (Hübner et al. 2021 ). There are no data available on the current market value of emodepside. The compound is only administered with another synthetic anthelmintic agent, praziquantel, in veterinary medicine (Fig. 5 ).

Chemical structures of nematicidal agents

Traditional Chinese medicines (TCM)

Traditional Chinese medicine (TCM) has existed for almost 5000 years and was historically used by the Chinese to combat diseases. Until now, interest and acceptance of TCM continue to grow worldwide (Lin et al. 2018 ). Fungi have always been part of TCM, and the most explored species belong to the genera Ophiocordyceps, Cordyceps, and Ganoderma (Paterson 2006 , 2008 ).

Ophiocordyceps/Cordyceps preparations are commonly used in traditional Chinese medicines in China, Japan, Korea, and other eastern Asian countries (Zhu et al. 1998 ). They were used in China for at least 2000 year (Zhu et al. 1998 ). Ophiocordyceps sinensis , Cordyceps militaris, Cordyceps guangdongensis, and Isaria cicadae are the most explored and commercialized “ Cordyceps ” fungi in China (Zhu et al. 1998 ; Zhang et al. 2014 ). There is a huge demand in Chinese markets for O. sinensis, with an annual yield of 100–150 tons (of which approximately 100 tons are produced in China). The market price is USD 8 to USD 25 each or USD 25,000 to 60,000 per kilogram (Zha et al. 2018 ). The national annual value of natural Chinese Cordyceps is estimated to be over 10 billion RMB (USD 1.5 billion) in 2004 (Dong et al. 2015 ).

Ganoderma -based products are marketed in many Asian, European and North American countries, although South Asian countries such as China, Japan, Korea, Malaysia and Singapore are the main producers of the food products (Chang and Miles 2004b ). The annual global market turnover of Ganoderma -based products was approximately USD 2.16 billion in 2002 (Lai et al. 2004 ). In 2019, the global market value of Ganoderma- based products was USD 3.1 billion , and projected to reach USD 5.1 billion in 2027 (Allied Market Research 2022a ).

Fungi have long been explored due to their medicinal value (De Silva et al. 2013 ; Hyde et al. 2019 ; Badalyan and Rapior 2021 ) and many fungal products are produced as nutraceuticals, including cosmeceuticals (Bandara et al. 2015 ; Badalyan et al. 2022 ). Nutraceuticals are a group of products from edible sources, proven to be safe for consumption and can be considered a supplement to effective pharmacological treatment (Niego et al. 2021 ). Fungal products are being consumed as functional foods, dietary supplements, and sources of bioactive compounds with a range of medicinal applications (Hyde et al. 2019 ; Badalyan et al. 2021 ). Some of the most explored genera are Agaricus, Hericium, Lentinula, and Ophiocordyceps (Valverde et al. 2015 ; Hyde et al. 2019 ; Niego et al. 2021 ). The reported bioactivities include antibacterial, antidiabetic, antifungal, antihypercholesterolemial, antioxidant, antiparasitic, antitumor, antiviral, cardiovascular, hepatoprotective, immunomodulatory and radical scavenging effects (Hyde et al. 2019 ; Zeb and Lee 2021 ; Niego et al. 2021 ). The bioactive principles causative for those effects are polysaccharides, proteins (El Enshasy and Hatti-Kaul 2013 ), glycoproteins (Cohen et al. 2014 ), phenolic compounds (Gupta et al. 2018 ; Hamwenye et al. 2022 ), ergosterol (Diallo et al. 2020 ) and other terpenoids, or unsaturated fatty acids (Galappaththi et al. 2022 ). The health-promoting properties of many fungi are the driving force for increasing their market value and incorporation into the nutraceutical industry (Niego et al. 2021 ). The following are economically important fungi explored in the nutraceutical and functional food industries (Table 4 ).

The bioactivities of Agaricus bisporus have been quite established. It is one of the most explored mushrooms for nutraceuticals (Gopalakrishnan et al. 2005 ). Agaricus bisporus was valued globally at USD 16.69 billion in 2022 (Market Data Forecast 2022a ).

Hericium species (and in particular H. erinaceus ) are highly valued for their medicinal properties. China is the main exporter of Hericium with a market value of USD 270 million, followed by Italy (USD 90.97 million) and Poland (USD 65.83 million) in 2021 (Tridge 2022a ). The global export value of Hericium was USD 963.4 million and the import value was USD 978.83 million in 2021 (Tridge 2022b ).

The nutraceutical properties of Lentinula edodes have been well documented (Liu et al. 2020b ; Wang et al. 2020 ; Raghoonundon et al. 2021 ). The global market value of Lentinula (Shiitake) is USD 4.31 billion in 2022 and is expected to reach USD 4.7 billion in 2030 at a CAGR of 8.3% (The Brainy Insights 2022 ).

Red yeast rice was created by fermenting Monascus purpureus on white rice. In traditional Chinese medicine, the powdered yeast-rice mixture has been utilized as a staple food in Asia (Nguyen et al. 2017 ). For heart disease and high cholesterol, people consume red yeast rice orally as a supplement. (Dufossé 2019 ). The cholesterol-lowering properties and other beneficial effects that have been proven for the product is, however, not due to the pigments (Li et al. 1998 ). The substance monacolin K (see above under “statins”) is one of them (Nguyen et al. 2017 ). The red yeast rice had a market value of USD 380.6 million in 2022 (Persistence Market Research 2022 ).

Ophiocordyceps sinensis is one of the most well-known and most expensive fungal species worldwide, with a market price that reached up to USD 60,000/kg in 2015 (Lei et al. 2015 ). Various nutraceutical products have been derived from Ophiocordyceps sinensis. In China, more than 1000 nutraceutical products are prepared from this species (Rakhee et al. 2021 ). The global market value of O. sinensis was around USD 5–11 billion in 2018 (He 2018 ).

The market value of these economically important mushrooms, however, was mainly due to the fact that they are also food sources and not solely functional food or nutraceuticals, particularly A. bisporus and L. edodes which are also some of the most cultivated mushrooms for food, thus will not be included in the summation for functional food to avoid double inclusion. By definition, functional food refers to food or food ingredients providing health benefits beyond basic nutrition (Colorado State University Extension 2023). The global functional mushroom market was then valued at USD 12.34 billion.

Fungi are beneficial to humans both industrially and commercially, especially in food and beverage production (Koul and Farooq 2020 ). Fermented foods and alcoholic beverages have long been an essential part of the human diet around the world. These foods are often well preserved with high nutrient content such as proteins, vitamins, minerals, and other nutrients (Tamang et al. 2020 ). The availability of nutrients in fermented foods and beverages is mainly due to the fermentation process performed by living microorganisms such as bacteria and fungi (Rezac et al. 2018 ). These microorganisms may also improve gastrointestinal health and provide other health benefits as probiotics (Rezac et al. 2018 ) that contribute to the popularity of fermented products (Fortune Business Insight 2022a ). Saccharomyces cerevisiae , Candida spp., Debaryomyces spp. and Wickerhamomyces anomalus are the most common yeasts associated with traditional fermentations of fermented foods and beverages (Bhalla and Savitri 2017 ). Saccharomyces cerevisiae is the yeast species most frequently involved in alcoholic fermentation and is also used in most other fermented products (Walker and Stewart 2016 ). Fermentation of foods and beverages changes the flavor and taste of the product, as well as its nutritional value and digestibility (Fortune Business Insight 2022a ). Table 5 lists some economically important fermented foods and beverages along with their global market value.

Baked goods are an essential part of human food. Saccharomyces cerevisiae is a very important fungus in the food production industry and is known as Baker's Yeast. It converts sugars and starches into alcohol and carbon dioxide during the fermentation process. Saccharomyces cerevisiae has been bred into various strains which produce significantly more CO 2 , and thus can achieve faster fermentation than wild type strains (Lahue et al. 2020 ). The global value of the market for bakery products reached USD 497.50 billion in 2022 (Imarc 2023 ). The top exporters are Germany (USD 4.3 billion), Canada (USD 3.14 billion), Italy (USD 2.73 billion), France (USD 2.28 billion) and the USA (USD 2.04 billion) (OEC 2022b ).

Cheese is a widely consumed dairy product with numerous varieties such as cheddar, feta, gouda, parmesan and camembert, produced in different parts of the world. Filamentous fungi and yeasts are used in the production of different kinds of cheese (Metin 2018 ; Chourasia et al. 2021 ). Filamentous fungi that thrive in cheese include Aspergillus, Cladosporium, Geotrichum, Mucor, Penicillium , and Trichoderma (Hymery et al. 2014 ) . There are two categories of mold-ripened cheeses: internally ripened and surface-ripened. The most well-known examples of internally ripened cheeses are the blue cheeses caused by Penicillium roqueforti creating blue veins, as the name suggests. The ripening of the famous Camembert-type soft cheeses relies on Penicillium camemberti (Lessard et al. 2014 ; Ropars et al. 2020 ) . Another example is Norwegian Gamalost with its ripening caused by Mucor with a yellow–brown color. Surface-ripened cheeses can be hard, semi-hard, and soft. The most common examples of hard surface-ripened cheeses are French Cantal, Salers, and Rodez, which involve species of Scopulariopsis or Sporendonema casei . The Saint Nectaire semi-hard cheese uses Mucor spp. and Bisifusarium domesticum as ripening molds (Zhang and Zhao 2010 ; Zhang et al. 2012 ; Metin 2018 ).

The global value of the cheese market was USD 231.65 billion in 2022 (The Business Research Company 2023 ). The top exporters were Germany (USD 4.79 billion), Netherlands (USD 4.11 billion), Italy (USD 3.57 billion), France (USD 3.49 billion), and Denmark (USD 1.71 billion) (OEC 2022c ). Blue cheese production had a total net trade of USD 624 million in 2020 wherein the top exporters are Italy (USD 185 million), Denmark (USD 122 million), France (USD 121 million, Germany (USD 94.8 million), and the United Kingdom (USD 17.1 million) (OEC 2022a ).

Alcoholic beverage and spirits

Yeasts, especially Saccharomyces cerevisiae, play a vital role in the production of fermented beverages. They are involved in the production of all alcoholic beverages and the selection of suitable yeast strains is essential not only to maximize alcohol yield, but also to maintain beverage sensory quality (Walker and Stewart 2016 ). Saccharomyces cerevisiae is the most common yeast used for many years in the production of beverages associated with the fermentation process (de Almeida Silva Vilela et al. 2020 ). Fermented beverages are made by yeast by converting sugar together with other essential nutrients such as amino acids, minerals, and vitamins into alcohol and carbon dioxide (primary fermentation metabolites) and other chief secondary metabolites (Walker and Stewart 2016 ; Vilela 2019 ). Secondary metabolites produced by yeast act as important flavor congeners of beverages that influence the final taste and aroma of alcoholic beverages such as beer and wine, as well as other distilled beverages such as brandy, rum, and whiskey (Walker and Stewart 2016 ). Yeasts are also involved in the production of white spirits such as gin and vodka (Pauley and Maskell 2017 ).

Beer is the most popular low-alcohol beverage consumed in huge amounts globally each year. It is important in the economy of many countries around the world (Maicas 2020 ). In Europe, traditional distinctions of beers are influenced by the different brewing practices, the water properties used in brewing, the type of malt and the yeast strains used (Britannica 2022a ). The European countries are some of the major producers and consumers of beers. The primary yeast involved in beer making is S. cerevisiae; however, other species such as S. eubayanus were also recently utilized in the brewing process (Gibson et al. 2017 ). Brewing is a complex fermentation process used in the beverage industries with huge annual revenues (Maicas 2020 ). Global beer production was valued at USD 605.20 billion in 2020 (Allied Market Research 2022b ).

Wine is an alcoholic drink made from fermented grapes. The varieties of grapes are the main factor in wine production. S. cerevisiae is also the main microorganism used in wine making. The global wine market size was USD 339.53 billion in 2020 which is projected to grow to USD 456.76 billion in 2028 (Fortune Business Insight 2022b ).

White spirits

Vodka is the most popular beverage in eastern Europe, traditionally made from potatoes and grain, especially rye (Menezes et al. 2016 ; Britannica 2022b ). The alcohol content of vodka is unaged and high concentration (40%) alcohol by volume (ABV), thus, can cause poisoning if over-consumed (Kanny et al. 2015 ; Pauley and Maskell 2017 ). The vodka industry was worth USD 39 billion in 2020 (Conway 2022 ).

Gin is an alcoholic beverage from junipers that has evolved greatly over the past 300 years from its origin with an alcohol content of 35–55% ABV (Abel 2001 ; Alcohol Rehab Guide 2022 ). It was previously invented as a remedy for military troops suffering from ‘East Indian fevers' (Abel 2001 ). The global gin market was valued at USD 14.03 billion in 2020 , and is expected to reach USD 20.17 billion by 2028 (Allied Market Research 2022c ).

Tequila is a distilled beverage made from the blue agave plant ( Agave tequilana ) originating from the western Mexican state of Jalisco (Graham 2022). It has an alcohol concentration of 38–55% ABV (Alcohol Rehab Guide 2022 ). The size of the tequila market has reached USD 12.89 billion in 2021 (The Spirit Business 2022 ).

Dark spirits

A dark spirit is a type of alcohol that has been aged in oak barrels, resulting in a dark color. With the exception of brandy, which is distilled from wine, dark spirits are beverages distilled from grains.

Rum is a distilled drink made from fermented sugarcane or molasses that originated in the Americas. It has an alcohol concentration of 40% ABV, but can go “overproof” with an alcohol concentration of 57.5–75% ABV (Alcohol Rehab Guide 2022 ). The highest valued rum comes from countries such as Barbados, Cuba, Guyana, Jamaica, and the Philippines (Brick 2022 ). The global value of the rum market was USD 15 billion in 2021, and is expected to reach USD 21.5 billion by 2027 (Market Data Forecast 2022b ).

Whiskey is a spirit made from fermented grains such as barley, corn, rye, and wheat aged in wooden casks. It has a typical alcohol concentration of 40–50% ABV (Alcohol Rehab Guide 2022 ). The global whiskey market was valued at USD 59.63 billion in 2019, and is projected to reach USD 86.39 billion by 2027 (Allied Market Research 2022d ).

Brandy is produced by distilling wine with an alcohol concentration of 35–60% ABV (Alcohol Rehab Guide 2022 ). Brandy varieties are well known throughout the world, such as Cognac and Armagnac from southwestern France (BBC 2022 ). The global brandy market was valued at USD 23.17 billion in 2020 and is expected to reach USD 25.32 billion by the end of 2027 (Globe Newswire 2022a ).

Our economy is significantly impacted by alcohol. Beer and wine were the most popular alcoholic beverages on the market. The total market value for alcoholic beverages made from fungi was USD 1,108.45 billion , demonstrating how popular drinking is worldwide.

Chocolate is primarily produced from cocoa beans ( Theobroma cacao ). Cocoa trees are native to the tropical regions of Central and South America. Most of the flavors that we recognize in chocolate only start to develop during the fermentation of beans (Carvalho 2016 ). Yeasts are also involved in nonalcoholic fermentation of chocolate production to improve chocolate flavor properties and add a new collection of aromas to chocolate (Maicas 2020 ). Yeasts are also responsible to produce several secondary metabolites that are flavor-active that add different fruity aromas to chocolate. These fruity volatile compounds include aldehydes, esters, fatty acids, higher alcohols, organic acids, phenols and sulphur-containing compounds (Carvalho 2016 ). The main yeasts involved in cocoa fermentation are Hanseniaspora opuntiae , Pichia kudriavzevii , and S. cerevisiae , which produce higher alcohols and acetyl-CoA to make acetate esters that give floral and fruity aromas (Gutiérrez-Ríos et al. 2022 ). Other yeasts found in cocoa formation that produce aromas are Galactomyces Galactomyces geotrichus and Wickerhamomyces anomalus (Koné et al. 2016 ). Saccharomyces, Pichia, Hanseniaspora, Kluyveromyces, Hansenula, Wickerhamomyces and Torulaspora are the predominant wild genera of yeast in cocoa fermentation in Colombia (Sandoval-Lozano et al. 2022 ).

There are approximately 4 million exporters of cocoa beans worldwide (International Trade Center 2022 ). The top exporters of chocolate and other food preparations containing cocoa in 2021 with their exported value were Germany (USD 32.82 billion), Belgium (USD 5.37 billion), Italy (USD 2.47 billion), Poland (USD 2.34 billion) and The Netherlands (USD 2.12 billion) (International Trade Center 2022 ). The global chocolate market value reached USD 130.56 billion in 2019 (Grand View Research 2022a ).

Coffee is one of the most consumed nonalcoholic beverages in the world. Due to its popularity and high demand for coffee quality from consumers, constant improvement and increase in variety are deemed necessary (Martinez et al. 2019 ; Ruta and Farcasanu 2021 ). Yeast processing can help to improve the quality and complexity of coffee (Walbank 2022 ). Coffee is an essential commodity exported to around 70 tropical and subtropical countries (FAO 2015 ). The global coffee market was valued at USD 102.02 billion in 2020 (Research and Markets 2022a ). The top coffee exporters according to net traded value were Brazil (USD 5.08 billion), Switzerland (USD 2.71 billion), Germany (USD 2.59 billion), Colombia (USD 2.54 billion), and Vietnam (USD 2.24 billion) (OEC 2022d ). However, like with other goods, it is unclear how much of the world coffee output is dependent on yeast fermentation, so the value provided here is an overestimation, but without more detailed market information, no further breakdowns of this amount can be calculated.

Vinegar is a liquid product formed from alcoholic fermentation and subsequent acetous fermentation of carbohydrate sources. It has been reported as medication in many cultures and documented for its beneficial health effects when consumed regularly (Ho et al. 2017 ) . Vinegar is the product of a two-step fermentation: In the first step, yeast converts sugars from fruits and grains to ethanol anaerobically, while in the second step, ethanol is oxidized to acetic acid aerobically by bacteria of the genera Acetobacter and Gluconobacter . However, alcohol formation is necessary to produce the required percentage of acetic acid (4–5%) for the liquid not to spoil (Tonkinson 2022 ). Several yeast species are involved in the fermentation of vinegar such as in traditional balsamic vinegar including Candida lactis-condensi, C. stellata, Hanseniaspora osmophila, H. valbyensis, S. cerevisiae, Saccharomycodes ludwigii as well as Zygosaccharomyces bailii, Z. bisporus, Z. lentus, Z. mellis, Z. pseudorouxii and Z. rouxii (Solieri and Giudici 2008 ). Vinegar has been an important household commodity and is used around the world. The global vinegar market reached a value of USD 2.27 billion in 2021 (Imarc 2022c ).

Kombucha is a beverage produced from the fermentation of tea ( Camellia sinensis ) by acetic bacteria and osmophilic yeasts (De Filippis et al. 2018 ). It is a good source of probiotics with numerous health benefits. Kombucha is an ancient drink dating back to as early as 220 B.C (Jayabalan et al. 2014 ). The yeasts involved are Brettanomyces/Dekkera , Candida , Torulaspora , Pichia , Schizosaccharomyces , Saccharomyces and Zygosaccharomyces (May et al. 2019 ) . The global market value for kombucha was USD 2.59 billion in 2021 (Polaris Market Research 2021 ).

Organic acids are one of the fastest growing sectors of the fermentation market globally, in which biotechnological production is an imperative economic alternative to chemical synthesis (Sharma et al. 2021 ). Organic acids are extensively used as ingredients in modern food processing (Copetti 2019 ). They are used mostly as buffers, preservatives, flavor enhancers, and adjuvants (Copetti 2019 ). Some organic acids from fungi have already made it to the market, such as citric, gluconic, and lactic acids made from fermentation of glucose or sucrose by Aspergillus niger (Copetti 2019 ). Other organic acids produced mostly by filamentous fungi but used to a lesser extent are fumaric, itaconic, malic, oxalic, succinic, and tartaric acids (Magnuson and Lasure 2004 ; Liaud et al. 2014 ; Dörsam et al. 2017 ).

Citric acid is an α-hydroxylated tricarboxylic acid found in citrus fruits. It has numerous applications in the food, pharmaceutical, and industrial fields. It is used to prevent the crystallization of sucrose in candies, as an acidulant in carbonated drinks, buffering agent, pH stabilizer in fruit and vegetable juices, and antioxidant as preservatives (Grewal and Kalra 1995 ; Magnuson and Lasure 2004 ; Carocho et al. 2018 ). Commercial citric acid production was initially done using A. niger in surface fermentation in 1916 by James Currie (Grewal and Kalra 1995 ; Show et al. 2015 ). Over the years, a variety of other microorganisms were studied, but A. niger is still superior among them in terms of production yield (Show et al. 2015 ). Citric acid production using this filamentous fungus is a multi-billion dollar industry (Cairns et al. 2018 ). Citric acid market was valued at USD 2.81 807 billion in 2021 and reached to USD 2.89 billion in 2022 due to increased demand (Future Market Insights 2022a ).

Fumaric acid is an unsaturated dicarboxyclic acid with various industrial applications in the food industry as a food acidulant in beverages and baking powders, feedstock for the synthesis of polymeric resins, and renewable energy resources (Yang et al. 2011 ). Fumaric acid is produced from glucose fermentation by the filamentous fungus Rhizopus arrhizus via a reductive tricarboxylic acid (TCA) pathway (Pan et al. 2016 ; Martin-Dominguez et al. 2022 ) and had an estimated market value of USD 654.4 million in 2020 (Reportlinker 2022 ).

Gluconic acid was first isolated from a strain of A. niger in 1922 (Goldberg and Rokem 2009 ). The production of gluconic acid can be done on an industrial scale using the enzymes glucose oxidase and catalase, derived from A. niger, which can efficiently convert glucose to gluconic acid (Copetti 2019 ). Other filamentous fungi, such as Gonatobotrys, Gliocladium, Penicillium and Scopulariopsis were found to produce gluconic acid (Goldberg and Rokem 2009 ; Dowdells et al. 2010 ). Gluconic acid and its derivatives have various applications in pharmaceuticals, cosmetics, and food products as additive or buffer salts (National Center for Biotechnology Information 2022b ; Yadav et al. 2022 ). The market size of gluconic acid was more than USD 50 million in 2017(Global Market Insights 2022b ).

Itaconic acid is a dicarboxylic acid with an α,β-unsaturated functionality that can be used as a monomer for the production of resins, plastics, paints, and synthetic fibers (Steiger et al. 2013 ). Aspergillus species such as A. itaconicus and A. terreus have the ability to synthesize itaconic acid (Steiger et al. 2013 ) as well as Ustilago maydis (Wierckx et al. 2020 ). The value of the itaconic acid market was USD 98.4 million in 2021 (Market Data Forecast 2022c ).

Lactic acid is also one of the most common organic acids produced from renewable substrates using bacteria and fungi such as Rhizopus species (Zhang et al. 2007 ). Lactic acid became also interesting due to its high added value (Castillo Martinez et al. 2013 ). It has various applications in the food and food-related industries (Ameen and Caruso 2017 ). Other potential applications are the production of biobased itaconic acid substitutes for petrochemical derivatives, such as biodegradable and biocompatible polylactate polymers (Zhang et al. 2007 ; Teleky and Vodnar 2019 ). Lactic acid market value was estimated to be USD 1.1 billion in 2020 (Markets and Markets 2022a ).

