Fungal Literature Sampler
The scientific literature on fungi and fungal biotechnology is rich and growing quickly. We’ve put together and summarized a sampler of papers on general views of fungi and how people can work with fungi to solve problems, with specifics on mycelium materials and Ganoderma fungi useful for mycomaterials. The bibliography is far from exhaustive, but offers points of entry to key ideas and lines of research to follow farther.
Most of the papers are open access. If a paper isn’t open, it’s here because it has been especially helpful.
Download a full PDF of Open Fung’s Fungal Literature Sample here.
General fungal reviews
“Fungal impacts on Earth’s ecosystems”
Fungi are a vast and ancient group of organisms – currently estimated at upwards of two million species after a billion years of evolution. This paper is an extensive review of how these fungi can impact the planet, the beneficial and harmful ways fungi can impact humans, and how humans are trying to work with fungi to address problems.
“The Mycelium as a Network”
Fungal hyphal cells can form a mycelium network that is complex, communicative, dynamic and adaptive with impacts from the micron- to the ecosystem scale. This paper reviews the network properties of mycelium, covering biochemical, microscopy, molecular and mathematical modeling methods for understanding. The graphics are particularly strong.
“The contribution of fungi to the global economy”
Fungi provide valuable products and environmental services to people, but we don't yet have conclusive estimates capturing how much money all of these fungal products and services are worth. The authors of this paper describe different economic roles fungi play, then draw from many data sources and modeling methods to estimate that fungi contribute 54.57 trillion US dollars per year to the global economy.
“Mycorrhizal mycelium as a global carbon pool”
The close relationships between many land plants and mycorrhizal fungi are strong drivers of carbon movement through global environments. Using almost 200 datasets, the authors estimate that about 13.12 Gt of carbon dioxide equivalents fixed by plants are, at least temporarily, stored underground in mycorrhizal fungal mycelium per year – over a third of the quantity of annual carbon dioxide emissions from fossil fuels.
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“Mycology: A neglected megascience”
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Hawksworth, D. L. (2009). In: Applied Mycology. TEXT AVAILABLE - Discussion from a foundational academic mycologist about why the study of fungi hasn’t alwayreceived
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Fungal applications and biotechnology: reviews and theory
“Partnerships with Indigenous Peoples for an ethical bioeconomy”
Not specifically about fungal biotechnology, but covers issues of access and benefit-sharing, collaboration and engagement with Indigenous and local communities that touch all biotechnologists reaching towards an ethical and sustainable bioeconomy.
“Unlocking the magic in mycelium: Using synthetic biology to optimize filamentous fungi for biomanufacturing and sustainability”
Informative review of how filamentous fungi – those that grow in threadlike hyphal cells and form mycelium, as opposed to single-celled fungi, like yeasts – can be applied in the bioeconomy and the environment, and how we might boost those contributions with cutting-edge synthetic biology.
“Leather-like material biofabrication using fungi”
An excellent review of the promise and challenges of growing leatherlike materials using mycelium. My favorite line: “Inherent biological variation in fungal growth makes uniform thickness and surface texture difficult to achieve in mycelium-derived leather-like materials.”
“Growing a circular economy with fungal biotechnology: A white paper.”
This call to action covers how we can incorporate fungi into a circular bioeconomy and how fungi can help meet UN sustainable development goals. If you only read one paper on this list, this could be the one to pick!
“A review of the material and mechanical properties of select Ganoderma fungi structures as a source for bioinspiration”
There are a lot of solid reviews about fungal biomaterials out there, but I find the slant in this one to be unique and extremely helpful. It focuses on the properties of Ganoderma fungi, and the paper’s graphics showing the structure of Ganoderma, its three different types of hyphal cells and spores are extra enlightening.
“Fungi Fabrics and Living Colors: Toward Ecocentric Biodesign?”
A thought-provoking consideration of how we can incorporate living beings into designed materials without repeating the same patterns that led to our overlapping environmental crises. Can we acknowledge the centrality of non-human living things in biodesign? Gives three examples of ways to do this: rethinking “biomimicry,” growing leather from responsive mycelium, and dying fabric with bacteria that co-determine how the colors turn out.
“Current state and future prospects of pure mycelium materials”
A great overview of materials made from fungal mycelium.