Fungi-based protein also called mycoprotein, is a sustainable protein source produced from single-cell protein (SCP) production technologies that could address the demands of protein worldwide (Nigam and Singh 2014 ). The global demand on vegetarian opportunities is currently rising, so the market share of mycoprotein is bound to increase. Moreover, mycoprotein consumption has been associated with numerous bioactivities such as antioxidant, antidiabetic, reduced cholesterol level, and fungal proteins are a potential source of amino acids that could induce muscle protein synthesis (Derbyshire and Delange 2021 ; Stoffel et al. 2021 ). Formerly, mycoproteins were produced for animal consumption, but the market for human feed is constantly growing (Nigam and Singh 2014 ; Derbyshire 2022 ). The commercially marketed Quorn™ mycoprotein is derived from Fusarium venenatum (Nigam and Singh 2014 ), which remains the most important source. Pleurotus albidus can also be used to produce mycoproteins flour that was used to replace wheat flour to produce cookies (Stoffel et al. 2021 ). The company “Nature Fynd” promotes Fy produced by a strain of Fusarium flavolapis , collected in the Yellowstone National Park, as an alternative producer. Due to the numerous food applications of mycoprotein, it was valued at USD 156.6 million in 2020 (Allied Market Research 2022e ) and is estimated to attain a market worth of USD 397.5 million in 2029 (Globe Newswire 2022b ) (Table 3 ).

Food enhancers/staple food

Soy sauce is the oldest food flavoring ingredient that is eminent, particularly in Asian countries (Hong et al. 2015 ). It is an all-purpose seasoning used in around 100 countries around the world (Ito and Matsuyama 2021 ). Fungi responsible for brewing soy sauce are known as koji molds and belong to the genus Aspergillus (e.g. A. flavus var. oryzae , A. sojae, and A. tamarii ) (Terada et al. 1981 ). Soy sauce is produced by fermenting soybean, wheat, and salt by the action of koji mold, halophilic lactic acid bacteria, and salt-tolerant yeasts such as Candida etchellsii , C. versatilis, and Zygosaccharomyces rouxii (Ito and Matsuyama 2021 ). The market value of soy sauce was around USD 39.5 billion in 2019 (Verified Market Research 2022b ).

Miso is another condiment, a fermented soybean paste made of Aspergillus species that are involved in making soy sauce, specifically A. flavus var. oryzae and A. sojae (Yokotsuka and Sasaki 1998 ; Allwood et al. 2021 ). Other fungi involved in accelerating miso ripening process are Zygosaccharomyces spp. together with the lactic acid bacterium, Pediococcus halophilus (Wilson 1995 ; Batt and Tortorello 2014; Allwood et al. 2021 ). There are several kinds of miso products depending on the type of koji used in fermentation (e.g. rice, barley, soy bean) (Wilson 1995 ; Allwood et al. 2021 ). The market size for miso was valued at USD 67.8 million in 2022 (Future Market Insights 2022b ).

Indonesian tempeh is a mold-facilitated fermented product produced from fermented soy beans mostly consumed as stable food in Indonesia (Dinesh Babu et al. 2009 ), but is widely accepted globally (Ahnan‐Winarno et al. 2021 ). The fungi involved in soybean fermentation for tempeh production are Rhizopus species such as R. microporus (formerly R. oligosporus ), and R. arrhizus (formerly R. oryzae ) (Maheshwari et al. 2021 ; Sjamsuridzal et al. 2021 ). The mycelia of the Rhizopus species hold the soy beans together, thus forming a compact structure like a cake (Gandjar 1999 ; Handoyo and Morita 2006 ). The global tempeh market was valued at USD 4.53 billion in 2021 (The Business Research Company 2022 ).

Food colorants are pigments added to food to enhance quality and add color to the food to make it more presentable and attractive to consumers (Hyde et al. 2019 ). Natural colors are generally regarded safer than synthetic ones and some may even have medicinal benefits. Furthermore, food colorants are also added to preserve food products (Christiana 2016 ). Nevertheless, the compounds should undergo extensive toxicity tests before they can be added to food, as some fungal toxins are also pigments. With the growing demand for natural food colorants globally, filamentous fungi have become important and readily available sources of food coloring (Dufossé et al. 2014 ; Hyde et al. 2019 ). Several genera of filamentous fungi such as Fusarium , Monascus, Penicillium, and Talaromyces have been explored for colorant production (Caro et al. 2017 ; Lagashetti et al. 2019 ). Marine fungi are also potential sources of pigments with various color hues and atypical chemical structures (Dufossé et al. 2014 ). Filamentous fungi can produce an extraordinary range of pigments, including some chemical classes such as carotenoids, melanins, flavins, phenazines, quinones, azaphilones, and violacein or indigo (Dufossé et al. 2014 ; Poorniammal et al. 2021 ). For instance, the strain Penicillium oxalicum var. armeniaca CCM 8242 isolated from soil was able to produce the first commercial red color Arpink red™ pigment (Natural red™) (Caro et al. 2017 ). The high demand for natural food colorants was reflected in its global market value of USD 1.6 billion in 2020 (Mordor Intelligence 2022 ) (Table 4 ).

Astaxanthin is a red–orange carotenoid belonging to the xanthophyll family (Aneesh et al. 2022 ). The natural sources of astaxanthin include bacteria, fungi, algae, crustaceans, and certain fishes. Important fungal astaxanthin producers are the yeasts Phaffia rhodozyma (Pandey et al. 2015 ) and Xanthophyllomyces dendrorhous (Rodríguez-Sáiz et al. 2010 ). The market price of astaxanthin ranges from USD 2500 to USD 7000/kg with the global market value of USD 1.94 billion in 2022 (Grand View Research 2023a ). However, we did not find a way to discriminate between fungal and non-fungal sources during our literature search.

Carotenoids are terpenoid pigments commonly produced by vascular plants, algae, bacteria, and fungi (Avalos and Carmen Limón 2015 ). Various fungi produced various carotenoids. Among these carotenoid-producing fungi are Blakeslea trispora (zygomycete) , Neurospora crassa (sordariomycete) and Xanthophyllomyces dendrorhous (basidiomycetous yeast) (Avalos and Carmen Limón 2015 ; Sandmann 2022 ). The provitamin β-carotene has also been described in Aspergillus giganteus , Penicillium sp., Rhodosporidium sp., Sclerotium rolfsii , Sclerotinia sclerotiorum , and Sporidiobolus pararoseus. Aside from the food market, there are potential applications for carotenoids in the textile industries (Venil et al. 2020 ). The global market size for carotenoids was USD 1.4 billion in 2019 (Fortune Business Insight 2022c ), but once again, the share of fungal products remains obscure.

Monascus pigments are natural food colorants widely used in the food industries worldwide, but especially in China and Japan (Feng et al. 2012 ). They are produced by fungi belonging to the genus Monascus . The six main pigments produced by strains of the genus Monascus are orange (monascorubrine, rubropunctatin), red (monascorubramine, rubropuntantamine), and yellow (monascine, ankaflavin) (Agboyibor et al. 2018 ). Furthermore, Monascus pigments have a variety of biological activities, such as antimutagenic, anticancer properties, antimicrobial activities, and potential anti-obesity activities. Monascus pigments already had a market value of USD 1 million in Japan at the end of the 1990s (Dufossé 2019 ), but we did not find hard data on the current market (Fig. 6 ).

Chemical structures of Monascus pigments

Another application of natural dyes from fungi is the textile industry (Chadni et al. 2017 ). Some wood rotting fungi are utilized in the production of textile dyes (Hernández et al. 2019 ). Other filamentous fungi can also produce various pigments with application in dyeing cotton, silk, and wool (Sudha et al. 2016; Kalra et al. 2020 ). Neurospora spp. are ascomycetous yeasts that can also be a great source of pigments (Lagashetti et al. 2019 ; Kalra et al. 2020 ).

Agarwood is a dark resinous heartwood that forms in mainly Aquilaria and Gyrinops trees naturally infected with endophytic fungi such as Cladosporium, Fusarium, Melanospora spp ., and “Mycelia sterilia” (Hyde et al. 2019 ), however, fungi reported have not been identified accurately according to the current standards. Agarwood is used as a fragrance in cosmetics and some important religious rituals (Chowdhury et al. 2016 ). The global market value of the agarwood chip market is worth USD 8.30 billion in 2018 (Straits Research 2021 ). Agarwood oil also has applications in the chemical, cosmetics, and personal care product industries with a global market value of USD 278.03 million in 2021 (Maximize Market Research 2021 ) (Tables 6 , 7 ).

Fungi synthesize natural active biomolecules that can be applied to develop cosmetic products due to their important biological functions and applications such as antiaging, antioxidants, skin revitalization, skin whitening, and hair products (Hyde et al. 2010 ; Wu et al. 2016 ; Badalyan et al. 2022 ). Numerous bioactive metabolites such as polysaccharides and glycoproteins, as well as other metabolites such as phenolic compounds, terpenoids, and several lipid components, have been explored due to their functions in skin health (Usman et al. 2021 ).

Lentinula edodes and Ganoderma species are two of the most popular fungal sources used in skin care products. Lentinan extracted from L. edodes has the potential to protect the skin against environmental pollutants (Zi et al. 2020 ). Agaricus subrufescens residues were also used as cosmeceutical ingredients (Hyde et al. 2010 ; Wisitrassameewong et al. 2012 ). Other macrofungal species utilized in skin care preparations include Taiwanofungus camphoratus, Grifola frondosa, Inonotus obliquus, Ophiocordyceps sinensis , and Schizophyllum commune (Wu et al. 2016 ). The following compounds are economically important isolated from fungi.

β-Glucans are polysaccharides that constitute around 30–80% of the fungal cell wall (Free 2013 ). Among these β-glucans, lentinan had a market value of USD 10 million in 2022 (Market Watch 2023b ). However, yeast is the main source of β-glucans in the market with a global value of USD 174.2 million in 2021 (Grand View Research 2023b ). The global β-glucan market was over USD 329.4 million in 2019 (Global Market Insights 2022c ) and increased to USD 403.8 million in 2020 (Markets and Markets 2022b ).

Ergothioneine is a natural sulphur derivative of histidine, which was originally isolated from the ascomycete Claviceps purpurea (Tanret 1909 ). It has antioxidant activity and is nowadays more commonly produced from edible mushrooms (Fu and Shen 2022 ). Some sources of ergothioneine are A. bisporus, Boletus edulis Flammulina “ velutipes ” (probably F. filiformis according to the current taxonomy established by Wang et al. 2018), Pleurotus eryngii, P. ostreatus and Lentinula edodes (Lee et al. 2009 ; Kalač 2016 ). The global market value of l -ergothioneine was around USD 15.02 million in 2021 (Market Data Forecast 2022d ).

Kojic acid , a pyrone derivative, is a byproduct in the fermentation process of producing sake, an alcoholic drink made from rice and almost exclusively produced in Japan (Goldberg and Rokem 2009 ). It is a metabolite of Aspergillus species, including A. oryzae and A. flavus. Kojic acid inhibits tyrosinase in the synthesis of melanin and is documented for its numerous bioactivities (Saeedi et al. 2019 ). It is known for its radioprotective and skin lightening effect and therefore has been used as ingredients in the production of skin creams, lotions, soaps and dental care products (Saeedi et al. 2019 ; Phasha et al. 2022 ). Kojic acid has a global market value of USD 36 million in 2022 (Market Watch 2022c ).

Fungi have been used as biological control agents of crop pests in agricultural and urban areas for several decades. (Wraight and Carruthers 1998). Almost 50% of the registered microbial biopesticides are based on fungi (Mascarin et al. 2018). Mycopesticides have been on the market since 1981, and the first of these products was registered in the United States as Mycar. It was composed of Hirsutella thompsonii , which causes epizootics in some species of spider mites (Rechcigl and Rechcigl 2000). Mycoinsecticides are interesting organic alternatives to chemical insecticides because they tend to be less harmful to human health and the environment. They are also target-specific with less detrimental effects to human and pets (Moore et al. 2011).

The most explored entomopathogenic fungal genera are Akanthomyces, Beauveria, and Metarhizium, which can infect a wide variety of insect hosts such as dipterans, lepidopterans, and coccidae. Numerous studies have explored the efficacy of these genera as biocontrol agents. Beauveria bassiana is one of the most explored entomopathogenic species worldwide (Fancelli et al. 2013 ). It has high genetic variability, virulence and adaptability to local conditions (Fancelli et al. 2013 ). Metarhizium, on the other hand, has been widely studied because of its general safety, narrow host range, environmentally friendliness and ease of mass production (Aw and Hue 2017). The global biopesticides market is estimated to be valued at $ 5.5 billion in 2022 (Research and Markets 2022b ). Mycopesticides account for around 10% of the global market value of biopesticides (Zaki et al. 2020 ), thus the global market value of mycopesticides can be estimated to be approximately USD 550 million in 2022.

Strobilurins are economically important bioactive compounds first reported by Anke et al. ( 1977 ) from cultures of the basidiomycete Strobilurus tenacellus. These compounds have later also been extracted from numerous Basidiomycota such as Oudemamsiella canari and Mucidula mucida (Rosa et al. 2003 , 2005 ; Iqbal et al. 2018 ). Strobilurins became an inspiration in developing the β-methoxyacrylate class of agricultural fungicides (Nofiani et al. 2018 ) such as azoxystrobin (Quadris), trifloxystrobin (Flint), pyraclostrobin (Cabrio), or Pristine (pyraclostrobin and boscalid, a premix of a Group 11 and a Group 7 fungicide). The global market value for strobilurin fungicides was estimated to be USD 4.65 billion in 2022 (Market Watch 2022d ).

Mycorrhizal fungi such as arbuscular mycorrhiza and ectomycorrhiza are considered natural biofertilizers since they provide important nutrients to the plant in exchange for photosynthetic products (Berruti et al. 2016 ; Domínguez-Núñez et al. 2020 ). A mycorrhiza-based biofertilizer is a substrate containing fungi colonizing the rhizosphere or the interior of the plant when applied to seeds, plant surfaces, or soil (Hyde et al. 2019 ; Karima and Samia 2020 ). It helps in plant growth by enhancing the supply of primary nutrients to the host plant and promoting growth hormones (Berruti et al. 2016 ).

The most widespread EcM product inoculum is Pisolithus tinctorius with a wide host range that can be applied as a vegetative mycelium peat vermiculite carrier used in nursery and forestry plantations (Gentili and Jumpponen 2006 ; Sebastiana et al. 2018 ). Other fungi used are Rhizophagus (formerly Glomus ), Sebacinales and Trichoderma species (Kaewchai et al. 2009 ; Molla et al. 2012 ). The market value of mycorrhiza-based biofertilizers was USD 271.8 million in 2021 (Market Data Forecast 2022e ).

Fungi grow ubiquitously on complex organic matter, prone to recycle nutrients and decompose complex substrates. In order to successfully accomplish this endeavor, they are equipped with a great portfolio of enzymes. Compared to chemical reactions, enzymes, used as biocatalysts, are highly specific, have few side products, and show mild reaction conditions with a low environmental and toxicological impact. Besides highly specific ones, also enzymes with a broad substrate spectrum are present. In general enzymes are classified in six classes by the International Enzyme Commission: EC1 oxidoreductases, EC2 transferases, EC3 hydrolases, EC4 lyases, EC5 isomerases, and EC6 ligases. In 2019 fungal enzymes had an overall marked share of 50% on all enzymes available (Kango et al. 2019 ). Despite of the large share, solely few species, namely Aspergillus , Trichoderma , Rhizopus , and Penicillium, contribute significantly to the market (El-Gendi et al. 2021 ).

In the food processing industry, fungi are the primary source for enzymes. They are applied to influence for example food quality, shelf life, texture, color, aroma but may also simplify production steps and yields (Zhang et al. 2018 ). Due to their unique properties, they are also used in the pharmaceutical, chemical, textile, detergent, cosmetic, and flavor or fragrance industry (He et al. 2017 ; Haile and Ayele 2022 ; El-Gendi et al. 2021 ). Whereas in 1983 about 30 enzyme classes, half of fungal origin, were commercialized, in 2020, the industrial enzymes market was valued at USD 5.7 billion (PS Market Research 2022 ). Though no detailed information on product origins is available, several global acting companies for enzyme production use fungi as production organisms. Among them are Novozymes (Denmark), DSM (Netherlands), Chr. Hansen (Denmark), DuPont (USA), BASF (Germany), Advanced Enzymes (India), and AB Enzymes (Germany). Here, production processes are diverse. Besides cultivation of the native producer strain, also heterologous hosts are widely spread. Second helps to overcome to long cultivation times, process in homogenities and simplifies the overall process. However, in case of technical enzymes, partially purified enzyme mixtures decrease production costs significantly. In bioremediation, wastewater treatment and bioethanol production, the use of whole-cell bioconversion of non engineered strains in mixed cultures is preferred to enzyme preparations (El-Gendi et al. 2021 ). Continuous fermentation simplifies processes and keeps them at low costs. Modern bioinformatics pushes the identification of novel enzymes with unique properties. Furthermore, genome mining approaches are on the verge, which increases product portfolios drastically.

Oxidoreductases are catalyzing redox reactions, therefore are typical housekeeping enzymes i.e., involved in the glycolysis and amino acid metabolism. Despite the great potential and the formation of several products of industrial interest, oxidoreductases are nowadays solely used in niche applications (Gygli and van Berkel 2015 ). However, fungi have a large share of this market, especially in bioremediation applications and wastewater treatment (Verma et al. 2020 ). This is due to the fact that mainly white rot Basidiomycota grow on lignocellulolytic biomass hitherto secretes robust enzymes of high redox-potential for the substrate degradation (Martínez et al. 2018 ). Especially Laccases, monooxygenases, and dioxygenases are of commercial use (El-Gendi et al. 2021 ).

Laccases have a great potential to oxidize phenolic and non-phenolic substrates cofactor independently. They are commercially used in destaining processes, paper industry, medical applications, and for biosensors. Usually, fungi are applied in microbial consortia to degrade pollution i.e. Aspergillus consortia that are used in wastewater treatment to degrade heavy metals chromium and cadmium (Talukdar et al. 2020 ). The global market value for laccase was USD 3 million in 2021 (Business Research Insights 2022 ).

Reactions involving the transfer of functional groups are catalyzed by transferases. Especially enzymes, catalyzing the transformation of higher sugars, are of commercial use. Invertases are breaking down sucrose into fructose and glucose (Nadaroglu and Polat 2022 ). Fungi that produce invertases are Aspergillus, Penicillium , Rhizopus and yeasts such as Saccharomyces (Kulshrestha et al. 2013 ; Matei et al. 2017 ) . Invertase is prevalent in the food industry, especially in making confectioneries and baking as well as extending the shelf life of the food products (Infita 2022 ). Inverted sugar prevents crystallization in food products and increases sweetness of soft fillings like soft caramel (Verma et al. 2020 ). Moreover, fructosyltransferase activities are used to produce fructooligosaccharides (FOS) which are known for their health promoting effects. FOS are applied as an alternative sweetener with a viscous texture in food industry (Lateef et al. 2012 ). The overall invertase market was valued at USD 57.6 million in 2022 (Future Market Insights 2022c ).

Hydrolases can cleave substrates via the addition of water. They display the most extensively studied enzyme class with the largest potential for commercialization (Gurung et al. 2013 ). Especially fungal amylases, proteases, cellulases and lipases are widely commercialized, but also other enzymes of industrial interest are known.

Among them, α -amylases are randomly hydrolyzing α-1,4-glycosidic bonds converting complex carbohydrate to simple sugars (Akinfemiwa et al. 2022 ). This unique property allows cleavage of non-reduced sugar residues unlike other amylolytic enzymes (Gupta et al. 2003 ). α - Amylases have the largest share of enzyme preparations used in industry (Monteiro de Souza and De Oliveira e Magalhães 2010 ). The global market size was valued at USD 278.2 million in 2018 and expected to increase up to USD 353 million by 2026 (Fior Market 2022 ). Among others, they are applied in the food, detergent, pulp and paper, and textile industry to hydrolyze starch (Gupta et al. 2003 ). The main applications of α -amylases are in the production of bakery products and food processing i.e. for glucose production (Copetti 2019 ). Herein, fungal enzymes are preferred to other sources due to their GRAS-status (Gupta et al. 2003 ). In bakeries, they are applied since 1995 and lead to a higher dough volume, better color, softer doughs, higher toasting quality, and a softer crumb (Pritchard 1992 ). They are commercially produced by Aspergillus (e.g. A. awamori , A. niger and A. oryzae ), Penicillium and Rhizopus (Pandey et al. 2000 ; Copetti 2019 ).

Proteases hydrolyze peptide bonds of proteins into smaller polypeptides or single amino acids (Flores-Gallegos et al. 2019 ). Various fungi such as Aspergillus, Humicola, Mucor, Penicillium, Rhizopus, Thermoascus, Thermomyces , etc. can produce proteases (Monteiro de Souza et al. 2015 ). These enzymes are easier to purify than their bacterial counterparts (Vishwanatha et al. 2010 ). Proteases have various industrial applications in the food industry, as pharmaceuticals and in biomedical therapies (Monteiro de Souza et al. 2015 ; Agbowuro et al. 2018 ; Naeem et al. 2022 ). Fungal proteases are cost effective and have faster production compared to other microbial sources (Monteiro de Souza et al. 2015 ). The global market value of fungal proteases was valued at USD 1.3 billion in 2021 (Dataintelo 2022a ).

Increasing demand on biofuels opens the potential of cellulose as cost effective substrate. Degradation is catalyzed by cellulases . Apart from biofuels, cellulases are also applied in combination with pectinases to clarify juices and in biopolishing of textiles (Gurung et al. 2013 ). Because of its high product titers, exceeding 100 g/L in submerged processes, Trichoderma reesei is widely used in industry (Gupta et al. 2016 ). The global market value of cellulases was USD 1.62 billion in 2022 (Future Market Insights 2022d ).

Pectinases are catalyzing the hydrolysis of pectin and pectic substances. Pectinases have wide application in food industry especially in fruit juice extraction to increase pressing efficiency and to clarify the former turbid drink (Jayani et al. 2005 ). Enzyme treatment in combination with cellulases decreases filtration times by 50% (Blanco 1999 ). During textile production an enzymatic cocktail containing among others pectinases is applied to remove non-cellulolytic impurities from clothes, called biosourcing (Hoondal et al. 2002 ). Some of the most efficient fungal species in producing pectinases are Aspergillus awamori, A. niger, Mucor piriformis, Penicillium restrictum, Trichoderma viride, and Yarrowia lipolytica (Haile and Ayele 2022 ). About 25% of global share of food enzyme in 2005 sales pectinases with the market value of USD 1.4 billion in 2020 based on the global industrial market value of industrial enzymes (Jayani et al. 2005 ; PS Market Research 2022 ).

Lipases hydrolyze triacylglycerols to glycerol and fatty acids which is of commercial interest as biological laundry detergents, cosmetics additive, fine chemical production, paper pitching, leather defatting, wastewater treatment, and biodiesel production (El-Gendi et al. 2021 ). Fungal lipases stand out due to its alkaline and temperature stability (Singh and Mukhopadhyay 2012 ). In food production i.e. offered by Rohm and Haas Co. USA, lipases are used to improve cheese aroma (Singh and Mukhopadhyay 2012 ). Important sources for fungal lipases are for example Aspergillus , Penicillium , and Fusarium (El-Gendi et al. 2021 ). The global lipase market value in 2022 was USD 694 million (Future Market Insights 2022e ).

Galactosidases , specifically β -galactosidases isolated from bacteria, filamentous fungi and yeasts, are responsible to hydrolyze lactose (Saqib et al. 2017 ). Yeasts, such as Kluyveromyces lactis and Guehomyces pullulans , but also filamentous fungi like Aspergillus niger are commercially important sources of β -galactosidases for food processing, waste disposal, and prebiotic production (Hu et al. 2010 ; Saqib et al. 2017 ). This enzyme is also used in whey disposal, a byproduct of cheese industry (Karasova et al. 2002 ). Enzymatically treated waste streams are rich in ethanol and a sweet syrup that is used further in food production including bakery (Zhou and Chen 2001 ). Lactase is a β -galactosidase, with a special importance for the production of lactose-free milk and dairy products (Copetti 2019 ). This enzyme is industrially obtained from fungal sources such as Aspergillus , Kluyveromyces, Trichoderma, Penicillium, Rhizopus, and Fusarium (Mahoney 1985 ; Zhou and Chen 2001 ; Seyis and Aksoz 2004 ; Jesus and Guimarães 2021 ).. The lactases produced by filamentous fungi are more heat-stable compared with the ones originating yeasts (Copetti 2019 ). Whereas the global market of galactosidases was valued at USD 1.5 billion in 2021, lactase take about 12% of the overall market share with around USD 185.2 million in 2019 (Grand View Research 2022b ; Dataintelo 2022b ).

Lyases mediate elimination reactions. Among them is the pectin lyase of industrial interest, catalyzing the non-hydrolytic cleavage of pectin. As previously described for pectinases, the lyases are present in preparations for fruit and juice processing (Suhaimi et al. 2021 ). These enzymes are also applied in a cocktail with various hydrolases in wine making as well as paper production processes. As for pectinases fungal sources play an important role in production due to their unique enzyme properties, namely temperature and pH stability. Prominent sources are Aspergillus , Candida , Penicillium , Trichoderma , and some yeasts.

Isomerases catalyze rearrangement processes. The most notable is the isomerization of glucose to generate high fructose corn syrup (HFCS), which has various applications in the food sector owing to its sweetness. HFCS has a high solubility, low crystallization tendency, and a high freezing point depression, which makes its use in drinks and frozen desserts advantageous. Besides food, HFCS is used in the detergent, and pharmaceutical industries (Singh et al. 2017 ). Commercially available pectin lyases have been solely reported from Aspergillus .

Ligases catalyze the formation of new bonds. Despite the high importance in intercellular mechanisms (besides niche applications in diagnostics, and molecular biology), hardly any commercialization is reported. Solely yeasts were reported as promising producer strains.

Edible mushrooms

Wild and commercial fungi can be resources of low-calorie functional food and nutraceuticals (Hyde et al. 2019 ; Lu et al. 2020 ). The popularity among consumers is due to the pleasant and unique flavors and health benefits of these mushrooms (Barros et al. 2008 ; Ache et al. 2021 ). Fungi are rich in crude fibers, proteins, vitamins, and minerals; they contain low fat, calories and high quality carbohydrates (Bandara et al. 2017 ; Thakur 2020 ). Fresh fungi have a moisture content of about 90% while dry matter consists of carbohydrates (50–65%), proteins (19–35%), essential fatty acids (2–6%) with some vitamins and minerals (Rathore et al. 2017 ; Jacinto-Azevedo et al. 2021 ). Consumption of edible fungi promotes good health caused by synergistic effects of the bioactive compounds present such as β-glucans, ascorbic acid, lectins, unsaturated fatty acids, phenolic compounds, tocopherols, and carotenoids (Varghese et al. 2019 ). Cultivated (54%), medicinal (38%), and wild fungi (8%) are the three major components of the global mushroom industry (Royse et al. 2017 ).

The world production of mushrooms was 42,590 thousand tons from the 20 largest producing countries, mainly contributed by China (93.93%) in 2020 (Table 8 ) (FAOSTAT 2022a ). The mushroom and truffle production and yield in Asia was around 7 million tons and 3.2 million hg/ha yield, respectively, in 2018. China is the largest contributor with 6.7 million tons production (FAOSTAT 2022a ). The total value for each country was determined by multiplying the production by the 2020 mushroom price (USD/ton) (FAOSTAT 2022b ). The total value of mushroom was USD 45.42 billion in 2020. The value of mushrooms being traded for food which only include cultivable and wild edible fungi constitute to 62% of the mushroom production or USD 28.16 billion in 2020.

Cultivable fungi

Cultivated edible mushroom production is attributed to 54% of global production, amounting to USD 34 billion in 2013 (Royse et al. 2017 ). Around 60 species of fungi are commercially cultivated (Chang and Miles 2004a ). The most cultivated mushroom genera in global production are Agaricus bisporus , followed by “ Lentinus” (Lentinula) edodes , Pleurotus spp., and Flammulina “velutipes” (or F. filiformis ) (cf. Valverde et al. 2015 with the names corrected to the currently valid taxonomy). These genera accounted for 85% of global mushroom production in the world (Royse et al. 2017 ; Kant Raut 2019 ) . The market value of edible cultivated fungi in 2020 accounted to 54% of global production was USD 24.53 billion .

Wild edible fungi

Approximately 2000 species of wild fungi are edible and can be consumed safely (Li et al. 2021 ). About 470 species have medicinal values, and another 180 species have been attributed value in other activities, such as religious purposes. In Tanzania, for instance, macrofungal collectors were able to collect up to 1500 kg of fruiting bodies of wild fungi that can earn around USD 500 to 600 per season (Tibuhwa 2013 ; Chelala et al. 2014 ). The retailers were also able to sell 750 to 800 kg with seasonal earnings of USD 750 to 1000 (Chelala et al. 2014 ). Wild edible fungi are collected each year in several million tons, accounting for 8% of global macrofungal production (Royse et al. 2017 ), therefore it was accounted to around USD 3.63 billion in 2020 .

Facing the transition towards a sustainable economy based on reduction of non-renewable materials and minimization of waste, fungal biotechnology offers a fascinating opportunity with high innovation potential (Meyer et al. 2020 ). In the last two decades, fungal biomaterials attracted growing attention in academic and commercial fields. Transferring organic agricultural or industrial waste into new bio-based products, they constitute a new form of a cost-effective and low energy bio-fabrication (Jones et al. 2020 ). Fungal mycelium has the capacity to degrade lignocellulosic materials and form composite networks (Sun et al. 2019 ) with insulating, non-flammable or hydrophobic properties and an improved mechanical strength (Jones et al. 2020 ). The mycelium binder constituent interfaces a dispersed phase of agricultural residue (substrate filler) and functions as a load transfer medium between the typically fibrous agricultural residue within the composite in a manner similar to the matrix phase of a polymer composite (Jones et al. 2020 ). Produced as an eco-friendly alternative to petroleum-based products, mycelium-based materials are considered to play a pivotal role in construction, automotive, transportation, electronic, and fashion sectors as well as design or applied arts. They comprise a great opportunity for substitution of traditional materials, such as timber, steel, concrete, foams or leather might (Karana et al. 2018 ; Attias et al. 2020 ; Silverman et al. 2020 ; Stelzer et al. 2021 ). Across the diverse disciplines, potential applications for mycelium-based products are reaching from packaging materials, bricks, acoustic panels, insulation panels, wall panels, fireproof material to leather and clothes.

Consistent with an increasing number of patent documents and scientific publications since 2007, the field of mycelium-based materials is currently under rapid development (Cerimi et al. 2019 ; Sydor et al. 2021 ). Sydor et al. ( 2021 ) identified 153 patent applications and 55 patents granted until April 2021, demonstrating the immense attraction for commercialization. US-based companies, such as Ecovative (Ecovative Design LLC 2023 ) and MycoWorks (MycoWorks 2023 ), pioneer companies focusing on dehydrated mycelium materials for packaging, insulation, fire protection or the production of a fungal leather alternative, respectively, have become key players on the market during the last fifteen years. Another up-coming label based in the US and Netherlands is Bolt Threads. In 2018, nine years after their foundation, they have developed the sustainable leather-like material Mylo™ attracting the attention of serious partners like Adidas, Kering, Lululemon or Stella McCartney. Furthermore, the success of European startups indicates the surging demand on the market for economical and eco-friendly sustainable technologies (Jones et al. 2020 ; Bitting et al. 2022 ). Striking examples are MycoTex (NEFFA 2023 ), the winner in 2018 of the Global Change Award created by the H&M foundation, or Grown.Bio (GrownBio 2022 ), featured as the most innovative SME (small and medium-sized enterprise) of the Netherlands in 2021.

Industries like packaging, plastic, and fashion have shown great interest in eco-friendly products (Research and Markets 2022c ). This shift goes hand in hand with an increasing economic impact of mycelium-based products. According to market analyses, the global market for mycelium materials, including food and beverage products, is valued at USD 2.48 billion in 2020 (Research and Markets 2022c ). A year later, the global mycelium market has increased to USD 2.95 billion with North America having the highest share of the mycelium market in 2021, followed by Europe and the Asia Pacific (InsightAce Analytic 2021 ). Driven by an ongoing trend to a bio-based circular economy around the world, governmental regulations, such as the styrofoam and polystyrene bans in various nations, increasing R&D investments and further application opportunities (e.g. mycorrhizal fungal concentrate for organic farming and tree nurseries, health-conscious products), the demand for eco-friendly mycelium-based products is continuously increasing. In 2030, the mycelium market is expected to reach USD 5.49 billion, with a CAGR of 7.3% during a forecasted period of 2022–2030 (InsightAce Analytic 2021 ). One of the key factors in hampering the market growth is the conversion from a small-scale of commercial products into an industrial-size scale. Limiting factors can be depicted by the lack of infrastructure facilities and challenging scale-up processes. Applicable solutions are needed for deficiencies in scalability (e.g. long cycles of production, risk of contamination, complex multi-step manufacturing processes), reproducibility (e.g. missing levels of standardization), and automation (InsightAce Analytic 2021 ; Bitting et al. 2022 ). The monopoly of mycelium-related patents in the industry together with a disconnect between industry and academia is further strengthenig this dilemma and prevent the distribution of knowledge and progress (Bitting et al. 2022 ).

Recreational mushroom picking is another service provided by forest ecosystems in which people can also sell the fungi they collected at the local market (Martínez de Aragón et al. 2011 ; Cutler II et al. 2021 ). Mushroom pickers benefited both from the recreational experience and the product collected (Martínez de Aragón et al. 2011 ; Debnath et al. 2019 ). Public demand for recreational mushroom picking has increased in the past decades in the forest ecosystems, affecting forest land owners (Frutos et al. 2009 ). Many people pick mushrooms for recreational purposes and not for economic reasons; thus, environmental valuation methods allow the benefits of this recreation activity to be assessed. The most common method used to measure the monetary value of recreational mushroom picking is the travel cost method. This method is used to approximate the difference between the willingness to pay for the good of the consumer (demand function) and the actual spending (price) (Bockstael 2007 ).

Mushroom picking is an upward leisure activity in European forests (Schulp et al. 2014 ; Marini Govigli et al. 2019 ). Along with the economic growth of European countries, fungal picking has transformed its profile to that of a primarily recreational activity (Kotowski 2016 ). Most studies of mushroom picking activities are done in Spain (Table 4 ). It shows that Europeans enjoy mushroom foraging activities as leisure activities. For instance, in Spain many studies were conducted in forested areas using the travel cost method to estimate the amount paid for mushroom hunting, which ranged from 10 to 60 €/trip (10.5–63 USD/trip) (Frutos et al. 2009 ). The estimated recreational surplus in Catalonia, Spain was €39 (40.9 USD) per journey (Martínez de Aragón et al. 2011 ). The average recreational surplus in Spain is USD 33.0 per trip, with 23% of the adult population experiencing mushroom picking at least once a year (Martínez de Aragón et al. 2011 ). Spain has around 38.6 million adults (World Population Review 2022 ), which, multiplied to 0.23 is equal to 8.9 million adults, which are experiencing mushroom picking once a year. Each of these 8.9 million adults has a recreational surplus of USD 33.0 per trip, thus the total recreational surplus is USD 293 million per year in Spain.

With filamentous fungi and yeasts being veritable cellular factories, they are major contributors to biotechnology, providing virtuous outlets for the protection of the environment and health by breaking down various industrial waste.

The spent mushroom substrate (SMS) is composed of fungal mycelia, extracellular enzymes, and unused lignocellulosic substrates. Production of industrially important hydrolytic enzymes by fungi has various economic applications, which can be an inexpensive alternative energy source (Grujić et al. 2015 ). SMS is suitable in a fermentative process with easy and low-energy sugar conversion through various hydrolysis methods, such as that leading to the production of economically important bioethanol (Antunes et al. 2020 ). SMS use could be a strategy to improve yield and reduce the cost of the production process (Corrêa et al. 2016 ). It could also be incorporated as a substrate for biogas generation (Pérez-Chávez et al. 2019 ).

Biofuels produced from renewable lignocellulosic biomass are projected to meet increasing energy demands without increasing greenhouse gas emissions as do fossil fuels (Saini and Sharma 2021 ), with butanol as the most promising for its high energy density (Cascone 2008 ). The spent mushroom substrate was also used as a potential substrate for butanol production using biodiesel removal in situ (Antunes et al. 2020 ). The old macrofungal-growing substrates harbor bacteria that can convert cellulose to biobutanol. Biobutanol is an alternative fuel that can directly replace gasoline in engines (Zhen et al. 2020 ). However, traditional methods in producing biofuels require a lot of energy and water. They are costly and require complicated pretreatments (Saini and Sharma 2021 ).

The global biofuels market was valued at nearly USD 110 billion in 2021 and is projected to continue to increase until 2030 with a forecast value of USD 201.2 billion (Statista 2022a ).

Environmental pollutions, especially synthetic organic compounds such as xenobiotics, have become a major problem worldwide. These compounds are recalcitrant and many microorganisms cannot degrade them (Embrandiri et al. 2016 ). Fungi can act as remediation tools due to their ability to enzymatically convert different type of pollutants into less complex components, even including a wide range of xenobiotics (Kulshreshtha et al. 2014 ; Zhen et al. 2022 ). White-rot fungi are able to produce extracellular ligninolytic enzymes consisting of three groups: lignin peroxidase (LiP), manganese-dependent peroxidases (MnP) and laccases, which play important roles in the transformation and mineralization of organic contaminants (Wang et al. 2009 ; Bulkan et al. 2020 ).

In agriculture, contaminated soil with heavy metals has negative effects on crop quality and yield (Kumar et al. 2018). There are three general effective strategies involving fungi to address contaminated soil: biodegradation, bioconversion, and biosorption. Toxicity reduction by fungi depends on their ability to produce different enzymes to degrade pollutants. Fungi can degrade pollutants and transform them into simpler mineral constituents (Chugh et al. 2022 ). They can also convert waste and other pollutants into more useful forms. Fungi can also absorb heavy metals from substrates without production of secondary pollution produced, which could be a very effective method to reclaim polluted land (Prenafeta-Boldú et al. 2019 ). Fungi have the potential to accumulate heavy metals in their network of hyphae and convert them into their mineral constituents. Fungi can also degrade recalcitrant hydrocarbon contaminants. They can turn hydrocarbon pollutants present in the environment into an energy source, increasing the selection of the hydrocarbon-metabolizing fungal population (Prenafeta-Boldú et al. 2019 ). Some fungal species are effective degraders of high molecular weight polycyclic aromatic hydrocarbons (Al Farraj et al. 2020 ). Some genera of fungi used for degradation of polycyclic aromatic hydrocarbons and polycyclic aromatic hydrocarbons are Agaricus (Li et al. 2010 ), Armillaria (Hadibarata and Kristanti 2013 ), Ganoderma (Agrawal et al. 2018 ), Marasmiellus (Vieira et al. 2018 ), Phanerochaete (Wang et al. 2009 ), Pleurotus (Li et al. 2010 ; Hadibarata and Teh 2014 ), and Trametes (as Coriolus ) (Jang et al. 2009 ). However, these projects are all at experimental stages. Among the different potential fungi, only Phanerochaete chrysosporium has been explored in large-scale biodegradation programs. Although global bioremediation has a large market value of USD 12.38 billion in 2021 (Grand View Research 2023c ), contribution of fungi is yet to be determined.

Fungi have been utilized in different industries (Hyde et al. 2019 ). Fungal products and services have a great impact in the global economy. High-value products biosynthesized by fungi can generate millions of dollars, which can help the country’s economy. The valuation of fungi at the global level is exceptionally difficult, and, in this account, we do not claim to accurately estimate this. However, we have used our knowledge and the available literature to derive a starting figure that can be used and built upon openly. Without such a figure, the value of fungi will not be recognized and there will be no monetary value for these amazing organisms, extrapolate decisions from stakeholders to further protect and conserve the mycobiota. This is our first attempt to give monetary value to the contributions of fungi in global economy that focus on marketed products and services. Our evaluation is carried out to derive an internationally accepted figure. The most significant fungal products and services with great economic value are listed in Table 9 . Fungi can be primary source in making products such as medicines and pharmaceuticals, TCM, functional food and nutraceuticals, pigments, and others or they can be involved in the production process especially in fermentation. For instance, production of fermented food and beverages such as baked goods, cheese, alcoholic beverages (e.g. beer, wine, whiskey) and non-alcoholic beverages (e.g. chocolate and coffee) with high market value will not be possible without the activity of yeast and filamentous fungi in fermentation in combination with bacteria. Therefore, involvement of fungi in this important process is also considered as contribution of fungi to global economy.

Our attempt to give monetary value to the contributions of fungi includes the definite value of fungi itself (e.g. wild and cultivated mushrooms), product value from the products made from fungi or through the involvement in the production process, and traded value from fungal services such as carbon stock and recreational foraging. Table 10 is the summation of the market values of fungal products and services that are economically of great importance. The contribution of fungi to the global market through fungal products and services is estimated to be USD 54.59 trillion .

The traded value of fungal services accounted for 95.90% of the total estimated value of fungi, or USD 52.33 trillion (Table 11 ). However, because this is just a monetary estimate of services rendered by fungi and has not been in the actual market, it can only be considered potential market value and not actual market value that affect the global Gross Domestic Product (GDP). The actual market value is made up of definite and product values that are already considered market revenue, totaling USD 2.24 trillion . The GDP is USD 88.44 trillion (Statista 2022b ), thus fungal products constitute for 2.53% of the GDP .

Fungi are being exploited in the forest and other ecosystems for their food production, medicinal and pharmaceutical applications; hence, they represent a major economic source worldwide (Zotti et al. 2013 ). Fungal products are sustainable, which could also create a biobased circular economy, which aims to eliminate waste and transform the production and use of goods (Valavanidis 2018 ). The circular economy is estimated to offer up to USD 4.5 trillion in global economic benefits by 2030 (World Economic Forum 2022 ).

Some services offered by fungi are eminent, especially those with direct effect to humans such as food source, medicines, commodities, and cultural services. These services also have available market values and thus are quantifiable. However, ecosystem services provided by fungi and their role in ecosystem functions, are not recognized, especially those which belong to supporting and regulating services. These services are not generally considered within policy appraisal at present. They represent an area where a greater and more methodical focus would be very valuable. To recognize and improve their strategic importance, several conservation strategies are necessary (Zotti et al. 2013 ). Institutional environment moves towards regulating mycological resources, with estimating the value of this ecosystem service becoming a key tool for policy-makers and rural entrepreneurs (Marini Govigli et al. 2019 ).

A formal ecosystem assessment could provide the necessary information on the larger suite of ecosystem services contributed by fungi in the ecosystems and for beneficiaries to value these services. Smaller and more specific studies should also be performed for each service to capture the true values of fungi. The under-representation of fungal ecosystem services and functions in ecosystem assessments potentially limits the information available for decision-making about regional and global activities that impact forest ecosystems. The concept of ecosystem services was developed as part of the Millennium Ecosystem Assessment project in the early 2000s to analyze and quantify the true and total value of an ecosystem. Valuation of fungi is essential, as it will contribute toward better decision making in policy appraisals considering the full costs and benefits to the natural environment and human wellbeing, while providing policy development with new insights (DEFRA 2006 ). Monetary valuation, or monetarization, is used in translating measures of social and biophysical impacts into monetary units so that they can be compared against each other and against the costs and benefits already expressed in monetary units (Marquina et al. 2022 ). This is very helpful in assessing the global value of fungi. Humans tend to give value to expensive things because they enjoy the pleasure of price. Studies show that people tend to value expensive items over their cheaper or free counterparts (Schmidt et al. 2017 ). The global value of fungi in ecosystems is missing or is not recognized since we get them for free. No studies have been conducted yet to give the overall value of fungi. Under-representation of the services rendered by fungi can lead to an underestimation of the global significance of the contribution of fungi in forest ecosystems. Moreover, many potential applications of fungi in biotechnology and industry have not been fully explored. Fungi have the potential to generate money, especially in their role as a bioremediator, mycoinsecticide, mycofertilizer, and biofuel. Quantifying ecosystem services provided by fungi and giving them a monetary price can help appraise their value to be taken into account in policy making and development. Formal and comprehensive ecosystem assessment would require considerable investment but could substantially improve coordination between management bodies, such as legislators and beneficiaries that could help in the conservation of fungal resources both on the regional and global scale.

In this study, using data from a number of industries, we estimated the global market value of fungi at USD 54.57 trillion . We acknowledge this is the first attempt to evaluate fungi in monetary terms, and that future assessments building on our work will perhaps be able to improve on the accuracy of this value. We have had to assume the value of fungi for some key industries, especially the food and beverages industries, as the total value of the products produced in those industries was based on the assumption that the final products would not be able to be produced, without the contribution of fungi.

This study emphasizes that fungi have an enormous market value, having an undeniable impact on the global economy. Monetary valuation of global products and services by fungi as well as their roles in ecosystem functioning is crucial to drive policy regarding the conservation and industrialization of this natural resource. With the rapid advancement of new technologies and related industries, the market presence of fungi will only increase, and thus fungi are prone to unsuitable exploitation if not properly monitored and managed in the future. The huge financial value of fungi adds weight to the argument that landscapes need to be conserved in order to conserve the natural resources found within. We have only discovered a small percentage of the fungi found in nature; thus, it is highly probable that there are untold billions of dollars’ worth of fungal resources yet to be discovered, or lost if their habitats are destroyed.