Fungi and art
“Merging science and art through fungi”
Historically, art and science have been complementary facets of the same endeavors. Here, pioneering fungal biotechnologist and artist, Vera Meyer, writes about blending art, culture, community, science and technology, and how fungi can bridge these fields.
“The beauty and the morbid: Fungi as source of inspiration in contemporary art”
Instances of fungi inhabiting the places where art and science meld.
Mycomaterial research
“Fabrication factors influencing mechanical, moisture- and water-related properties of mycelium-based composites”
Study showing how different fungi, substrates and processing methods impact the characteristics of composite materials made of mycelium and substrate.
“Development of a circular sustainable culturing process for natural leather-like materials based on fungal mycelium”
Interesting project looking at the growth impacts of fungal strain and fungal substrate (and accidentally, other microbes!) on a very specific process for making mycelium mats. Not as much about the impact of environmental conditions, but cool.
“Uncovering the transcriptional landscape of Fomes fomentarius during fungal-based material production through gene co-expression network analysis“
Cutting-edge study of gene transcription of the polypore basidiomycete fungus, Fomes fomentarius, on different substrates and in different conditions relevant to composite biomaterial growth. Presents exciting and unique insights that may also relate to the genetics underpinning growth and biomaterial processes using related fungi.
“Collection and characterization of wood decay fungal strains for developing pure mycelium mats”
Comparison of the pure mycelial mats grown from 21 different strains of wild-type Italian wood-decay fungi in liquid and semi-solid growth media, concluding Fomitopsis iberica performed best.
“Establishment of the basidiomycete Fomes fomentarius for the production of composite materials”
Study testing a well-performing wild-type strain of the basidiomycete polypore, Fomes fomentarius, on different substrates, then characterizing the mechanical properties and electron micrographs of the resulting composite mycomaterials.
“Mycofabrication of mycelium-based leather from brown-rot fungi”
Study comparing the effects of different types of wooddecay fungi, growth conditions and post-growth processing regimes on the properties of pure mycelial leatherlike fungal mats. Mats showed leatherlike properties, and high performance was observed after post-harvest treatment with 20% polyethylene glycol.
“Life cycle assessment of MycoWorks’ ReishiTM: the first low-carbon and biodegradable alternative leather”
Cradle-to-gate assessment of MycoWorks mycelium leatherlike material, showing improvement in sustainability over a cow leather benchmark in several current and planned iterations.
Mycomaterial patent landscape reviews
“Recent technological innovations in mycelium materials as leather substitutes: a patent review”
Overview of the rapidly growing number of patents for leatherlike materials made from mycelium, covering fungal species, biomass generation and post-processing conditions. Highlights gaps in knowledge.
“Patent landscape analysis for materials based on fungal mycelium: a guidance report on how to interpret the current patent situation.”
To reduce uncertainty and highlight areas open for sustainable innovation in the field, this review aims to summarize the patent landscape for composite mycelium-based materials, particularly those for construction and packaging.
Ganoderma biology, phylogeny, cultivation
“Mushrooms for the tropics: Growing Ganoderma”
Practical overview of Ganoderma and methods for growing it.
“Genome sequence of the model medicinal mushroom Ganoderma lucidum”
This is the first full-genome sequence published for a Ganoderma. It details genetic features: 13 chromosomes, 43.3 Mb size, ~16,000 predicted protein coding genes, as well as information about differences in gene expression between vegetative mycelium, primordia and mature sporocarps.
“Elucidating “lucidum”: Distinguishing the diverse laccate Ganoderma species of the United States”
Many evolutionarily distinct Ganoderma species are sometimes described by the names “reishi,” “lingzhi” or “Ganoderma lucidum,” but they can behave quite differently when growing mycomaterials. This indispensable paper uses DNA sequencing and multi-locus phylogenetic analysis to investigate this diversity. Reports 18 phylogenetically supported species of shiny Ganoderma within the lucidum complex, geographic range info for US taxa, including an intercontinental introduction, and physical characteristics.
“Cultural characterization and chlamydospore function of the Ganodermataceae present in the eastern United States”
Compares several isolates of each of nine species of Eastern US fungi in Ganoderma lucidum complex plus one related fungus. The authors report which ones make chlamydospores (tough resting spores) in culture, and calculate curves showing growth rate on agar plates over a wide range of temperatures. G. sessile and Tomophagus colossus grew fastest, G. meredithiae, G. ravenelii, and G. tsugae grew slowest, and those that made chlamydospores were able to survive high temperatures.