Data availability

No data were generated for this study.

Abel EL (2001) The gin epidemic: much ado about what? Alcohol Alcohol 36:401–405. https://doi.org/10.1093/alcalc/36.5.401

Article CAS PubMed Google Scholar

Ache NT, Manju EB, Lawrence MN, Tonjock RK (2021) Nutrient and mineral components of wild edible mushrooms from the Kilum-Ijim forest, Cameroon. Afr J Food Sci 15:152–161. https://doi.org/10.5897/ajfs2021.2089

Article CAS Google Scholar

Agbowuro AA, Huston WM, Gamble AB, Tyndall JDA (2018) Proteases and protease inhibitors in infectious diseases. Med Res Rev 38:1295–1331. https://doi.org/10.1002/med.21475

Agboyibor C, Kong W-B, Chen D et al (2018) Monascus pigments production, composition, bioactivity and its application: a review. Biocatal Agric Biotechnol 16:433–447. https://doi.org/10.1016/j.bcab.2018.09.012

Article Google Scholar

Agrawal N, Verma P, Shahi SK (2018) Degradation of polycyclic aromatic hydrocarbons (phenanthrene and pyrene) by the ligninolytic fungi Ganoderma lucidum isolated from the hardwood stump. Bioresour Bioprocess 5:11. https://doi.org/10.1186/s40643-018-0197-5

Ahnan-Winarno AD, Cordeiro L, Winarno FG et al (2021) Tempeh: a semicentennial review on its health benefits, fermentation, safety, processing, sustainability, and affordability. Compr Rev Food Sci Food Saf 20:1717–1767. https://doi.org/10.1111/1541-4337.12710

Akinfemiwa O, Zubair M, Muniraj T (2022) Amylase. StatPearls Publishing, Treasure Island

Google Scholar

Al Farraj DA, Hadibarata T, Elshikh MS et al (2020) Biotransformation and degradation pathway of pyrene by filamentous soil fungus Trichoderma sp. F03. Water Air Soil Pollut 231:168. https://doi.org/10.1007/s11270-020-04514-0

Alcohol Rehab Guide (2022) Types of alcohol. https://www.alcoholrehabguide.org/alcohol/types/#:~:text=Vodka%2C a liquor usually made,ABV in the United States. accessed 5 Sep 2022

Allied Market Research (2022a) Reishi mushroom market. https://www.alliedmarketresearch.com/reishi-mushroom-market-A10352 . accessed 23 Aug 2022

Allied Market Research (2022b) Beer market. https://www.alliedmarketresearch.com/beer-market . accessed 2 Sep 2022

Allied Market Research (2022c) Gin market. https://www.alliedmarketresearch.com/gin-market-A11469#:~:text=The global gin market was, forecast period 2021 to 2028. accessed 5 Sep 2022

Allied Market Research (2022d) Whiskey market. https://www.alliedmarketresearch.com/whiskey-market-A06652#:~:text=The global whiskey market was,share in the whiskey market. accessed 5 Sep 2022

Allied Market Research (2022e) Fungal protein market by type. https://www.alliedmarketresearch.com/fungal-protein-market-A12366 . accessed 15 Dec 2022c

Allwood JG, Wakeling LT, Bean DC (2021) Fermentation and the microbial community of Japanese koji and miso: a review. J Food Sci 86:2194–2207. https://doi.org/10.1111/1750-3841.15773

Ameen SM, Caruso G (2017) Lactic acid in the food industry. Springer, Cham

Book Google Scholar

Aneesh PA, Ajeeshkumar KK, Lekshmi RGK et al (2022) Bioactivities of astaxanthin from natural sources, augmenting its biomedical potential: A review. Trends Food Sci Technol 125:81–90. https://doi.org/10.1016/j.tifs.2022.05.004

Anke T, Werle A, Bross M, Steglich W (1977) The strobilurins-new antifungal antibiotics from the basidiomycete Strobilurus tenacellus (Pers. Ex Fr.) Sing. J Antibiotics 30:806–810. https://doi.org/10.7164/antibiotics.30.806

Antunes F, Marçal S, Taofiq O et al (2020) Valorization of mushroom by-products as a source of value-added compounds and potential applications. Molecules 25:2672. https://doi.org/10.3390/molecules25112672

Article CAS PubMed PubMed Central Google Scholar

Asbel LE, Levison ME (2000) Cephalosporins, carbapenems, and monobactams. Infect Dis Clin North Am 14:435–447. https://doi.org/10.1016/S0891-5520(05)70256-7

Attias N, Danai O, Abitbol T et al (2020) Mycelium bio-composites in industrial design and architecture: comparative review and experimental analysis. J Clean Prod 246:119037. https://doi.org/10.1016/j.jclepro.2019.119037

Avalos J, Carmen Limón M (2015) Biological roles of fungal carotenoids. Curr Genet 61:309–324. https://doi.org/10.1007/s00294-014-0454-x

Badalyan SM, Rapior S (2021) The neurotrophic and neuroprotective potential of macrofungi. In: Agrawal DC, Dhanasekaran M (eds) Medicinal herbs and fungi. Springer, Singapore, pp 37–77

Chapter Google Scholar

Badalyan S, Badalyan S, Barkhudaryan A (2021) The cardioprotective properties of Agaricomycetes mushrooms growing in the territory of Armenia (review). Int J Med Mushrooms 23:21–31. https://doi.org/10.1615/IntJMedMushrooms.2021038280

Article PubMed Google Scholar

Badalyan SM, Barkhudaryan A, Rapior S (2022) Medicinal macrofungi as cosmeceuticals: a review. Int J Med Mushrooms 24:1–13. https://doi.org/10.1615/IntJMedMushrooms.2022043124

Bandara AR, Rapior S, Bhat DJ et al (2015) Polyporus umbellatus , an edible-medicinal cultivated mushroom with multiple developed health-care products as food, medicine and cosmetics: a review. Cryptogam Mycol 36:3–42. https://doi.org/10.7872/crym.v36.iss1.2015.3

Bandara AR, Karunarathna SC, Mortimer PE et al (2017) First successful domestication and determination of nutritional and antioxidant properties of the red ear mushroom Auricularia thailandica (Auriculariales, Basidiomycota). Mycol Prog 16:1029–1039. https://doi.org/10.1007/s11557-017-1344-7

Barros L, Cruz T, Baptista P et al (2008) Wild and commercial mushrooms as source of nutrients and nutraceuticals. Food Chem Toxicol 46:2742–2747. https://doi.org/10.1016/j.fct.2008.04.030

Batt CA, Lou TM (2014) Encyclopedia of food microbiology, 2nd edn. Academic Press, Massachusetts

BBC (2022) Brandy. https://www.bbc.co.uk/food/brandy . accessed 5 Sep 2022

Bentley R (2000) Mycophenolic acid: A one hundred year odyssey from antibiotic to immunosuppressant. Chem Rev 100:3801–3826. https://doi.org/10.1021/cr990097b

Benz F, Knüsel F, Nüesch J et al (1974) Stoffwechselprodukte von Mikroorganismen 143. Mitteilung. Echinocandin B, ein neuartiges Polypeptid-Antibioticum aus Aspergillus nidulans var. echinulatus  : Isolierung und Bausteine. Helv Chim Acta 57:2459–2477. https://doi.org/10.1002/hlca.19740570818

Berruti A, Lumini E, Balestrini R, Bianciotto V (2016) Arbuscular mycorrhizal fungi as natural biofertilizers: let’s benefit from past successes. Front Microbiol 6:1559. https://doi.org/10.3389/fmicb.2015.01559

Article PubMed PubMed Central Google Scholar

Bhalla TC, Savitri (2017) Yeasts and traditional fermented foods and beverages. In: Satyanarayana T, Kunze G (eds) Yeast diversity in human welfare. Springer, Singapore, pp 53–82

Bills GF, Gloer JB (2016) Biologically active secondary metabolites from the fungi. Microbiol Spectr 4:1–32. https://doi.org/10.1128/microbiolspec.FUNK-0009-2016

Bitting S, Derme T, Lee J et al (2022) Challenges and opportunities in scaling up architectural applications of mycelium-based materials with digital fabrication. Biomimetics 7:44. https://doi.org/10.3390/biomimetics7020044

Blanco P (1999) Production of pectic enzymes in yeasts. FEMS Microbiol Lett 175:1–9. https://doi.org/10.1016/S0378-1097(99)00090-7

Bockstael N (2007) Environmental and resource valuation with revealed preferences. Springer, Dordrecht

Brick J (2022) The world’s best rum comes from these countries. https://www.thrillist.com/drink/nation/best-rum-regions-worlds-best-rum . accessed 5 Sep 2022

Britannica (2022a) Types of beer. https://www.britannica.com/topic/beer/Types-of-beer . accessed 31 Aug 2022

Britannica (2022b) Learn how to make vodka. https://www.britannica.com/video/179764/Overview-vodka#:~:text=Traditionally%2C vodka is made from,the distillation process can begin. accessed 1 Sep 2022

Brown AG, Butterworth D, Cole M et al (1976) Naturally-occurring beta-lactamase inhibitors with antibacterial activity. J Antibiotics 29:668–669. https://doi.org/10.7164/antibiotics.29.668

Bulkan G, Ferreira JA, Taherzadeh MJ (2020) Removal of organic micro-pollutants using filamentous fungi. In: Varjani S, Pandey A, Tyagi RD, Nho HH, Larroche C (eds) Current developments in biotechnology and bioengineering. Elsevier, Amsterdam, pp 363–395

Business Research Insights (2022) Laccase market size, share, growth, and industry analysis by type. https://www.businessresearchinsights.com/market-reports/laccase-market-101221 . accessed 14 Feb 2023

Cairns TC, Nai C, Meyer V (2018) How a fungus shapes biotechnology: 100 years of Aspergillus niger research. Fungal Biol Biotechnol 5:13. https://doi.org/10.1186/s40694-018-0054-5

Caro Y, Venkatachalam M, Lebeau J et al (2017) Pigments and colorants from filamentous fungi. In: Mérillon JM, Ramawat KG (eds) Fungal metabolites. Springer, Cham, pp 499–568

Carocho M, Morales P, Ferreira ICFR (2018) Antioxidants: reviewing the chemistry, food applications, legislation and role as preservatives. Trends Food Sci Technol 71:107–120. https://doi.org/10.1016/j.tifs.2017.11.008

Carvalho Â (2016) Yeasts and chocolate, the new “BFFs.” https://yeastcell.eu/2016/02/16/yeasts-and-chocolate-the-new-bffs/ . accessed 5 Sep 2022

Cascone R (2008) Biobutanol-a replacement for bioethanol? Chem Eng Prog 104:S4–S9

CAS Google Scholar

Castillo Martinez FA, Balciunas EM, Salgado JM et al (2013) Lactic acid properties, applications and production: a review. Trends Food Sci Technol 30:70–83. https://doi.org/10.1016/j.tifs.2012.11.007

Cerimi K, Akkaya KC, Pohl C et al (2019) Fungi as source for new bio-based materials: a patent review. Fungal Biol Biotechnol 6:17. https://doi.org/10.1186/s40694-019-0080-y

Chadni Z, Rahaman MH, Jerin I et al (2017) Extraction and optimisation of red pigment production as secondary metabolites from Talaromyces verruculosus and its potential use in textile industries. Mycology 8:48–57. https://doi.org/10.1080/21501203.2017.1302013

Chang ST, Miles P (2004a) Pleurotus : a mushroom of broad adaptability. Mushrooms: cultivation, nutritional value, medicinal effect, and environmental impact, 2nd edn. CRC Press, Boca Raton, pp 315–325

Chang ST, Miles PG (2004b) Mushrooms: cultivation, nutritional value, medicinal effect, and environmental impact, 2nd edn. CRC Press, Boca Raton

ChEBI (2022) Lovastatin. http://www.ebi.ac.uk/chebi/searchId.do?chebiId=CHEBI:40303 . accessed 20 Aug 2022

Chelala BL, Chacha M, Matemu A (2014) Wild edible mushroom value chain for improved livelihoods in Southern Highlands of Tanzania. Am J Res Commun 2:1–14

Cheng T, Kolařík M, Quijada L, Stadler M (2022) A re-assessment of Taxomyces andreanae , the alleged taxol-producing fungus, using comparative genomics. IMA Fungus 13:17. https://doi.org/10.1186/s43008-022-00103-4

Chourasia R, Abedin MM, Chiring Phukon L et al (2021) Biotechnological approaches for the production of designer cheese with improved functionality. Compr Rev Food Sci Food Saf 20:960–979. https://doi.org/10.1111/1541-4337.12680

Chowdhury M, Hussain MD, Chung SO et al (2016) Agarwood manufacturing: a multidisciplinary opportunity for economy of Bangladesh-a review. Agric Eng Int CIGR J 18:171–178

Christiana NO (2016) Production of food colourants by filamentous fungi. African J Microbiol Res 10:960–971. https://doi.org/10.5897/AJMR2016.7904

Chugh M, Kumar L, Bharadvaja N (2022) Chapter 4-Fungal diversity in the bioremediation of toxic effluents. In: Rodriguez-Couto S, Shah M (eds) Development in wastewater treatment research and processes. Elsevier, Amsterdam, pp 61–88

Clemmensen KE, Bahr A, Ovaskainen O et al (2013) Roots and associated fungi drive long-term carbon sequestration in boreal forest. Science 339:1615–1618. https://doi.org/10.1126/science.1231923

Cohen N, Cohen J, Asatiani MD et al (2014) Chemical composition and nutritional and medicinal value of fruit bodies and submerged cultured mycelia of culinary-medicinal higher basidiomycetes mushrooms. Int J Med Mushrooms 16:273–291. https://doi.org/10.1615/IntJMedMushr.v16.i3.80

Colorado State University Extension (2023) Functional Foods for Health–9.391. https://extension.colostate.edu/topic-areas/nutrition-food-safety-health/functional-foods-for-health-9-391/#:~:text=The term functional foods is,chronic disease and condition development. accessed 8 May 2023

Conway J (2022) Vodka industry- statistics & facts. In: Statista. https://www.statista.com/topics/3741/vodka-industry/#:~:text=World market,9 liter cases in 2020. accessed 5 Sep 2022

Copetti MV (2019) Fungi as industrial producers of food ingredients. Curr Opin Food Sci 25:52–56. https://doi.org/10.1016/j.cofs.2019.02.006

Corrêa RCG, da Silva BP, Castoldi R et al (2016) Spent mushroom substrate of Pleurotus pulmonarius : a source of easily hydrolyzable lignocellulose. Folia Microbiol (praha) 61:439–448. https://doi.org/10.1007/s12223-016-0457-8

Cutler WD II, Bradshaw AJ, Dentinger BTM (2021) What’s for dinner this time?: DNA authentication of “wild mushrooms” in food products sold in the USA. PeerJ 9:e11747. https://doi.org/10.7717/peerj.11747

Data Bridge Market Research (2022) Global echinocandins market – Industry trends and forecast to 2029. https://www.databridgemarketresearch.com/reports/global-echinocandins-market . accessed 3 Feb 2023

Dataintelo (2022a) Global fungal protease market by type. https://dataintelo.com/report/global-fungal-protease-market/ . accessed 31 Oct 2022

Dataintelo (2022b) Global galactosidase market by type. https://dataintelo.com/report/global-galactosidase-market/ . accessed 29 Oct 2022

de Almeida Silva Vilela J, de Figueiredo Vilela L, Ramos CL, Schwan RF (2020) Physiological and genetic characterization of indigenous Saccharomyces cerevisiae for potential use in productions of fermented maize-based-beverages. Braz J Microbiol 51:1297–1307. https://doi.org/10.1007/s42770-020-00271-8

De Filippis F, Troise AD, Vitaglione P, Ercolini D (2018) Different temperatures select distinctive acetic acid bacteria species and promotes organic acids production during Kombucha tea fermentation. Food Microbiol 73:11–16. https://doi.org/10.1016/j.fm.2018.01.008

De Silva DD, Rapior S, Sudarman E et al (2013) Bioactive metabolites from macrofungi: ethnopharmacology, biological activities and chemistry. Fungal Divers 62:1–40. https://doi.org/10.1007/s13225-013-0265-2

Debnath S, Debnath B, Das P, Saha AK (2019) Review on an ethnomedicinal practices of wild mushrooms by the local tribes of India. J Appl Pharm Sci 9:144–156. https://doi.org/10.7324/JAPS.2019.90818

DEFRA (2006) An introductory guide to valuing ecosystem services. London

Demain AL, Sanchez S (2009) Microbial drug discovery: 80 years of progress. J Antibiotics 62:5–16. https://doi.org/10.1038/ja.2008.16

Derbyshire E (2022) Fungal-derived mycoprotein and health across the lifespan: a narrative review. J Fungi 8:653. https://doi.org/10.3390/jof8070653

Derbyshire EJ, Delange J (2021) Fungal protein – What is it and what is the health evidence? A systematic review focusing on mycoprotein. Front Sustain Food Syst 5:581682. https://doi.org/10.3389/fsufs.2021.581682

Diallo I, Morel S, Vitou M, et al (2020) Ergosterol and amino acids contents of culinary-medicinal Shiitake from various culture conditions. Proceedings 70:78. https://doi.org/10.3390/foods_2020-07702

Dinesh Babu P, Bhakyaraj R, Vidhyalakshmi R (2009) A low cost nutritious food “Tempeh”-a review. World J Dairy Food Sci 4:22–27

Domínguez-Núñez JA, Berrocal-Lobo M, Albanesi A (2020) Ectomycorrhizal fungi as biofertilizers in forestry. In: Mirmajlessi SM, Radhakrishnan R (eds) Biostimulants in plant science. IntechOpen, Rijeka

Dong C, Guo S, Wang W, Liu X (2015) Cordyceps industry in China. Mycology 6:121–129. https://doi.org/10.1080/21501203.2015.1043967

Dörsam S, Fesseler J, Gorte O et al (2017) Sustainable carbon sources for microbial organic acid production with filamentous fungi. Biotechnol Biofuels 10:242. https://doi.org/10.1186/s13068-017-0930-x

Dowdells C, Jones RL, Mattey M et al (2010) Gluconic acid production by Aspergillus terreus . Lett Appl Microbiol 51:252–257. https://doi.org/10.1111/j.1472-765X.2010.02890.x

Drugbank Online (2022a) Griseofulvin. https://go.drugbank.com/drugs/DB00400 . accessed 21 Aug 2022

Drugbank Online (2022b) Fingolimod. https://go.drugbank.com/drugs/DB08868 . Accessed 22 Aug 2022

Dufossé L (2019) Pigments, microbial. In: Schmidt T (ed) Encyclopedia of microbiology, 4th edn. Academic Press, Oxford, pp 579–594

Dufossé L, Fouillaud M, Caro Y et al (2014) Filamentous fungi are large-scale producers of pigments and colorants for the food industry. Curr Opin Biotechnol 26:56–61. https://doi.org/10.1016/j.copbio.2013.09.007

Ecovative Design LLC (2023) Ecovative design. https://www.ecovative.com/ . accessed 2 Feb 2023

Edy N, Barus HN, Finkeldey R, Polle A (2022) Host plant richness and environment in tropical forest transformation systems shape arbuscular mycorrhizal fungal richness. Front Plant Sci 13:1004097. https://doi.org/10.3389/fpls.2022.1004097

El-Gendi H, Saleh AK, Badierah R et al (2021) A comprehensive insight into fungal enzymes: structure, classification, and their role in mankind’s challenges. J Fungi 8:23. https://doi.org/10.3390/jof8010023

El Enshasy HA, Hatti-Kaul R (2013) Mushroom immunomodulators: unique molecules with unlimited applications. Trends Biotechnol 31:668–677. https://doi.org/10.1016/j.tibtech.2013.09.003

Embrandiri A, Kiyasudeen SK, Rupani PF, Ibrahim MH (2016) Environmental xenobiotics and its effects on natural ecosystem. In: Singh A, Prasad S, Singh R (eds) Plant responses to xenobiotics. Springer, Singapore, pp 1–18

Endo A (1979) Monacolin K, a new hypocholesterolemic agent produced by a Monascus species. J Antibiotics 32:852–854. https://doi.org/10.7164/antibiotics.32.852

Endo A (2008) A gift from nature: the birth of the statins. Nat Med 14:1050–1052. https://doi.org/10.1038/nm1008-1050

Endo A (2010) A historical perspective on the discovery of statins. Proc Jpn Acad Ser B Phys Biol Sci 86(5):484−463. https://doi.org/10.2183/pjab.86.484

Endo A, Kuroda M, Tsujita Y (1976) ML-236A, ML-236B, and ML-236C, new inhibitors of cholesterogenesis produced by Penicillium citrinium . J Antibiotics 29:1346–1348. https://doi.org/10.7164/antibiotics.29.1346

Espinel-Ingroff A, Dannaoui E (2021) Special issue: antifungal agents recently approved or under development. J Fungi 7:239. https://doi.org/10.3390/jof7030239

European Medicines Agency (2013) Fourth European surveillance of veterinary antimicrobial consumption (ESVAC) report. Sales of veterinary antimicrobial agents in 26 EU/EEA countries in 2012. Eur Med Agency, Amsterdam

FAO (2015) FAO coffee pocketbook. http://www.fao.org/3/a-i4985e.pdf . accessed 5 Sep 2022

FAOSTAT (2022a) Mushroom and truffles. https://www.fao.org/faostat/en/#search/Mushrooms and truffles. accessed 13 May 2022

FAOSTAT (2022b) Mushroom & truffles: Producer prices. https://www.fao.org/faostat/en/#search/mushroom . accessed 10 Jan 2023

Feng Y, Shao Y, Chen F (2012) Monascus pigments. Appl Microbiol Biotechnol 96:1421–1440. https://doi.org/10.1007/s00253-012-4504-3

Fernandes P (2016) Fusidic acid: A bacterial elongation factor inhibitor for the oral treatment of acute and chronic staphylococcal infections. Cold Spring Harb Perspect Med 6:a025437. https://doi.org/10.1101/cshperspect.a025437

Fior Market (2022) Alpha-amylase baking enzyme market by source. https://www.fiormarkets.com/report/alpha-amylase-baking-enzyme-market-by-source-fungi-bacteria-396099.html . accessed 29 Oct 2022

Flores-Gallegos AC, Delgado-García M, Ascacio-Valdés JA et al (2019) Hydrolases of halophilic origin with importance for the food industry. In: Kuddos M (ed) Enzymes in food biotechnology. Elsevier, Amsterdam, pp 197–219

Food Data Central (2019) Mushroom raw. https://fdc.nal.usda.gov/fdc-app.html#/food-details/169251/nutrients

Fortune Business Insight (2022a) Fermented food and drink market. https://www.fortunebusinessinsights.com/fermented-food-and-drinks-market-103255 . accessed 5 Sep 2022

Fortune Business Insight (2022b) Wine market. https://www.fortunebusinessinsights.com/wine-market-102836 . accessed 4 Sep 2022

Fortune Business Insight (2022c) Carotenoids market size, share & Covid-impact analysis, by type. https://www.fortunebusinessinsights.com/industry-reports/carotenoids-market-100180 . accessed 2 Nov 2022

Free SJ (2013) Fungal cell wall organization and biosynthesis. Adv Genet 81:33–82. https://doi.org/10.1016/B978-0-12-407677-8.00002-6

Frutos P, Martínez Peña F, Ortega Martínez P, Esteban S (2009) Estimating the social benefits of recreational harvesting of edible wild mushrooms using travel cost methods. For Syst 18:235. https://doi.org/10.5424/fs/2009183-01065

Fu TT, Shen L (2022) Ergothioneine as a natural antioxidant against oxidative stress-related diseases. Front Pharmacol 13:850813. https://doi.org/10.3389/fphar.2022.850813

Future Market Insights (2022a) Citric acid market. https://www.futuremarketinsights.com/reports/citric-acid-market . accessed 18 Sep 2022

Future Market Insights (2022b) Miso market. https://www.technavio.com/report/miso-market-industry-analysis . accessed 31 Oct 2022

Future Market Insights (2022c) Invertase market outlook (2022c-2032). https://www.futuremarketinsights.com/reports/invertase-market . accessed 29 Oct 2022

Future Market Insights (2022d) Cellulases market. https://www.futuremarketinsights.com/reports/cellulase-market . accessed 14 Feb 2023

Future Market Insights (2022e) Lipase market. https://www.futuremarketinsights.com/reports/lipase-market . accessed 14 Feb 2023

Galappaththi MCA, Patabendige NM, Premarathne BM et al (2022) A review of Ganoderma triterpenoids and their bioactivities. Biomolecules 13:24. https://doi.org/10.3390/biom13010024

Gandjar I (1999) Fermented foods | Fermentations of the far East. In: Robinson R (ed) Encyclopedia of food microbiology. Elsevier, Oxford, pp 767–773

Gao YG, Selmer M, Dunham CM et al (2009) The structure of the ribosome with elongation factor G trapped in the posttranslocational state. Science 326:694–699. https://doi.org/10.1126/science.1179709

Gentili F, Jumpponen A (2006) Potential and possible uses of bacterial and fungal biofertilizers. In: Rai M (ed) Handbook of microbial biofertilizers. CRC Press, Florida, pp 29–56

Ghannoum M, Arendrup MC, Chaturvedi VP et al (2020) Ibrexafungerp: A novel oral triterpenoid antifungal in development for the treatment of Candida auris infections. Antibiotics 9:539. https://doi.org/10.3390/antibiotics9090539

Gibson B, Geertman J-MA, Hittinger CT et al (2017) New yeasts—new brews: modern approaches to brewing yeast design and development. FEMS Yeast Res 17:fox038. https://doi.org/10.1093/femsyr/fox038

Global Market Insights (2022a) Onychomycosis treatment market size. https://www.gminsights.com/industry-analysis/onychomycosis-treatment-market . accessed 22 Aug 2022

Global Market Insights (2022b) Gluconic acid market. https://www.gminsights.com/industry-analysis/gluconic-acid-market . accessed 21 Sep 2022

Global Market Insights (2022c) Global beta glucan market. https://www.gminsights.com/industry-analysis/beta-glucans-market . accessed 13 Nov 2022

Globe Newswire (2022a) Brandy market growth segments 2022a. https://www.globenewswire.com/en/news-release/2022/01/31/2375510/0/en/Brandy-Market-Growth-Segments-2022-Trends-Regional-Analysis-Top-Players-Revenue-USD-25320-million-Forecast-by-2027-Report-by-Absolute-Reports.html#:~:text= The global Brandy market was,1. accessed 4 Sep 2022

Globe Newswire (2022b) Fungal protein market worth $397.5 million by 2029. https://www.globenewswire.com/en/news-release/2022b/06/22/2466987/0/en/Fungal-Protein-Market-Worth-397-5-Million-by-2029-Exclusive-Report-by-Meticulous-Research.html . accessed 11 Nov 2022

GlobeNewswire (2022) SCYNEXIS reports third quarter 2022 financial results and provides corporate update. https://www.globenewswire.com/en/news-release/2022/11/09/2551800/0/en/SCYNEXIS-Reports-Third-Quarter-2022-Financial-Results-and-Provides-Corporate-Update.html . accessed 3 Feb 2023

Godtfredsen WO, Jahnsen S, Lorck H et al (1962) Fusidic acid: a new antibiotic. Nature 193:987–987. https://doi.org/10.1038/193987a0

Goldberg I, Rokem JS (2009) Organic and fatty acid production, microbial. In: Schaechter M (ed) Encyclopedia of microbiology, 3rd edn. Academic Press, Cambridge, pp 421–442

Gopalakrishnan C, Pawar RS, Bhutani KK (2005) Development of Agaricus bisporus as a nutraceutical of tomorrow. Acta Hortic 680:45–47. https://doi.org/10.17660/ActaHortic.2005.680.5

Gosio B (1893) Sperimentate su culture pure di bacilli del carbonchio demonstrarano notevole potere antisettica. CR Acad Med Torino 61:484

Gosio B (1896) Richerche batteriologiche e chemiche sulle alterazoni del mais. Riv D’igiene Sanita Pub Ann 7:825–849

Graham C (2022e) What is tequila? https://www.thespruceeats.com/all-about-tequila-760706 . accessed 5 Sep 2022

Grand View Research (2022a) Chocolate market size, share & trends analysis report by product. https://www.grandviewresearch.com/industry-analysis/chocolate-market#:~:text=Report Overview,4.6%25 from 2020 to 2027. accessed 4 Sep 2022

Grand View Research (2022b) Lactase market. https://www.grandviewresearch.com/industry-analysis/lactase-market . accessed 29 Oct 2022

Grand View Research (2023a) GVR report cover astaxanthin market size, share & trends report astaxanthin market size, share & trends analysis report by product. https://www.grandviewresearch.com/industry-analysis/global-astaxanthin-market . accessed 14 Jan 2023

Grand View Research (2023b) Yeast beta-glucan market size, share & trends analysis report by application. https://www.grandviewresearch.com/industry-analysis/yeast-beta-glucan-market-report . accessed 19 Jan 2023

Grand View Research (2023c) Bioremediation market size, share & trends analysis report by type. https://www.grandviewresearch.com/industry-analysis/bioremediation-market-report . accessed 12 Jan 2023

Grewal HS, Kalra KL (1995) Fungal production of citric acid. Biotechnol Adv 13:209–234. https://doi.org/10.1016/0734-9750(95)00002-8

GrownBio (2022) 100% natural alternative to plastic foam. https://www.grown.bio/ . accessed 2 Feb 2023

Grubb M, Vrolijk C, Brack D (1999) The Kyoto protocol: a guide and assessment. Royal Institute for International Affairs, London

Grujić M, Dojnov B, Potočnik I et al (2015) Spent mushroom compost as substrate for the production of industrially important hydrolytic enzymes by fungi Trichoderma spp. and Aspergillus niger in solid state fermentation. Int Biodeterior Biodegradation 104:290–298. https://doi.org/10.1016/j.ibiod.2015.04.029

Gupta AK, Foley KA, Versteeg SG (2017) New antifungal agents and new formulations against dermatophytes. Mycopathologia 182:127–141. https://doi.org/10.1007/s11046-016-0045-0

Gupta R, Gigras P, Mohapatra H et al (2003) Microbial α-amylases: a biotechnological perspective. Process Biochem 38:1599–1616. https://doi.org/10.1016/S0032-9592(03)00053-0

Gupta S, Summuna B, Gupta M, Annepu SK (2018) Edible mushrooms: cultivation, bioactive molecules, and health benefits. In: Mérillon JM, Ramawat K (eds) Bioactive molecules in food. Reference series in phytochemistry. Springer, Cham, pp 1–33

Gupta VK, Kubicek CP, Berrin J-G et al (2016) Fungal enzymes for bio-products from sustainable and waste biomass. Trends Biochem Sci 41:633–645. https://doi.org/10.1016/j.tibs.2016.04.006

Gurung N, Ray S, Bose S, Rai V (2013) A broader view: Microbial enzymes and their relevance in industries, medicine, and beyond. Biomed Res Int 2013:329121. https://doi.org/10.1155/2013/329121

Gutiérrez-Ríos HG, Suárez-Quiroz ML, Hernández-Estrada ZJ et al (2022) Yeasts as producers of flavor precursors during cocoa bean fermentation and their relevance as starter cultures: a review. Fermentation 8:331. https://doi.org/10.3390/fermentation8070331

Gygli G, van Berkel JH (2015) Oxizymes for biotechnology. Curr Biotechnol 4:100–110. https://doi.org/10.2174/2211550104666150423202036

Hadibarata T, Kristanti RA (2013) Biodegradation and metabolite transformation of pyrene by basidiomycetes fungal isolate Armillaria sp. F022. Bioprocess Biosyst Eng 36:461–468. https://doi.org/10.1007/s00449-012-0803-4

Hadibarata T, Teh ZC (2014) Optimization of pyrene degradation by white-rot fungus Pleurotus pulmonarius F043 and characterization of its metabolites. Bioprocess Biosyst Eng 37:1679–1684. https://doi.org/10.1007/s00449-014-1140-6

Haile S, Ayele A (2022) Pectinase from microorganisms and its industrial applications. Sci World J 2022:1–15. https://doi.org/10.1155/2022/1881305

Hamwenye KKN, Ueitele ISE, Kadhila NP et al (2022) Towards medicinal tea from untapped Namibian Ganoderma : Phenolics and in vitro antioxidant activity of wild and cultivated mushrooms. S Afr J Sci 118:9357. https://doi.org/10.17159/sajs.2022/9357

Handoyo T, Morita N (2006) Structural and functional properties of fermented soybean (Tempeh) by using Rhizopus oligosporus . Int J Food Prop 9:347–355. https://doi.org/10.1080/10942910500224746

Harder A, Pham T, Jarling R, et al (2011) Method for producing optically active, cyclic depsipeptides comprising lactic acid and phenyl lactic and having 24 ring atoms, using fungus strains of Rosellinia type, and further species of Xylariaceae (Patent No. WO2010072323A1). European Patent Office13/133,611

He J (2018) Harvest and trade of caterpillar mushroom ( Ophiocordyceps sinensis ) and the implications for sustainable use in the Tibet Region of Southwest China. J Ethnopharmacol 221:86–90. https://doi.org/10.1016/j.jep.2018.04.022

He L, Mao Y, Zhang L et al (2017) Functional expression of a novel α-amylase from Antarctic psychrotolerant fungus for baking industry and its magnetic immobilization. BMC Biotechnol 17:22. https://doi.org/10.1186/s12896-017-0343-8

Helaly SE, Thongbai B, Stadler M (2018) Diversity of biologically active secondary metabolites from endophytic and saprotrophic fungi of the ascomycete order Xylariales. Nat Prod Rep 35:992–1014. https://doi.org/10.1039/C8NP00010G

Hernández VA, Galleguillos F, Thibaut R, Müller A (2019) Fungal dyes for textile applications: testing of industrial conditions for wool fabrics dyeing. J Text Inst 110:61–66. https://doi.org/10.1080/00405000.2018.1460037

Ho CW, Lazim AM, Fazry S et al (2017) Varieties, production, composition and health benefits of vinegars: a review. Food Chem 221:1621–1630. https://doi.org/10.1016/j.foodchem.2016.10.128

Hong S-B, Kim D-H, Samson RA (2015) Aspergillus associated with meju, a fermented soybean starting material for traditional soy sauce and soybean paste in Korea. Mycobiology 43:218–224. https://doi.org/10.5941/MYCO.2015.43.3.218

Hoondal G, Tiwari R, Tewari R et al (2002) Microbial alkaline pectinases and their industrial applications: a review. Appl Microbiol Biotechnol 59:409–418. https://doi.org/10.1007/s00253-002-1061-1

Houbraken J, Frisvad JC, Samson RA (2011) Fleming’s penicillin producing strain is not Penicillium chrysogenum but P. rubens . IMA Fungus 2:87–95. https://doi.org/10.5598/imafungus.2011.02.01.12

Hu X, Robin S, O’Connell S et al (2010) Engineering of a fungal beta-galactosidase to remove product inhibition by galactose. Appl Microbiol Biotechnol 87:1773–1782. https://doi.org/10.1007/s00253-010-2662-8

Hübner MP, Townson S, Gokool S et al (2021) Evaluation of the in vitro susceptibility of various filarial nematodes to emodepside. Int J Parasitol Drugs Drug Resist 17:27–35. https://doi.org/10.1016/j.ijpddr.2021.07.005

Hüttel W (2021) Echinocandins: structural diversity, biosynthesis, and development of antimycotics. Appl Microbiol Biotechnol 105:55–66. https://doi.org/10.1007/s00253-020-11022-y

Hyde KD, Bahkali AH, Moslem MA (2010) Fungi—an unusual source for cosmetics. Fungal Divers 43:1–9. https://doi.org/10.1007/s13225-010-0043-3

Hyde KD, Xu J, Rapior S et al (2019) The amazing potential of fungi: 50 ways we can exploit fungi industrially. Fungal Divers 97:1–136. https://doi.org/10.1007/s13225-019-00430-9

Hymery N, Vasseur V, Coton M et al (2014) Filamentous fungi and mycotoxins in cheese: a review. Compr Rev Food Sci Food Saf 13:437–456. https://doi.org/10.1111/1541-4337.12069

Imarc (2022a) Cephalosporin market: Global industry trends, share, size, growth, opportunity and forecast 2023–2028. https://www.imarcgroup.com/cephalosporin-market . accessed 3 Feb 2023

Imarc (2022b) Statin market: Global industry trends, share, size, growth, opportunity and forecast 2022b-2027. https://www.imarcgroup.com/statin-market . accessed 3 Feb 2023

Imarc (2022c) Global vinegar market outlook 2022c- 2027. https://www.imarcgroup.com/vinegar-manufacturing-plant#:~:text=The global vinegar market was,US%24 2.27 Billion in 2021. accessed 5 Sep 2022

Imarc (2023) Bakery products market: Global industry trends, share, size, growth, opportunity and forecast 2023–2028. https://www.imarcgroup.com/bakery-products-market#:~:text=The global bakery products market,3.7%25 during 2023–2028. accessed 25 Jan 2023

Infita (2022) Uses of invertase enzymes. https://infinitabiotech.com/blog/uses-of-invertase-enzyme/#:~:text=Invertase is one such enzyme,to extend their shelf life. accessed 29 Oct 2022

InsightAce Analytic (2021) Global mycelium market. https://www.insightaceanalytic.com/report/global-mycelium-market-/1219 . accessed 21 Jun 2022

International Trade Center (2022) Trade map. https://www.trademap.org/Index.aspx . accessed 5 Sep 2022

IPCC (2005) Chapter 9: Implications of carbon dioxide capture and storage for greenhouse gas inventories and accounting. In: Hiraishi T, Miguez JD (eds) IPCC special report on carbon dioxide capture and storage. pp 363–379

Iqbal Z, Han LC, Soares-Sello AM et al (2018) Investigations into the biosynthesis of the antifungal strobilurins. Org Biomol Chem 16:5524–5532. https://doi.org/10.1039/c8ob00608c

Ito K, Matsuyama A (2021) Koji molds for Japanese soy sauce brewing: characteristics and key enzymes. J Fungi 7:658. https://doi.org/10.3390/jof7080658

Iwamoto T, Fujie A, Nitta K et al (1994) WF11899A, B and C, novel antifungal lipopeptides. II. Biological properties. J Antibiotics 47:1092–1097. https://doi.org/10.7164/antibiotics.47.1092

Jacinto-Azevedo B, Valderrama N, Henríquez K et al (2021) Nutritional value and biological properties of Chilean wild and commercial edible mushrooms. Food Chem 356:129651. https://doi.org/10.1016/j.foodchem.2021.129651

Jain A, Utley L, Parr TR et al (2018) Tebipenem, the first oral carbapenem antibiotic. Expert Rev Anti Infect Ther 16:513–522. https://doi.org/10.1080/14787210.2018.1496821

Jallow S, Govender NP (2021) Ibrexafungerp: a first-in-class oral triterpenoid glucan synthase inhibitor. J Fungi 7:163. https://doi.org/10.3390/jof7030163

Jang K-Y, Cho S-M, Seok S-J et al (2009) Screening of biodegradable function of indigenous ligno-degrading mushroom using dyes. Mycobiology 37:53. https://doi.org/10.4489/MYCO.2009.37.1.053

Jayabalan R, Malbaša RV, Lončar ES et al (2014) A review on Kombucha tea—microbiology, composition, fermentation, beneficial effects, toxicity, and tea fungus. Compr Rev Food Sci Food Saf 13:538–550. https://doi.org/10.1111/1541-4337.12073

Jayani RS, Saxena S, Gupta R (2005) Microbial pectinolytic enzymes: a review. Process Biochem 40:2931–2944. https://doi.org/10.1016/j.procbio.2005.03.026

Jesus LFMC, Guimarães LHS (2021) Production of β-galactosidase by Trichoderma sp. through solid-state fermentation targeting the recovery of galactooligosaccharides from whey cheese. J Appl Microbiol 130:865–877. https://doi.org/10.1111/jam.14805

Jiang W, Hu J-W, He X-R et al (2021) Statins: a repurposed drug to fight cancer. J Exp Clin Cancer Res 40:241. https://doi.org/10.1186/s13046-021-02041-2

Johnston RJ, Wainger LA (2015) Benefit transfer for ecosystem service valuation: an introduction to theory and methods. In: Johnston RJ, Rolfe J, Rosenberger RS, Brouwer R (eds) Benefit transfer of environmental and resource values. Springer, Dordrecht, pp 237–273

Jones M, Mautner A, Luenco S et al (2020) Engineered mycelium composite construction materials from fungal biorefineries: a critical review. Mater Des 187:108397. https://doi.org/10.1016/j.matdes.2019.108397

Joseph S (2019) Nabriva prices two pneumonia antibiotic versions at over $200 per day. https://www.reuters.com/article/us-nabriva-fda-idUSKCN1V91QQ . accessed 13 Feb 2023

Kaewchai S, Soytong K, Hyde KD (2009) Mycofungicides and fungal biofertilizers. Fungal Divers 38:25–50

Kahan JS, Kahan FM, Goegelman R et al (1979) Thienamycin, a new beta-lactam antibiotic I. Discovery, taxonomy, isolation and physical properties. J Antibiotics 32:1–12. https://doi.org/10.7164/antibiotics.32.1

Kalač P (2016) Health-stimulating compounds and effects. In: Kalač P (ed) Edible mushrooms. Elsevier, Amsterdam, pp 137–153

Kalra R, Conlan XA, Goel M (2020) Fungi as a potential source of pigments: harnessing filamentous fungi. Front Chem 8:369. https://doi.org/10.3389/fchem.2020.00369

Kango N, Jana UK, Choukade R (2019) Fungal enzymes: sources and biotechnological applications. In: Satyanarayana T, Deshmukh SK, Deshpande MV (eds) Advancing frontiers in mycology & mycotechnology. Springer, Singapore, pp 515–538

Kanny D, Brewer RD, Mesnick JB et al (2015) Vital signs: alcohol poisoning deaths-United States, 2010–2012. MMWR Morb Mortal Wkly Rep 63:1238–1242

PubMed PubMed Central Google Scholar

Kant Raut J (2019) Current status, challenges and prospects of mushroom industry in Nepal. Int J Agric Econ 4:154. https://doi.org/10.11648/j.ijae.20190404.13

Karana E, Blauwhoff D, Hultink EJ, Camere S (2018) When the material grows: a case study on designing (with) mycelium-based materials. Int J Des 12:119–136

Karasova P, Spiwok V, Malá Š et al (2002) Beta-galactosidase activity in psychrotrophic microorganisms and their potential use in food industry. Czech J Food Sci 20:43. https://doi.org/10.17221/3508-CJFS

Karima B, Samia T (2020) Native arbuscular mycorrhizal fungi and agro-industries in arid lands: productions, applications strategies and challenges. In: Radhakrishnan R (ed) mycorrhizal fungi. IntechOpen, Rijeka

Kawasaki Y (2009) Mizoribine: a new approach in the treatment of renal disease. Clin Dev Immunol 2009:681482. https://doi.org/10.1155/2009/681482

Keller NP (2019) Fungal secondary metabolism: regulation, function and drug discovery. Nat Rev Microbiol 17:167–180. https://doi.org/10.1038/s41579-018-0121-1

Kluepfel D, Bagli J, Baker H et al (1972) Myriocin, a new antifungal antibiotic from Myriococcum albomyces . J Antibiotics 25:109–115. https://doi.org/10.7164/antibiotics.25.109

Koné MK, Guéhi ST, Durand N et al (2016) Contribution of predominant yeasts to the occurrence of aroma compounds during cocoa bean fermentation. Food Res Int 89:910–917. https://doi.org/10.1016/j.foodres.2016.04.010

Kotowski M (2016) Differences between European regulations on wild mushroom commerce and actual trends in wild mushroom picking. Slovak Ethnol 64:169–178

Koul B, Farooq B (2020) Chapter 12-Mycotechnology: utility of fungi in food and beverage industries. In: Singh J, Gehlot P (eds) New and future developments in microbial biotechnology and bioengineering. Elsevier, Amsterdam, pp 133–153

Krücken J, Holden-Dye L, Keiser J et al (2021) Development of emodepside as a possible adulticidal treatment for human onchocerciasis-The fruit of a successful industrial-academic collaboration. PLoS Pathog 17:e1009682. https://doi.org/10.1371/journal.ppat.1009682

Kulshreshtha S, Mathur N, Bhatnagar P (2014) Mushroom as a product and their role in mycoremediation. AMB Express 4:29. https://doi.org/10.1186/s13568-014-0029-8

Kulshrestha S, Tyagi P, Sindhi V, Yadavilli KS (2013) Invertase and its applications–a brief review. J Pharm Res 7:792–797. https://doi.org/10.1016/j.jopr.2013.07.014

Lagashetti AC, Dufossé L, Singh SK, Singh PN (2019) Fungal pigments and their prospects in different industries. Microorganisms 7:604. https://doi.org/10.3390/microorganisms7120604

Lahue C, Madden AA, Dunn RR, Smukowski Heil C (2020) History and domestication of Saccharomyces cerevisiae in bread baking. Front Genet 11:584718. https://doi.org/10.3389/fgene.2020.584718

Lai T, Gao Y, Zhou S (2004) Global marketing of medicinal Ling Zhi mushroom Ganoderma lucidum (W.Curt.:Fr.) Lloyd (Aphyllophoromycetideae) products and safety concerns. Int J Med Mushrooms 6:189–194. https://doi.org/10.1615/IntJMedMushr.v6.i2.100

Laich F, Fierro F, Martín JF (2002) Production of penicillin by fungi growing on food products: identification of a complete penicillin gene cluster in Penicillium griseofulvum and a truncated cluster in Penicillium verrucosum . Appl Environ Microbiol 68:1211–1219. https://doi.org/10.1128/AEM.68.3.1211-1219.2002

Lateef A, Oloke J, Gueguim-Kana E, Raimi O (2012) Production of fructosyltransferase by a local isolate of Aspergillus niger in both submerged and solid substrate media. Acta Aliment 41:100–117. https://doi.org/10.1556/AAlim.41.2012.1.12

Lee WY, Park E-J, Ahn JK, Ka K-H (2009) Ergothioneine contents in fruiting bodies and their enhancement in mycelial cultures by the addition of methionine. Mycobiology 37:43. https://doi.org/10.4489/MYCO.2009.37.1.043

Lei W, Zhang G, Peng Q, Liu X (2015) Development of Ophiocordyceps sinensis through plant-mediated interkingdom host colonization. Int J Mol Sci 16:17482–17493. https://doi.org/10.3390/ijms160817482

Lenihan C, Tan JC (2020) Clinical management of the adult kidney transplant recipient. In: Yu ASL, Chertow GM, Luyckx VA et al (eds) Brenner and rector’s the kidney, 11th edn. Elsevier, Philadelphia, pp 2244–2287

Lessard M-H, Viel C, Boyle B et al (2014) Metatranscriptome analysis of fungal strains Penicillium camemberti and Geotrichum candidum reveal cheese matrix breakdown and potential development of sensory properties of ripened Camembert-type cheese. BMC Genomics 15:235. https://doi.org/10.1186/1471-2164-15-235

Li C, Zhu Y, Wang Y et al (1998) Monascus purpureus -fermented rice (red yeast rice): a natural food product that lowers blood cholesterol in animal models of hypercholesterolemia. Nutr Res 18:71–81. https://doi.org/10.1016/S0271-5317(97)00201-7

Li X, Lin X, Zhang J et al (2010) Degradation of polycyclic aromatic hydrocarbons by crude extracts from spent mushroom substrate and its possible mechanisms. Curr Microbiol 60:336–342. https://doi.org/10.1007/s00284-009-9546-0

Li H, Tian Y, Menolli N et al (2021) Reviewing the world’s edible mushroom species: a new evidence-based classification system. Compr Rev Food Sci Food Saf 20:1982–2014. https://doi.org/10.1111/1541-4337.12708

Liaud N, Giniés C, Navarro D et al (2014) Exploring fungal biodiversity: organic acid production by 66 strains of filamentous fungi. Fungal Biol Biotechnol 1:1. https://doi.org/10.1186/s40694-014-0001-z

Article PubMed Central Google Scholar

Lin X, Kück U (2022) Cephalosporins as key lead generation beta-lactam antibiotics. Appl Microbiol Biotechnol 106:8007–8020. https://doi.org/10.1007/s00253-022-12272-8

Lin AX, Chan G, Hu Y et al (2018) Internationalization of traditional Chinese medicine: current international market, internationalization challenges and prospective suggestions. Chin Med 13:9. https://doi.org/10.1186/s13020-018-0167-z

Lipsky JJ (1996) Mycophenolate mofetil. Lancet (london, England) 348:1357–1359. https://doi.org/10.1016/S0140-6736(96)10310-X

Liu Y, Li X, Kou Y (2020a) Ectomycorrhizal fungi: participation in nutrient turnover and community assembly pattern in forest ecosystems. Forests 11:453. https://doi.org/10.3390/f11040453

Liu Y, Luo M, Liu F et al (2020b) Effects of freeze drying and hot-air drying on the physicochemical properties and bioactivities of polysaccharides from Lentinula edodes . Int J Biol Macromol 145:476–483. https://doi.org/10.1016/j.ijbiomac.2019.12.222

Lobanovska M, Pilla G (2017) Penicillin’s discovery and antibiotic resistance: lessons for the future? Yale J Biol Med 90:135–145

CAS PubMed PubMed Central Google Scholar

Lu H, Lou H, Hu J et al (2020) Macrofungi: a review of cultivation strategies, bioactivity, and application of mushrooms. Compr Rev Food Sci Food Saf 19:2333–2356. https://doi.org/10.1111/1541-4337.12602

Magnuson JK, Lasure LL (2004) Organic acid production by filamentous fungi. In: Lange L (ed) Advances in fungal biotechnology for industry, agriculture, and medicine. Springer, Boston, pp 307–340

Maheshwari G, Ahlborn J, Rühl M (2021) Role of fungi in fermented foods. In: Zaragoza O, Casadevall A (eds) Encyclopedia of mycology. Elsevier, Amsterdam, pp 590–600

Mahoney RR (1985) Modification of lactose and lactose-containing dairy products with β-galactosidase. In: Fox PF (ed) Developments in dairy chemistry—3. Springer, Dordrecht, pp 69–109

Maicas S (2020) The role of yeasts in fermentation processes. Microorganisms 8:1142. https://doi.org/10.3390/microorganisms8081142

Mapook A, Hyde KD, Hassan K et al (2022) Ten decadal advances in fungal biology leading towards human well-being. Fungal Divers 116:547–614. https://doi.org/10.1007/s13225-022-00510-3

Marini Govigli V, Górriz-Mifsud E, Varela E (2019) Zonal travel cost approaches to assess recreational wild mushroom picking value: trade-offs between online and onsite data collection strategies. For Policy Econ 102:51–65. https://doi.org/10.1016/j.forpol.2019.02.003

Market Data Forecast (2022a) Button mushrooms market. https://www.marketdataforecast.com/market-reports/button-mushrooms-market . accessed 21 Dec 2022

Market Data Forecast (2022b) Rum market. https://www.marketdataforecast.com/market-reports/rum-market . accessed 2 Sep 2022

Market Data Forecast (2022c) Itaconic acid market. https://www.marketdataforecast.com/market-reports/itaconic-acid-market . accessed 31 Oct 2022

Market Data Forecast (2022d) L-ergothioneine market size 2022d: 29.76% CAGR | Current trend, share, competitors and forecast. https://www.marketwatch.com/press-release/l-ergothioneine-market-size-2022d-2976-cagr-current-trend-share-competitors-and-forecast-2022d-10-19 . accessed 12 Nov 2022

Market Data Forecast (2022e) Mycorrhiza-based biofertilizer market. https://www.marketdataforecast.com/market-reports/mycorrhiza-based-biofertilizer-market . accessed 3 Feb 2023

Market Watch (2022a) Penicillin market. https://www.marketwatch.com/press-release/penicillin-market-size-share-trends-massive-growth-is-expected-to-grow-at-a-cagr-of-38-cagr-forecast-by-2028-108-pages-report-2022a-07-21?tesla=y . accessed 21 Jul 2022

Market Watch (2022b) Fusidic acid market. https://www.marketwatch.com/press-release/fusidic-acid-market-size-2022b-spread-across-104-pages-with-technological-advancements-in-report-recent-innovations-value-chain-analysis-till-2028-2022b-07-13#:~:text=219 is 73.27%25.-,The Global Fusidic Acid Market. Accessed 21 Jul 2022

Market Watch (2023a) Mycophenolate Mofetil market and its economic impact on industry with forecast 2028. https://www.marketwatch.com/press-release/mycophenolate-mofetil-market-and-its-economic-impact-on-industry-with-forecast-2028-2023a-03-01 . accessed 12 Mar 2023

Market Watch (2023b) Lentinan market overview 2023b to 2030, future trends and forecast. https://www.marketwatch.com/press-release/lentinan-market-overview-2023b-to-2030-future-trends-and-forecast-ajinomoto-elicityl-nammex-2023b-01-09?mod=search_headline . accessed 19 Jan 2023

Market Watch (2022c) Kojic acid market analysis and insights. https://www.marketwatch.com/press-release/kojic-acid-market-size-2022c-2028-global-industrial-analysis-by-key-companies-profiled-sale-recent-developments-and-forecast-research-report-2022c-10-17#:~:text=Due to the COVID-19,the forecast period 2022c–2028. accessed 12 Nov 2022

Market Watch (2022d) Strobilurin fungicide market size 2023 global gross margin analysis, industry leading players update, development history, business prospect and industry research report 2028. https://www.marketwatch.com/press-release/strobilurin-fungicide-market-size-2023-global-gross-margin-analysis-industry-leading-players-update-development-history-business-prospect-and-industry-research-report-2028-2022b-11-02 . accessed 12 Jan 2023

Markets and Markets (2022a) Lactic acid market. https://www.marketsandmarkets.com/Market-Reports/polylacticacid-387.html?gclid=Cj0KCQjw--2aBhD5ARIsALiRlwDxrAlnCeieNtgEj25leT0adW95t_qWpjTRYnI1ZDUzcsBMxFumT64aApxxEALw_wcB . accessed 29 Oct 2022

Markets and Markets (2022b) Beta glucan market. https://www.marketsandmarkets.com/Market-Reports/beta-glucan-market-5191796.html . accessed 13 Nov 2022

Marquina T, Emery M, Hurley P, Gould RK (2022) The ‘quiet hunt’: the significance of mushroom foraging among Russian-speaking immigrants in New York City. Ecosyst People 18:226–240. https://doi.org/10.1080/26395916.2022.2055148

Martin-Dominguez V, Cabrera PIA, Eidt L et al (2022) Production of fumaric acid by Rhizopus arrhizus NRRL 1526: a simple production medium and the kinetic modelling of the bioprocess. Fermentation 8:64. https://doi.org/10.3390/fermentation8020064

Martin-Lopez B, Montes C, Benayas J (2008) Economic valuation of biodiversity conservation: the meaning of numbers. Conserv Biol 22:624–635. https://doi.org/10.1111/j.1523-1739.2008.00921.x

Martínez AT, Camarero S, Ruiz-Dueñas FJ, Martínez MJ (2018) Biological lignin degradation. In: Beckham G (ed) Lignin Valorization: Emerging Approaches. RSC, Washington DC, pp 199–225

Martinez SJ, Bressani APP, Dias DR et al (2019) Effect of bacterial and yeast starters on the formation of volatile and organic acid compounds in coffee beans and selection of flavors markers precursors during wet fermentation. Front Microbiol 10:1287. https://doi.org/10.3389/fmicb.2019.01287

Martínez de Aragón J, Riera P, Giergiczny M, Colinas C (2011) Value of wild mushroom picking as an environmental service. For Policy Econ 13:419–424. https://doi.org/10.1016/j.forpol.2011.05.003

Matei G, Matei S, Pele M et al (2017) Invertase production by fungi, characterization of enzyme activity and kinetic parameters. Rev Chim 68:2205–2208. https://doi.org/10.37358/RC.17.10.5856

Maximize Market Research (2021) Agarwood oil market: Global industry analysis and forecast (2022–2029). http://www.maximizemarketresearch.com/market-report/global-agarwood-oil-market/107023/ . accessed 9 Jan 2023

May A, Narayanan S, Alcock J et al (2019) Kombucha: a novel model system for cooperation and conflict in a complex multi-species microbial ecosystem. PeerJ 7:e7565. https://doi.org/10.7717/peerj.7565

Menezes AGT, Menezes EGT, Alves JGLF et al (2016) Vodka production from potato ( Solanum tuberosum L. ) using three Saccharomyces cerevisiae isolates. J Inst Brew 122:76–83. https://doi.org/10.1002/jib.302

Metin B (2018) Filamentous fungi in cheese production. In: Budak SO, Akal HC (eds) Microbial cultures and enzymes in dairy technology, 1st edn. IGI Global, Pennsylvania, pp 257–275

Meyer V, Basenko EY, Benz JP et al (2020) Growing a circular economy with fungal biotechnology: a white paper. Fungal Biol Biotechnol 7:5. https://doi.org/10.1186/s40694-020-00095-z

Miller J (2019) Novartis MS drug wins FDA’s blessing amid scrutiny of $88,000 annual price. In: Reuters. https://www.reuters.com/article/us-fda-mayzent-idUSKCN1R8019 . accessed 23 Aug 2022

Miyadera T, Sugimura Y, Hashimoto T et al (1983) Synthesis and in vitro activity of a new carbapenem, RS-533. J Antibiotics 36:1034–1039. https://doi.org/10.7164/antibiotics.36.1034

Mizuno K, Tsujino M, Takada M et al (1974) Studies on bredinin. I. Isolation, characterization and biological properties. J Antibiotics 27:775–782. https://doi.org/10.7164/antibiotics.27.775

Molla AH, Manjurul Haque M, Amdadul Haque M, Ilias GNM (2012) Trichoderma -enriched biofertilizer enhances production and nutritional quality of tomato ( Lycopersicon esculentum Mill.) and minimizes NPK fertilizer use. Agric Res 1:265–272. https://doi.org/10.1007/s40003-012-0025-7

Monteiro de Souza P, De Oliveira e Magalhães P (2010) Application of microbial α-amylase in industry-a review. Braz J Microbiol 41:850–861. https://doi.org/10.1590/S1517-83822010000400004

Monteiro de Souza P, de Bittencourt ML et al (2015) A biotechnology perspective of fungal proteases. Braz J Microbiol 46:337–346. https://doi.org/10.1590/S1517-838246220140359

Mordor Intelligence (2022) Natural food colorants market - growth, trend, Covid-19 impact, and forecast (2022–2027). https://www.mordorintelligence.com/industry-reports/global-natural-food-colorants-market . accessed 3 Nov 2022

Mortimer PE, Karunarathna SC, Li Q et al (2012) Prized edible Asian mushrooms: ecology, conservation and sustainability. Fungal Divers 56:31–47. https://doi.org/10.1007/s13225-012-0196-3

MycoWorks (2023) Grow the future of materials. https://www.mycoworks.com/ . accessed 2 Feb 2023

Nadaroglu H, Polat MS (2022) Microbial extremozymes: novel sources and industrial applications. In: Kuddos M (ed) Microbial extremozymes. Elsevier, Amsterdam, pp 67–88

Naeem M, Manzoor S, Abid M-U-H et al (2022) Fungal proteases as emerging biocatalysts to meet the current challenges and recent developments in biomedical therapies: an updated review. J Fungi 8:109. https://doi.org/10.3390/jof8020109

National Center for Biotechnology Information (2022a) PubChem compound summary for CID 441140, Griseofulvin. https://pubchem.ncbi.nlm.nih.gov/compound/441140 . accessed 20 Aug 2022

National Center for Biotechnology Information (2022b) PubChem compound summary for CID 10690, Gluconic acid. https://pubchem.ncbi.nlm.nih.gov/compound/Gluconic-acid . accessed 21 Sep 2022

NEFFA (2023) The future of fashion manufacturing. https://www.mycotex.nl/ . accessed 2 Feb 2023

Neu HC, Chin NX, Saha G, Labthavikul P (1986) In vitro activity against aerobic and anaerobic gram-positive and gram-negative bacteria and beta-lactamase stability of RS-533, a novel carbapenem. Antimicrob Agents Chemother 30:828–834. https://doi.org/10.1128/AAC.30.6.828

Newman DJ, Cragg GM (2020) Natural products as sources of new drugs over the nearly four decades from 01/1981 to 09/2019. J Nat Prod 83:770–803. https://doi.org/10.1021/acs.jnatprod.9b01285

Newton GG, Abraham EP (1955) Cephalosporin C, a new antibiotic containing sulphur and d -alpha-aminoadipic acid. Nature 175:548. https://doi.org/10.1038/175548a0

Nguyen T, Karl M, Santini A (2017) Red yeast rice. Foods 6:19. https://doi.org/10.3390/foods6030019

Niego AG, Rapior S, Thongklang N et al (2021) Macrofungi as a nutraceutical source: promising bioactive compounds and market value. J Fungi 7:397. https://doi.org/10.3390/jof7050397

Niego AGT, Rapior S, Thongklang N et al (2023) Reviewing the contributions of macrofungi to forest ecosystem processes and services. Fungal Biol Rev 44:100294. https://doi.org/10.1016/j.fbr.2022.11.002

Nigam PS, Singh A (2014) Single cell protein | Mycelial fungi. In: Batt CA, Tortorello ML (eds) Encyclopedia of Food microbiology, 2nd edn. Elsevier, Amsterdam, pp 415–424

Nofiani R, de Mattos-Shipley K, Lebe KE et al (2018) Strobilurin biosynthesis in basidiomycete fungi. Nat Commun 9:3940. https://doi.org/10.1038/s41467-018-06202-4

OEC (2022a) Blue Cheeses. https://oec.world/en/profile/hs/cheese-blue-veined . accessed 27 Aug 2022

OEC (2022b) Baked goods. https://oec.world/en/profile/hs/baked-goods . accessed 27 Aug 2022

OEC (2022c) Cheese. https://oec.world/en/profile/hs/cheese . accessed 27 Aug 2022

OEC (2022d) Coffee. https://oec.world/en/profile/hs/coffee#exporters-importers . accessed 5 Sep 2022

Oxford AE, Raistrick H, Simonart P (1939) Studies in the biochemistry of micro-organisms: Griseofulvin, C(17)H(17)O(6)Cl, a metabolic product of Penicillium griseo-fulvum Dierckx. Biochem J 33:240–248. https://doi.org/10.1042/bj0330240

Pan X, Liu H, Liu J et al (2016) Omics-based approaches reveal phospholipids remodeling of Rhizopus oryzae responding to furfural stress for fumaric acid-production from xylose. Bioresour Technol 222:24–32. https://doi.org/10.1016/j.biortech.2016.09.101

Pandey A, Höfer R, Taherzadeh M et al (2015) Industrial biorefineries & white biotechnology. Elsevier, Amsterdam

Pandey A, Nigam P, Soccol CR et al (2000) Advances in microbial amylases. Biotechnol Appl Biochem 31:135. https://doi.org/10.1042/BA19990073

Papp-Wallace KM, Endimiani A, Taracila MA, Bonomo RA (2011) Carbapenems: past, present, and future. Antimicrob Agents Chemother 55:4943–4960. https://doi.org/10.1128/AAC.00296-11

Paterson RRM (2008) Cordyceps – A traditional Chinese medicine and another fungal therapeutic biofactory? Phytochemistry 69:1469–1495. https://doi.org/10.1016/j.phytochem.2008.01.027

Paterson RRM (2006) Ganoderma –a therapeutic fungal biofactory. Phytochemistry 67:1985–2001. https://doi.org/10.1016/j.phytochem.2006.07.004

Paukner S, Riedl R (2017) Pleuromutilins: potent drugs for resistant bugs-mode of action and resistance. Cold Spring Harb Perspect Med 7:a027110. https://doi.org/10.1101/cshperspect.a027110

Pauley M, Maskell D (2017) Mini-review: The role of Saccharomyces cerevisiae in the production of gin and vodka. Beverages 3:13. https://doi.org/10.3390/beverages3010013

Pedersen TR, Tobert JA (2004) Simvastatin: a review. Expert Opin Pharmacother 5:2583–2596. https://doi.org/10.1517/14656566.5.12.2583

Peláez F, Cabello A, Platas G et al (2000) The discovery of enfumafungin, a novel antifungal compound produced by an endophytic Hormonema species biological activity and taxonomy of the producing organisms. Syst Appl Microbiol 23:333–343. https://doi.org/10.1016/S0723-2020(00)80062-4

Pérez-Chávez AM, Mayer L, Albertó E (2019) Mushroom cultivation and biogas production: a sustainable reuse of organic resources. Energy Sustain Dev 50:50–60

Persistence Market Research (2022) Red yeast rice market. https://www.persistencemarketresearch.com/market-research/red-yeast-rice-market.asp . accessed 19 Jan 2023

Phasha V, Senabe J, Ndzotoyi P et al (2022) Review on the use of kojic acid—a skin-lightening ingredient. Cosmetics 9:64. https://doi.org/10.3390/cosmetics9030064

Polaris Market Research (2022) Carbapenem market share, size, trends, industry analysis report, by product type (Meropenem, Doripenem, Imipenem, Tebipenem, and others); by sales channel; by region; segment forecast, 2022–2030. https://www.polarismarketresearch.com/industry-analysis/carbapenem-market . accessed 3 Feb 2023

Polaris Market Research (2021) Kombucha market share, trends, size, industry analysis report, by product (organic, inorganic), by type (natural, flavored), by microbial, by distribution channel, by region; Segment forecast, 2022–2030. https://www.polarismarketresearch.com/industry-analysis/kombucha-market . accessed 6 May 2023

Poorniammal R, Prabhu S, Dufossé L, Kannan J (2021) Safety evaluation of fungal pigments for food applications. J Fungi 7:692. https://doi.org/10.3390/jof7090692

Prenafeta-Boldú FX, de Hoog GS, Summerbell RC (2019) Fungal communities in hydrocarbon degradation. In: McGenity TJ (ed) Microbial communities utilizing hydrocarbons and lipids: members, metagenomics and ecophysiology. Springer, Cham, pp 307–342

Pritchard PE (1992) Studies on the bread-improving mechanism of fungal alpha-amylase. J Biol Educ 26:12–18. https://doi.org/10.1080/00219266.1992.9655237

PS Market Research (2022) Industrial enzymes market research report. https://www.psmarketresearch.com/market-analysis/industrial-enzymes-market . accessed 31 Oct 2022

Raghoonundon B, Gonkhom D, Phonemany M et al (2021) Nutritional content, nutraceutical properties, cultivation methods and economical importance of Lentinula : a review. Fungal Biotec 1:88–100. https://doi.org/10.5943/FunBiotec/1/2/6

Rakhee MJ, Yadav RB et al (2021) Novel formulation development from Ophiocordyceps sinensis (Berk) for management of high-altitude maladies. 3 Biotech 11:9. https://doi.org/10.1007/s13205-020-02536-3

Rathore H, Prasad S, Sharma S (2017) Mushroom nutraceuticals for improved nutrition and better human health: a review. PharmaNutrition 5:35–46. https://doi.org/10.1016/j.phanu.2017.02.001

Refinitiv (2022) Carbon trading: exponential growth on record high. https://www.refinitiv.com/perspectives/market-insights/carbon-trading-exponential-growth-on-record-high/ . accessed 11 Jan 2023

Reportlinker (2022) Global fumaric acid industry. https://www.reportlinker.com/p05817775/Global-Fumaric-Acid-Industry.html?utm_source=GNW . accessed 31 Oct 2022

Research and Markets (2022a) Coffee market - growth, trends, COVID-19 impact, and forecasts (2022a - 2027). https://www.researchandmarkets.com/reports/5165416/coffee-market-growth-trends-covid-19-impact?utm_source=BW&utm_medium=PressRelease&utm_code=zj95v9&utm_campaign=1671722+-+%24102+Billion+Global+Coffee+Market+Growth%2C+Trends%2C+COVID-19+Impact%2C+and+Fore . accessed 5 Sep 2022

Research and Markets (2022b) Biopesticides market by type. https://www.researchandmarkets.com/reports/5116567/biopesticides-market-by-type-bioinsecticides?utm_source=GNOM&utm_medium=PressRelease&utm_code=g36rbx&utm_campaign=1736666+-+Global+Biopesticides+Market+Report+2022b-2027%3A+Pests+Developing+Resistance+to+C . accessed 12 Jan 2023

Research and Markets (2022c) Global mycelium market analysis and forecast, 2020–2021 & 2022c-2026 - adoption of mycelium in pharmaceutical and skin care industry. https://www.researchandmarkets.com/reports/5559925/mycelium-market-a-global-and-regional-analysis?utm_source=GNOM&utm_medium=PressRelease&utm_code=j2k7m4&utm_campaign=1685003+-+Global+Mycelium+Market+Analysis+and+Forecast%2C+2020-2021+%26+2022c-2026+-+Adop . accessed 21 Jun 2022

Rezac S, Kok CR, Heermann M, Hutkins R (2018) Fermented foods as a dietary source of live organisms. Front Microbiol 9:1785. https://doi.org/10.3389/fmicb.2018.01785

Rodríguez-Sáiz M, de la Fuente JL, Barredo JL (2010) Xanthophyllomyces dendrorhous for the industrial production of astaxanthin. Appl Microbiol Biotechnol 88:645–658. https://doi.org/10.1007/s00253-010-2814-x

Ropars J, Didiot E, Rodríguez de la Vega RC et al (2020) Domestication of the emblematic white cheese-making fungus Penicillium camemberti and its diversification into two varieties. Curr Biol 30:4441–4453. https://doi.org/10.1016/j.cub.2020.08.082

Rosa LH, Machado KMG, Jacob CC et al (2003) Screening of Brazilian basidiomycetes for antimicrobial activity. Mem Inst Oswaldo Cruz 98:967–974. https://doi.org/10.1590/S0074-02762003000700019

Rosa LH, Cota BB, Machado KMG et al (2005) Antifungal and other biological activities from Oudemansiella canarii (Basidiomycota). World J Microbiol Biotechnol 21:983–987. https://doi.org/10.1007/s11274-004-7553-7

Royse DJ, Baars J, Tan Q (2017) Current overview of mushroom production in the world. In: Diego CZ, Pardo-Giménez A (eds) Edible and medicinal mushrooms. Wiley, Chichester, pp 5–13

Rüegger A, Kuhn M, Lichti H et al (1976) Cyclosporin A, a peptide metabolite from Trichoderma polysporum (Link ex Pers.) Rifai, with a remarkable immunosuppressive activity. Helv Chim Acta 59:1075–1092. https://doi.org/10.1002/hlca.19760590412

Ruta LL, Farcasanu IC (2021) Coffee and yeasts: from flavor to biotechnology. Fermentation 7:9. https://doi.org/10.3390/fermentation7010009

Saeedi M, Eslamifar M, Khezri K (2019) Kojic acid applications in cosmetic and pharmaceutical preparations. Biomed Pharmacother 110:582–593. https://doi.org/10.1016/j.biopha.2018.12.006

Saini S, Sharma KK (2021) Fungal lignocellulolytic enzymes and lignocellulose: a critical review on their contribution to multiproduct biorefinery and global biofuel research. Int J Biol Macromol 193:2304–2319. https://doi.org/10.1016/j.ijbiomac.2021.11.063

San-Juan-Rodriguez A, Good CB, Heyman RA et al (2019) Trends in prices, market share, and spending on self-administered disease-modifying therapies for multiple sclerosis in medicare part D. JAMA Neurol 76:1386. https://doi.org/10.1001/jamaneurol.2019.2711

Sandargo B, Chepkirui C, Cheng T et al (2019) Biological and chemical diversity go hand in hand: Basidiomycota as source of new pharmaceuticals and agrochemicals. Biotechnol Adv 37:107344. https://doi.org/10.1016/j.biotechadv.2019.01.011

Sandmann G (2022) Carotenoids and their biosynthesis in fungi. Molecules 27:1431. https://doi.org/10.3390/molecules27041431

Sandoval-Lozano CJ, Caballero-Torres D, López-Giraldo LJ (2022) Screening wild yeast isolated from cocoa bean fermentation using volatile compounds profile. Molecules 27:902. https://doi.org/10.3390/molecules27030902

Saqib S, Akram A, Halim SA, Tassaduq R (2017) Sources of β-galactosidase and its applications in food industry. 3 Biotech 7:79. https://doi.org/10.1007/s13205-017-0645-5

Sasaki T, Takagi M, Yaguchi T et al (1992) A new anthelmintic cyclodepsipeptide, PF1022A. J Antibiotics 45:692–697. https://doi.org/10.7164/antibiotics.45.692

Sawistowska-Schröder ET, Kerridge D, Perry H (1984) Echinocandin inhibition of 1,3-beta- d -glucan synthase from Candida albicans . FEBS Lett 173:134–138. https://doi.org/10.1016/0014-5793(84)81032-7

Scherkenbeck J, Jeschke P, Harder A (2002) PF1022A and related cyclodepsipeptides-a novel class of anthelmintics. Curr Top Med Chem 2:759–777. https://doi.org/10.2174/1568026023393624

Schmidt L, Skvortsova V, Kullen C et al (2017) How context alters value: the brain’s valuation and affective regulation system link price cues to experienced taste pleasantness. Sci Rep 7:8098. https://doi.org/10.1038/s41598-017-08080-0

Schulp CJE, Thuiller W, Verburg PH (2014) Wild food in Europe: a synthesis of knowledge and data of terrestrial wild food as an ecosystem service. Ecol Econ 105:292–305. https://doi.org/10.1016/j.ecolecon.2014.06.018

Schwartz RE, Giacobbe RA, Bland JA, Monaghan RL (1989) L-671,329, a new antifungal agent. I. Fermentation and isolation. J Antibiotics 42:163–167. https://doi.org/10.7164/antibiotics.42.163

Scynexis (2022) SCYNEXIS reports fourth quarter and full year 2021 financial results and provides corporate update. https://www.scynexis.com/news-media/press-releases/detail/278/scynexis-reports-fourth-quarter-and-full-year-2021 . accessed 23 Aug 2022

Sebastiana M, da Silva AB, Matos AR et al (2018) Ectomycorrhizal inoculation with Pisolithus tinctorius reduces stress induced by drought in cork oak. Mycorrhiza 28:247–258. https://doi.org/10.1007/s00572-018-0823-2

Seyis I, Aksoz N (2004) Production of lactase by Trichoderma sp. Food Technol Biotechnol 42:121–124

Sharma N, Sahota P, Singh M (2021) Organic acid production from agricultural waste. In: Kashyap BK, Solanki MK, Kamboj DV, Pandey AK (eds) Waste to energy: prospects and applications. Springer, Singapore, pp 415–438

Show PL, Oladele KO, Siew QY et al (2015) Overview of citric acid production from Aspergillus niger . Front Life Sci 8:271–283. https://doi.org/10.1080/21553769.2015.1033653

Shrestha UB (2012) Asian medicine: a fungus in decline. Nature 482:35. https://doi.org/10.1038/482035b

Silverman J, Cao H, Cobb K (2020) Development of mushroom mycelium composites for footwear products. Cloth Text Res J 38:119–133. https://doi.org/10.1177/0887302X19890006

Singh AK, Mukhopadhyay M (2012) Overview of fungal lipase: a review. Appl Biochem Biotechnol 166:486–520. https://doi.org/10.1007/s12010-011-9444-3

Singh RS, Chauhan K, Singh RP (2017) Enzymatic approaches for the synthesis of high fructose syrup. In: Gahlawat S, Salar R, Siwach P, Duhan J, Kumar S, Kaur P (eds) Plant biotechnology: recent advancements and developments. Springer, Singapore, pp 189–211

Sjamsuridzal W, Khasanah M, Febriani R et al (2021) The effect of the use of commercial tempeh starter on the diversity of Rhizopus tempeh in Indonesia. Sci Rep 11:23932. https://doi.org/10.1038/s41598-021-03308-6

Ślusarczyk J, Adamska E, Czerwik-Marcinkowska J (2021) Fungi and algae as sources of medicinal and other biologically active compounds: a review. Nutrients 13:3178. https://doi.org/10.3390/nu13093178

Solieri L, Giudici P (2008) Yeasts associated to traditional balsamic vinegar: ecological and technological features. Int J Food Microbiol 125:36–45. https://doi.org/10.1016/j.ijfoodmicro.2007.06.022

Soudzilovskaia NA, Vaessen S, Barcelo M et al (2020) FungalRoot: global online database of plant mycorrhizal associations. New Phytol 227:955–966. https://doi.org/10.1111/nph.16569

Soudzilovskaia NA, van Bodegom PM, Terrer C et al (2019) Global mycorrhizal plant distribution linked to terrestrial carbon stocks. Nat Commun 10:5077. https://doi.org/10.1038/s41467-019-13019-2

Special Chem (2022) Parnika mushroom. https://cosmetics.specialchem.com/product/i-parnika-parnika-mushroom . accessed 2 Dec 2022

Statista (2022a) Global biofuels market size 2020–2030. https://www.statista.com/statistics/217179/global-biofuels-market-size/#:~:text=The global biofuels market was,to 201.2 billion U.S. dollars. accessed 20 Jun 2022

Statista (2022b) Global gross domestic product (GDP) at current prices from 1985 to 2027(in billion U.S. dollars). https://www.statista.com/statistics/268750/global-gross-domestic-product-gdp/#:~:text=The statistic shows global gross,trillion lower than in 2019. accessed 10 Mar 2023

Steiger MG, Blumhoff ML, Mattanovich D, Sauer M (2013) Biochemistry of microbial itaconic acid production. Front Microbiol 4:23. https://doi.org/10.3389/fmicb.2013.00023

Steinberg D (2006) Thematic review series: the pathogenesis of atherosclerosis. An interpretive history of the cholesterol controversy, part V: the discovery of the statins and the end of the controversy. J Lipid Res 47:1339–1351. https://doi.org/10.1194/jlr.R600009-JLR200

Stelzer L, Hoberg F, Bach V et al (2021) Life cycle assessment of fungal-based composite bricks. Sustainability 13:11573. https://doi.org/10.3390/su132111573

Stoffel F, de Santana W, O, Fontana RC, Camassola M, (2021) Use of Pleurotus albidus mycoprotein flour to produce cookies: Evaluation of nutritional enrichment and biological activity. Innov Food Sci Emerg Technol 68:102642. https://doi.org/10.1016/j.ifset.2021.102642

Straits Research (2021) Agarwood chip market. https://straitsresearch.com/report/agarwood-chip-market/ . accessed 14 Jan 2023

Sudha GC, Aggarwal S (2016) Dyeing wet blue goat nappa skin with a microbial colorant obtained from Penicillium minioluteum . J Clean Prod 127:585–590. https://doi.org/10.1016/j.jclepro.2016.03.043

Suhaimi H, Dailin DJ, Malek RA et al (2021) Fungal pectinases: production and applications in food industries. In: Dai X, Sharma M, Chen J (eds) Fungi in sustainable food production. Springer, Cham, pp 85–116

Sun W, Tajvidi M, Hunt CG et al (2019) Fully bio-based hybrid composites made of wood, fungal mycelium and cellulose nanofibrils. Sci Rep 9:3766. https://doi.org/10.1038/s41598-019-40442-8

Sunagawa M, Matsumura H, Inoue T et al (1990) A novel carbapenem antibiotic, SM-7338 structure-activity relationships. J Antibiotics 43:519–532. https://doi.org/10.7164/antibiotics.43.519

Sundelof JG, Hajdu R, Gill CJ et al (1997) Pharmacokinetics of L-749,345, a long-acting carbapenem antibiotic, in primates. Antimicrob Agents Chemother 41:1743–1748. https://doi.org/10.1128/AAC.41.8.1743

Survase SA, Kagliwal LD, Annapure US, Singhal RS (2011) Cyclosporin A–a review on fermentative production, downstream processing and pharmacological applications. Biotechnol Adv 29:418–435. https://doi.org/10.1016/j.biotechadv.2011.03.004

Sydor M, Bonenberg A, Doczekalska B, Cofta G (2021) Mycelium-based composites in art, architecture, and interior design: a review. Polymers (basel) 14:145. https://doi.org/10.3390/polym14010145

Takala T, Lehtinen A, Hujala T et al (2021) Forest owners as political actors. Environ Sci Policy 126:22–30. https://doi.org/10.1016/j.envsci.2021.09.009

Talukdar D, Jasrotia T, Sharma R et al (2020) Evaluation of novel indigenous fungal consortium for enhanced bioremediation of heavy metals from contaminated sites. Environ Technol Innov 20:101050. https://doi.org/10.1016/j.eti.2020.101050

Tamang JP, Cotter PD, Endo A et al (2020) Fermented foods in a global age: East meets West. Compr Rev Food Sci Food Saf 19:184–217. https://doi.org/10.1111/1541-4337.12520

Tanret C (1909) Sur une base nouvelle retiree du seigle ergote, l’ergothioneine. Comptes Rend Acad Sci 149:222–224

Teleky B-E, Vodnar D (2019) Biomass-derived production of itaconic acid as a building block in specialty polymers. Polymers (basel) 11:1035. https://doi.org/10.3390/polym11061035

Terada M, Hayashi K, Mizunuma T (1981) Enzyme productivity and identification of species of Koji-molds for soy sauce distributed in Japan. J Soy Sauce Res Technol 7:158–165

Thakur MP (2020) Advances in mushroom production: key to food, nutritional and employment security: a review. Indian Phytopathol 73:377–395

The Brainy Insights (2022) Shiitake mushroom market size by product type. https://www.thebrainyinsights.com/report/shiitake-mushroom-market-12716#:~:text=As per The Brainy Insights,USD 4.7 billion by 2030. accessed 20 Dec 2022

The Business Research Company (2022) Tempeh global market report 2022. https://www.thebusinessresearchcompany.com/report/tempeh-global-market-report#:~:text=This tempeh market research report,(CAGR) of 7.8%25. Accessed 31 Oct 2022

The Business Research Company (2023) Cheese global market report 2023. https://www.thebusinessresearchcompany.com/report/cheese-global-market-report . accessed 25 Jan 2023

The Spirit Business (2022g) Tequila sales to reach $24.19 billion by 2031. https://www.thespiritsbusiness.com/2022g/08/tequila-sales-to-reach-24-19-billion-by-2031/ . accessed 5 Sep 2022

Thomas K, Proschmann U, Ziemssen T (2017) Fingolimod hydrochloride for the treatment of relapsing remitting multiple sclerosis. Expert Opin Pharmacother 18:1649–1660. https://doi.org/10.1080/14656566.2017.1373093

Tibuhwa DD (2013) Wild mushroom- an underutilized healthy food resource and income generator: experience from Tanzania rural areas. J Ethnobiol Ethnomed 9:49. https://doi.org/10.1186/1746-4269-9-49

Tonkinson B (2022h) How to make vinegar. https://ice.edu/blog/making-vinegars#:~:text=The yeast consumes the natural,to take place%2C acetic fermentation. accessed 5 Sep 2022

Tridge (2022a) Hericium. https://www.tridge.com/intelligences/hericium-mushrooms/PL . accessed 30 Aug 2022

Tridge (2022b) Lion’s mane mushroom. https://www.tridge.com/intelligences/lions-mane-mushroom . accessed 21 Dec 2022

Tsuji M, Ishii Y, Ohno A et al (1998) In vitro and in vivo antibacterial activities of S-4661, a new carbapenem. Antimicrob Agents Chemother 42:94–99. https://doi.org/10.1128/AAC.42.1.94

Tulpule V, Brown S, Lim J, et al (1998) An economic assessment of the Kyoto Protocol using the global trade and environment model. In: The Economic Modelling of Climate Change: Background Analysis for the Kyoto Protocol OECD Workshop. Paris

Ubukata K, Hikida M, Yoshida M et al (1990) In vitro activity of LJC10,627, a new carbapenem antibiotic with high stability to dehydropeptidase I. Antimicrob Agents Chemother 34:994–1000. https://doi.org/10.1128/AAC.34.6.994

UNData (2022i) Mushroom and truffles. http://data.un.org/Data.aspx?d=FAO&f=itemCode:449 . accessed 13 May 2022

Usman M, Murtaza G, Ditta A (2021) Nutritional, medicinal, and cosmetic value of bioactive compounds in button mushroom ( Agaricus bisporus ): a review. Appl Sci 11:5943. https://doi.org/10.3390/app11135943

Valavanidis A (2018) Concept and practice of the circular economy. Sci Rev 143:2–5

Valverde ME, Hernández-Pérez T, Paredes-López O (2015) Edible mushrooms: improving human health and promoting quality life. Int J Microbiol 2015:376387. https://doi.org/10.1155/2015/376387

Varghese R, Dalvi YB, Lamrood PY et al (2019) Historical and current perspectives on therapeutic potential of higher basidiomycetes: an overview. 3 Biotech 9:362. https://doi.org/10.1007/s13205-019-1886-2

Venil CK, Velmurugan P, Dufossé L et al (2020) Fungal pigments: potential coloring compounds for wide ranging applications in textile dyeing. J Fungi 6:68. https://doi.org/10.3390/jof6020068

Verified Market Research (2022a) Cyclosporine market size and forecast. https://www.verifiedmarketresearch.com/product/cyclosporine-market/ . accessed 21 Aug 2022

Verified Market Research (2022b) Soy sauce market size and forecast. https://www.verifiedmarketresearch.com/product/soy-sauce-market/ . accessed 31 Oct 2022

Verma ML, Thakur M, Randhawa JS et al (2020) Biotechnological applications of fungal enzymes with special reference to bioremediation. In: Gothandam KM, Ranjan S, Dasgupta N, Lichtfouse E (eds) Environmental biotechnology, vol 2. Springer, Cham, pp 221–247

Vieira GAL, Magrini MJ, Bonugli-Santos RC et al (2018) Polycyclic aromatic hydrocarbons degradation by marine-derived basidiomycetes: optimization of the degradation process. Braz J Microbiol 49:749–756. https://doi.org/10.1016/j.bjm.2018.04.007

Vilela A (2019) The importance of yeasts on fermentation quality and human health-promoting compounds. Fermentation 5:46. https://doi.org/10.3390/fermentation5020046

Vishwanatha KS, Rao AGA, Singh SA (2010) Acid protease production by solid-state fermentation using Aspergillus oryzae MTCC 5341: optimization of process parameters. J Ind Microbiol Biotechnol 37:129–138. https://doi.org/10.1007/s10295-009-0654-4

Volpi C, Orabona C, Macchiarulo A et al (2019) Preclinical discovery and development of fingolimod for the treatment of multiple sclerosis. Expert Opin Drug Discov 14:1199–1212. https://doi.org/10.1080/17460441.2019.1646244

Walbank J (2022) Can yeast processing improve the quality of coffee?. https://mtpak.coffee/2022/04/yeast-processing-can-it-improve-quality-of-coffee/

Walker G, Stewart G (2016) Saccharomyces cerevisiae in the production of fermented beverages. Beverages 2:30. https://doi.org/10.3390/beverages2040030

Wang C, Sun H, Li J et al (2009) Enzyme activities during degradation of polycyclic aromatic hydrocarbons by white rot fungus Phanerochaete chrysosporium in soils. Chemosphere 77:733–738. https://doi.org/10.1016/j.chemosphere.2009.08.028

Wang J, Zhou Z, Dan D, Hu G (2020) Physicochemical properties and bioactivities of Lentinula edodes polysaccharides at different development stages. Int J Biol Macromol 150:573–577. https://doi.org/10.1016/j.ijbiomac.2020.02.099

Wierckx N, Agrimi G, Lübeck PS et al (2020) Metabolic specialization in itaconic acid production: a tale of two fungi. Curr Opin Biotechnol 62:153–159. https://doi.org/10.1016/j.copbio.2019.09.014

Willson J, Amliwala K, Harder A et al (2003) The effect of the anthelmintic emodepside at the neuromuscular junction of the parasitic nematode Ascaris suum . Parasitology 126:79–86. https://doi.org/10.1017/s0031182002002639

Wilson LA (1995) Practical handbook of soybean processing and utilization. Elsevier, Amsterdam

Wisitrassameewong K, Karunarathna S, Thongklang N et al (2012) Agaricus subrufescens : a review. Saudi J Biol Sci 19:131–146. https://doi.org/10.1016/j.sjbs.2012.01.003

Wittstein K, Cordsmeier A, Lambert C et al (2020) Identification of Rosellinia species as producers of cyclodepsipeptide PF1022 A and resurrection of the genus Dematophora as inferred from polythetic taxonomy. Stud Mycol 96:1–16. https://doi.org/10.1016/j.simyco.2020.01.001

World Economic Forum (2022) Why mushroom mycelium could be your next house, handbag, or ‘hamburger.’ https://www.weforum.org/agenda/2020/12/mycelium-mushroom-sustainable-packaging-fashion-meat/ . accessed 20 Jun 2022

World Population Review (2022) Spain population 2022. https://worldpopulationreview.com/countries/spain-population . accessed 19 Jun 2022

Wright AJ (1999) The penicillins. Mayo Clin Proc 74:290–307. https://doi.org/10.4065/74.3.290

Wu Y, Choi M-H, Li J et al (2016) Mushroom cosmetics: the present and future. Cosmetics 3:22. https://doi.org/10.3390/cosmetics3030022

Yadav P, Chauhan AK, Singh RB et al (2022) Organic acids: microbial sources, production, and applications. In: Singh RB, Watanabe S, Isaza AA (eds) Functional foods and nutraceuticals in metabolic and non-communicable diseases. Elsevier, Amsterdam, pp 325–337

Yang ST, Zhang K, Zhang B, Huang H (2011) 3.16 - Fumaric acid. In: Moo-Young M (ed) Comprehensive biotechnology, 2nd edn. Academic Press, Burlington, pp 163–177

Yokotsuka T, Sasaki M (1998) Fermented protein foods in the Orient: shoyu and miso in Japan. In: Wood BJB (ed) Microbiology of fermented foods. Springer, Boston, pp 351–415

Zaki O, Weekers F, Thonart P et al (2020) Limiting factors of mycopesticide development. Biol Control 144:104220. https://doi.org/10.1016/j.biocontrol.2020.104220

Zeb M, Lee CH (2021) Medicinal properties and bioactive compounds from wild mushrooms native to North America. Molecules 26:251. https://doi.org/10.3390/molecules26020251

Zha LS, Huang SK, Xiao YP et al (2018) An evaluation of common Cordyceps (Ascomycetes) species found in Chinese markets. Int J Med Mushrooms 20:1149–1162. https://doi.org/10.1615/IntJMedMushrooms.2018027330

Zhang N, Zhao X-H (2010) Study of Mucor spp. in semi-hard cheese ripening. J Food Sci Technol 47:613–619. https://doi.org/10.1007/s13197-010-0108-z

Zhang ZY, Jin B, Kelly JM (2007) Production of lactic acid from renewable materials by Rhizopus fungi. Biochem Eng J 35:251–263. https://doi.org/10.1016/j.bej.2007.01.028

Zhang N, Guo Q-Q, Zhao X-H (2012) A brief investigation on the proteolysis and textural modification of a semi-hard cheese ripened by Mucor spp. Biotechnology 12:54–60. https://doi.org/10.3923/biotech.2013.54.60

Zhang HW, Lin ZX, Tung YS et al (2014) Cordyceps sinensis (a traditional Chinese medicine) for treating chronic kidney disease. Cochrane Database Syst Rev 2014:CD008353. https://doi.org/10.1002/14651858.CD008353.pub2

Zhang J, Chen Q, Huang C, Gao W (2015) History, current situation and trend of edible mushroom industry development. Mycosystema 34:524–540. https://doi.org/10.13346/j.mycosystema.150076-en

Zhang Y, He S, Simpson BK (2018) Enzymes in food bioprocessing—novel food enzymes, applications, and related techniques. Curr Opin Food Sci 19:30–35. https://doi.org/10.1016/j.cofs.2017.12.007

Zhen X, Wang Y, Liu D (2020) Bio-butanol as a new generation of clean alternative fuel for SI (spark ignition) and CI (compression ignition) engines. Renew Energy 147:2494–2521. https://doi.org/10.1016/j.renene.2019.10.119

Zhen J, Wang J, Li L et al (2022) Biodegradation of PAHs by Trametes hirsuta zlh237 and effect of bioaugmentation on PAHs-contaminated soil. Polish J Environ Stud 31:2473–2484. https://doi.org/10.15244/pjoes/143010

Zhou QZK, Chen XD (2001) Effects of temperature and pH on the catalytic activity of the immobilized β-galactosidase from Kluyveromyces lactis . Biochem Eng J 9:33–40. https://doi.org/10.1016/S1369-703X(01)00118-8

Zhu J-S, Halpern GM, Jones K (1998) The scientific rediscovery of an ancient Chinese herbal medicine: Cordyceps sinensis Part I. J Altern Complement Med 4:289–303. https://doi.org/10.1089/acm.1998.4.3-289

Zi Y, Jiang B, He C, Liu L (2020) Lentinan inhibits oxidative stress and inflammatory cytokine production induced by benzo(a)pyrene in human keratinocytes. J Cosmet Dermatol 19:502–507. https://doi.org/10.1111/jocd.13005

Zotti M, Persiani AM, Ambrosio E et al (2013) Macrofungi as ecosystem resources: conservation versus exploitation. Plant Biosyst 147:219–225. https://doi.org/10.1080/11263504.2012.753133

Download references

The authors also acknowledge Dr. Eric Boa for sharing his studies and publications. We thank Dr. Olivier Raspé for helping in the initial conceptualization of the paper.

Open Access funding enabled and organized by Projekt DEAL. This work was in part supported by the Thailand Science Research and Innovation grant “Macrofungi diversity research from the Lancang-Mekong Watershed and surrounding areas” (Grant No. DBG6280009) and the National Research Council of Thailand (NRCT) grant “Total fungal diversity in a given forest area with implications towards species numbers, chemical diversity and biotechnology” (Grant No. N42A650547. We thank the Mushroom Research Foundation, Chiang Rai, Thailand for supporting the Ph.D studies of A.G. Niego. Niego would like to acknowledge the Mae Fah Luang University Thesis/Dissertation Support Grant (Reference No. Oh 7702(6)/49). EC-G was supported by the HZI POF IV Cooperativity and Creativity Project Call and CL was funded by a PhD stipend of the Life Science Foundation, Munich.

Author information

Authors and affiliations.

School of Science, Mae Fah Luang University, Chiang Rai, Thailand

Allen Grace T. Niego, Naritsada Thongklang, Arttapon Walker & Kevin D. Hyde

Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, Thailand

Iloilo Science and Technology University, La Paz, 5000, Iloilo, Philippines

Allen Grace T. Niego

Department Microbial Drugs, Helmholtz Centre for Infection Research, Inhoffenstrasse 7, 38124, Brunswick, Germany

Christopher Lambert, Miriam Grosse, Hedda Schrey, Esteban Charria-Girón & Marc Stadler

Institute of Microbiology, Technische Universität Braunschweig, Spielmannstraße 7, 38106, Brunswick, Germany

Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, 650201, Yunnan, People’s Republic of China

Peter Mortimer

Laboratory of Botany, Phytochemistry and Mycology, Faculty of Pharmacy, CS 14491, 15 Avenue Charles Flahault, 34093, Montpellier Cedex 5, France

Sylvie Rapior

CEFE, CNRS, Univ Montpellier, EPHE, IRD, CS 14491, 15 Avenue Charles Flahault, 34093, Montpellier Cedex 5, France

Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai, 50200, Thailand

Kevin D. Hyde

Institute for Plant Health, Zhongkai University of Agriculture and Engineering, Haizhu District, Guangzhou, 510225, People’s Republic of China

You can also search for this author in PubMed Google Scholar

Contributions

The first draft of the manuscript was written by AGN and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.

Corresponding authors

Correspondence to Kevin D. Hyde or Marc Stadler .

Ethics declarations

Competing interests.

The authors declare no competing interests.

Additional information

Handling Editor: R. Jeewon.

Rights and permissions

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/ .

Reprints and permissions

About this article

Niego, A.G.T., Lambert, C., Mortimer, P. et al. The contribution of fungi to the global economy. Fungal Diversity 121 , 95–137 (2023). https://doi.org/10.1007/s13225-023-00520-9

Download citation

Received : 15 March 2023

Accepted : 29 May 2023

Published : 12 July 2023

Issue Date : July 2023

DOI : https://doi.org/10.1007/s13225-023-00520-9

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

Fungi-based food
Medicinal mushrooms
Market value
Environmental biotechnology
Find a journal
Publish with us
Track your research

What is Medical Mycology?

Download PDF Copy

Introduction Medical mycology: A neglected area of microbiology? Evolution of anti-fungal therapies Recent advancements References Further reading

Fungal infections affect billions of people worldwide and, in severe cases can cause death. Yet medical mycology is a niche and neglected area of microbiology. Here we look at some common fungal infections, the therapeutic interventions used to treat them, how these have evolved and some of the more recent approaches that have served to aid our understanding of fungal pathology.

Medical mycology: A neglected area of microbiology?

The isolation and identification of fungi is an often-neglected area of medical microbiology. The science known as medical mycology began in the early 19 th century in Italy with the discovery of tinea favosa . The discipline concerns those infections in humans and animals that occur because of pathogenic fungi.

Fungi cause a range of diseases ranging from skin infections such as athlete’s foot, ringworm, dandruff, superficial cutaneous infections with dermophytes, to the more serious and invasive Candida and Aspergillus in severely immunocompromised hospital patients. Many important fungal pathogens seen in the immunocompromised also comprise part of the normal flora and this can lead to difficulties in detection.

It might seem surprising to learn that the global annual death toll due to fungal infections is greater than that for malaria, breast, or prostate cancer and more like those rates seen in tuberculosis (TB) and HIV (Gow, 2018). This toll runs to over a million people. In addition, around 10 million suffer from a severe fungal allergy; 100 million women annually fall foul to recurrent vulvovaginal infections and more than a billion people are afflicted by skin infections each year.

Candidas are ubiquitous in nature and can be found on many plants. They are also pathogens and saprophytes of both animals and humans that constitute the normal flora of the gut, the genitourinary tract, and the skin. The most common species in humans is Candida albicans but other species that can also cause trouble include G tropicalis, G glabrata, C krusei, G parapsilosis , and G pseudotropicalis . Of the several hundred Aspergillus spp. in nature there are just four linked to human pathogenicity: A fumigatus, A. flavus, A niger , and A. terreus .

Evolution of anti-fungal therapies

Gilchrist’s report of a medical case of blastomycosis saw the inception of medical mycology in the US in 1894. Over the course of the twentieth century further breakthroughs marked the discovery and characterization of dimorphic fungi, recognition of fungal pathogenicity and the part fungi play in systemic disease, the development of diagnostic tests in the laboratory, fungal classification, epidemiological and ecologic investigation.

The first kind of anti-fungal therapy, a saturated solution of potassium iodide (SSKI), turned out to be of limited clinical use and unsuccessful. The need for a broader spectrum anti-fungal, in addition to intravenous (IV) or oral anti-fungal agents was soon recognized, particularly as detection of fungal infections increased.

The first safe and effective anti-fungal treatments were discovered in Oxford in the late 1930s. In 1950 Hazen and Brown discovered nystatin, an important nod to the modern era of anti-fungal therapy. In the 1960s miconazole and clotrimazole were first introduced as topic agents. Then in the 1970s the broad-spectrum imidazole anti-fungals were developed and were active against dermatophytes, Candida, and other fungi.

Recent advancements

Researchers have now defined the immunopathology of many fungal diseases. The pathology may be driven by fungal invasion and virulence in some cases while in others it results from an over-active inflammatory response. Recent research has advanced our knowledge of immunopathology with respect to specific fungal diseases.

Meanwhile, next-generation sequencing has altered our understanding of the genes that predispose some patients to specific fungal infections. This in turn has advanced our knowledge about the fundamental mechanisms of immune surveillance and recognition.

In addition, we can now better appreciate the interrelationship between these processes and that of the modulatory influence enacted by the microbiome and the mycobiome. These new insights bring future hope for a vaccine, adjunct immunotherapy , and a personalised approach to the treatment of fungal infections.

Gillespie, S. et al. 1994. Medical Mycology. Medical Microbiology Illustrated . Pp. 113-124. Online: https://www.sciencedirect.com/topics/immunology-and-microbiology/medical-mycology .
Gow, N. et al. 2016. Medical Mycology and Fungal Immunology: New Research Perspectives Addressing a Major World Health Challenge. Phil. Trans. Roy. Soc., B.
Doi: 10.1098/rstb.2015.0462.
Gow, N. et al. 2018. Strategic Research Funding: A Success Story for Medical Mycology. Trends in Microbiology. Doi: 10.1016/j.tim.2018.05.014.
Gubbins, P. et al. 2009. Antifungal Therapy. Clinical Mycology, 2 nd Ed. pp. 161-195. Online: https://www.sciencedirect.com/topics/immunology-and-microbiology/medical-mycology.

Dr. Nicola Williams

Versatile science writer and content specialist (who can offer a unique historical twist too). I broadly focus on biology (including medicine), physics, and technology. I’m passionate about communicating the latest scientific research in an exciting, fresh, and accessible way. As a trained historian, I am also uniquely able to write content with a historical focus. I write about scientific news and research in a variety of formats, including articles, blogs, and scripts.

Please use one of the following formats to cite this article in your essay, paper or report:

Williams, Dr. Nicola. (2022, June 21). What is Medical Mycology?. News-Medical. Retrieved on September 14, 2024 from https://www.news-medical.net/health/What-is-Medical-Mycology.aspx.

Williams, Dr. Nicola. "What is Medical Mycology?". News-Medical . 14 September 2024. <https://www.news-medical.net/health/What-is-Medical-Mycology.aspx>.

Williams, Dr. Nicola. "What is Medical Mycology?". News-Medical. https://www.news-medical.net/health/What-is-Medical-Mycology.aspx. (accessed September 14, 2024).

Williams, Dr. Nicola. 2022. What is Medical Mycology? . News-Medical, viewed 14 September 2024, https://www.news-medical.net/health/What-is-Medical-Mycology.aspx.

Cancel reply to comment

Trending Stories
Latest Interviews
Top Health Articles

Vitamin D supplementation shows promising impact on cardiometabolic health

How can microdialysis benefit drug development

Ilona Vuist

In this interview, discover how Charles River uses the power of microdialysis for drug development as well as CNS therapeutics.

How can microdialysis benefit drug development

Global and Local Efforts to Take Action Against Hepatitis

Lindsey Hiebert and James Amugsi

In this interview, we explore global and local efforts to combat viral hepatitis with Lindsey Hiebert, Deputy Director of the Coalition for Global Hepatitis Elimination (CGHE), and James Amugsi, a Mandela Washington Fellow and Physician Assistant at Sandema Hospital in Ghana. Together, they provide valuable insights into the challenges, successes, and the importance of partnerships in the fight against hepatitis.

Global and Local Efforts to Take Action Against Hepatitis

Addressing Important Cardiac Biology Questions with Shotgun Top-Down Proteomics

In this interview conducted at Pittcon 2024, we spoke to Professor John Yates about capturing cardiomyocyte cell-to-cell heterogeneity via shotgun top-down proteomics.

Latest News

Antibody-drug conjugate shows promise in treating ovarian and endometrial cancers

Newsletters you may be interested in

Your AI Powered Scientific Assistant

Hi, I'm Azthena, you can trust me to find commercial scientific answers from News-Medical.net.

A few things you need to know before we start. Please read and accept to continue.

Use of “Azthena” is subject to the terms and conditions of use as set out by OpenAI .
Content provided on any AZoNetwork sites are subject to the site Terms & Conditions and Privacy Policy .
Large Language Models can make mistakes. Consider checking important information.

Great. Ask your question.

Azthena may occasionally provide inaccurate responses. Read the full terms .

While we only use edited and approved content for Azthena answers, it may on occasions provide incorrect responses. Please confirm any data provided with the related suppliers or authors. We do not provide medical advice, if you search for medical information you must always consult a medical professional before acting on any information provided.

Your questions, but not your email details will be shared with OpenAI and retained for 30 days in accordance with their privacy principles.

Please do not ask questions that use sensitive or confidential information.

Read the full Terms & Conditions .

Provide Feedback

IMAGES

Overview OF Mycology
SOLUTION: Fungi classification disease and its importance (Mb 3353
INTRODUCTION TO MYCOLOGY
Fungi, classification, Disease, and its importance (Mycology MB3353
Mycology 7
PPT

VIDEO

1. Mycology علم الفطريات
Importance of speciation in medical mycology
History of Mycology Part 1
Introduction to medical mycology in hindi
Anna Ronikier Importance of including type specimens in phylogenetic studies of myxomycetes
BLOSSOMS

COMMENTS

The importance of fungi and mycology for addressing major global challenges*
The time is right to make the world aware of the immense importance of fungi and mycology for sustainable global development, where land, water and biological materials are used in a more efficient and more sustainable manner. This is an opportunity for profiling mycology by narrating the role played by fungi in the bioeconomy.
(PDF) The importance of fungi and mycology for ...
The potential of fungi for a more sustainable world must be released. to address global challenges of climate change, higher demands on natural resources, and the increased burden of life-. style ...
The importance of fungi and of mycology for a global development of the
The vision of the European common research programme for 2014-2020, called Horizon 2020, is to create a smarter, more sustainable and more inclusive society. However, this is a global endeavor, which is important for mycologists all over the world because it includes a special role for fungi and fungal products. After ten years of research on industrial scale conversion of biowaste, the ...
Mycology What is it? Classification, in Medicine and How to Culture
Medical Mycology. Mycology is a wide field of study that is divided into several branches. This includes such divisions as forensic mycology, Ethnolichenology, and lichenology among others. These divisions allow mycologists to focus on specific areas of the field. In the food and alcohol industry, for instance, focus is directed towards the use ...
Celebrating the importance of mycological research
Celebrating the importance of mycological research. Although Kew is mostly known for its work on plants, a large part of the research is focused on the diversity and importance of fungi. Pepijn Kooij explains how mycologists at Kew are working to understand the role of fungi in plant diversity. By Dr Pepijn Kooij , Dr Tuula Niskanen and Dr ...
Introduction to Mycology
Mycology, the study of fungi, has shaped historical, cultural, and scientific narratives since ancient times. Journey through the ages, from fungi's mystical beginnings in prehistoric art to modern-day breakthroughs in medicine, technology, and sustainability. Explore the silent roles fungi have played in both tragic and transformative ...
Mycology 101: An Introduction to the Study of Fungi
Understand why fungi, often overlooked, are in fact indispensable to life as we know it. 1.1 What is Mycology: Definition and Overview. 1.2 The Importance of Fungi. 1.3 A Brief History of Mycology: From Discovery to Present Day. 1.4 Basics of Mycological Study. 1.5 Myths and Misconceptions about Fungi. 1.6 Fungi and Biodiversity.
Spotlight on: Mycology
There are two groups of people in mycology: those who specialised in it after doing a biology degree, but also many skilled amateurs and enthusiasts. We are indebted to them and their discoveries, and taxonomy is often in their hands. Their contribution is hugely important. Mycology is the study of fungi. It is closely associated with plant ...
Mycology
Toxicologists study mushroom and fungi for compounds which adversely affect other organisms. Pharmaceutical companies race to extract useful compounds from mushrooms. Careers in mycology are as diverse and complex as the field itself. Careers in Mycology. Mycology first became an important science in the agricultural industry, and remains so today.
Mycology
Mycology, the study of fungi, a group that includes the mushrooms and yeasts. Many fungi are useful in medicine and industry. Mycological research has led to the development of such antibiotic drugs as penicillin, streptomycin, and tetracycline, as well as other drugs, including statins.
PDF Kennesaw State University Department of Molecular & Cellular Biology
By the end of the course students will: Demonstrate knowledge in the basic understanding of the biology, taxonomy and phylogeny of fungi. Describe important impacts fungi have on humans and natural ecosystems. Describe the mechanisms and environmental conditions which facilitate fungal growth. Compare life histories of plant pathogens.
The importance of fungi and of mycology for a global ...
The vision of the European common research programme for 2014-2020, called Horizon 2020, is to create a smarter, more sustainable and more inclusive society. However, this is a global endeavor, which is important for mycologists all over the world because it includes a special role for fungi and fungal products. After ten years of research on ...
Introduction to Mycology
Welcome to "Introduction to Mycology: Exploring the Fascinating World of Fungi." This comprehensive course is designed to provide an in-depth understanding of fungi, their biology, ecology, and importance in various fields. Through a combination of engaging lessons, practical assignments, quizzes, and a final exam, you will embark on a journey to unravel the hidden mysteries of the fungal kingdom.
The importance of fungi and mycology for addressing major global
In the new bioeconomy, fungi play a very important role in addressing major global challenges, being instrumental for improved resource efficiency, making renewable substitutes for products from fossil resources. upgrading waste streams to valuable food and feed ingredients, counteracting life-style diseases and antibiotic resistance through strengthening the gut biota, making crop plants more ...
A Beginner's Guide to Mycology
Mycology is the study of fungus, which is an entire kingdom of life. Mycology is to fungi as botany is to plants. Most scientists consider mycologists to be a type of microbiologist. To date, scientists have named over 70,000 fungal species. Researchers believe there are up to 20 times that many fungus species out there.
Essays in mycology: Trends in Microbiology
Yeast. Genome. Mycology. Cambridge University Press, 1999. £57.50/$80.00 hbk, $21.95/$34.95 pbk (x + 377 pages) ISBN 0 521 56116 7 / ISBN 0 521 56784 X. Like many disciplines, mycology has had its ups and downs in the past 50 years. Currently, it is experiencing a decidedly exciting upward trend. In British universities, fungi were originally ...
The importance of fungi and of mycology for a global development of the
The importance of fungi and of mycology for a global development of the bioeconomy. The vision of the European common research programme for 2014-2020, called Horizon 2020, is to create a smarter, more sustainable and more inclusive society. However, this is a global endeavor, which is important for mycologists all over the world because it ...
Mycology
Mycology helps to differentiate between useful and harmful fungi and how the crops with fungal infections can be treated. Fungi produce various toxins that are harmful to other organisms. Most of the mushrooms produce toxins. While a few have medicinal properties. Yeat is another form of fungi which is used in brewing, distilling and bread making.
The importance of antimicrobial resistance in medical mycology
Cornely, O. A. et al. Defining breakthrough invasive fungal infection-position paper of the mycoses study group education and research consortium and the European Confederation of Medical Mycology ...
What is Mycology? A Rapidly Growing Field on the Study of Mushrooms
Mycology is the study of mushrooms, and it is a rapidly growing field with many intersections with other topics. From the classification and evolution of fungal species, to the use of magic mushrooms, to using mushroom mycelium for packaging material the field of mycology is diversifying. Historically mycology has primarily focused on fungal ...
The contribution of fungi to the global economy
Table 10 is the summation of the market values of fungal products and services that are economically of great importance. The contribution of fungi to the global market through fungal products and services is estimated to be USD 54.59 trillion. Table 10 Market value of fungal products and services by category.
What is Medical Mycology?
The isolation and identification of fungi is an often-neglected area of medical microbiology. The science known as medical mycology began in the early 19 th century in Italy with the discovery of ...
Mycology Essay
Mycology Essay. Mycology is an important and expansive area of biology that focuses on the study of fungi and their relationships to plants, animals, and the environment. ... Mycology is the study of fungi, according to a definition provided by the University of Toronto's Lab of Molecular and Cellular Biology (UofT). Fungi are a diverse group ...

The importance of fungi and mycology for addressing major global challenges

Abstract and Figures

Mycology What is it? Classification, in Medicine and How to Culture

Classification

Mold Fungi (Filamentous/Hyphal)

Geographical Classification

Epidemiologic Grouping

Observation

Recent Articles

Celebrating the importance of mycological research

Estimating numbers

How to investigate fungal diversity

Plant-fungal symbiosis and environmental change

Fungus-farming ants

References

Learn Mycology

Course Outline

The Basics of Fungi

Fungal Biology

Fungal Ecology

Human Uses of Fungi

Fungal Diseases

Fungal Conservation

Advanced Topics in Mycology

Amazon Storefront

Spotlight on: Mycology

Why is mycology important?

What careers are available?

How do I start?

Where can I get more information?

Web resources

At a glance

What led you to study fungus?

Describe a typical day?

What are you working on now?

What are the potential applications of your work?

Why are there so few mycologists?

Mycology Definition

History of Mycology

Careers in Mycology

Cite This Article

Save citation to file

Add to My Bibliography

The importance of fungi and of mycology for a global development of the bioeconomy

Similar articles

Related information

Introduction to Mycology

Note: This Living-Course is designed for Patrons who are curious about mycology and have a basic understanding of biology. No prior knowledge of mycology is required.

Target audiences

View Mycology Courses

The Mushroom Network is proudly affiliated with: SPORE ZE WORLD | Green Grass Grove | 3G-Camping Retreat | 3G-Prints | Project Aurora .

The importance of fungi and mycology for addressing major global challenges

Introduction to the Importance of Mycology

The Role of Fungi and Mycology in Addressing Major Global Challenges

The Importance of Mycological Research for Further Biological Solutions to Important Problems

Inspiration and Learning from Nature

The New Potential of Fungi and Fungal Products from Biorefineries

New Meta-Level Discoveries

New Sequence Analysis Tools, Making Order and Sense Out of Chaos

Fungal Inoculum for Stronger and More Robust Plant Growth

New Breakthroughs Needed

How to Strengthen International Mycology for a Biobased Future

Author information

Corresponding author

Additional information

Rights and permissions

About this article

Share this article

A Beginner's Guide to Mycology

How Are Mycology, Mycorrhizae, and Mycelia Different?

The Origins of Mycology

Trends in Mycology

What Makes a Fungus?

Areas of Mycology

Edible Fungi

Mychorrizae and the Environment

Medical Mycology

Pathogenic Fungi

Entheogenic (a.k.a. Magic) Mushrooms

Why Aren’t There More Mycologists?