“Global diversity of the Ganoderma lucidum complex (Ganodermataceae, Polyporales) inferred from morphology and multilocus phylogeny”
Similar to the Loyd et al. 2018 “Elucidating lucidum” paper, this earlier paper also takes an approach combining multi-locus DNA sequence phylogeny and morphology to show 13 supported species within the global “Ganoderma lucidum complex.”
Ganoderma genetic engineering
“The Cas9-gRNA ribonucleoprotein complex-mediated editing of pyrG in Ganoderma lucidum and unexpected insertion of contaminated DNA fragments”
Demonstrates successful genetic modification of the pyrG gene in Ganoderma using a CRISPR method, where the Cas9 protein and guide RNA are delivered together as a ribonucleoprotein complex. Interestingly, the authors also report that unexpected fragments of DNA from the host and the E. coli bacteria used to produce the Cas9 protein can be incorporated to the target site.
“CRISPR-Cas9 assisted gene disruption in the higher fungus Ganoderma species”
First CRISPR gene editing paper for Ganoderma. The authors successfully disrupted the ura3 gene by a method of expressing the Cas9 protein and guide RNA inside the Ganoderma, rather than delivering a ribonucleoprotein complex, as has become common in later studies.
“Targeted gene insertion and replacement in the basidiomycete Ganoderma lucidum by inactivation of nonhomologous end joining using CRISPR/Cas9”
Inactivated the non-homologous end joining machinery, a method for the cell to repair breaks in its DNA that cut through both DNA strands, in their Ganoderma strain by disrupting the ku70 gene using a CRISPR method. This genetic change allowed the authors to then insert and replace genes with much higher efficiency at the target site.
“Development of a 2A peptide-based multigene expression system and its application for enhanced production of ganoderic acids in Ganoderma lucidum”
Increasing levels of gene expression and expressing genes that did not originate in the target organism can be greater engineering challenges than reducing or eliminating gene expression. In this paper, authors employed required native intron sequences and a short viral peptide that allows multiple proteins to be expressed from a single mRNA sequence – the 2A peptide system – to simultaneously upregulate two genes in Ganoderma, leading to increases in medically significant ganoderic acids. This approach also allowed them to co-express multiple genes that did not originate from Ganoderma: two anti-fungal resistance genes and a fluorescent protein.
“Increased production and anti-senescence activity of exopolysaccharides in Ganoderma lingzhi by co-overexpression of β-1,3-glucan synthase and UDP-glucose pyrophosphorylase”
An elegant example of metabolic engineering in a basidiomycete fungus in which increased expression of two genes simultaneously in Ganoderma led to an increase in specific molecules thought to have human health benefits, and the increased bioactivity of extracts from the modified Ganoderma strain.
“Efficient expression of heterologous genes by the introduction of the endogenous glyceraldehyde-3-phosphate dehydrogenase gene intron 1 in Ganoderma lucidum”
Fascinating paper showing that native gene introns can be necessary for successful expression of genes inserted in Ganoderma that originated from another organism. Adds to the increasing body of literature showing that native introns may be necessary for expression of some outside genes in basidiomycete fungi – once again, the mushrooms are quirky and have cooler genetic regulation mechanisms than we knew!
“Breeding a new Ganoderma lucidum strain with increased contents of individual ganoderic acids by mono–mono crossing of genetically modified monokaryons”
Many basidiomycete, mushroom-forming fungi have an “n+n” genetic system: genetically distinct parent nuclei, each with one copy of each chromosome, can persist in a heterokaryotic individual, making the pursuit of genetic modifications complicated. In this paper, the authors genetically modify two different monokaryotic strains, then mate them to obtain a heterokaryon with increased amounts of medically significant ganoderic acid.
Complementary fungal research
“Hydrophobin gene deletion and environmental growth conditions impact mechanical properties of mycelium by affecting the density of the material”
Demonstrates the effects that genetic modification and changing temperature and CO2 concentration during growth can have on Schizophyllum commune mycelium. Most excitingly, the authors demonstrated that deleting a single hydrophobin gene among the many present in S. commune changed the properties of its mycelium from being similar to natural materials to being more similar to thermoplastic.
“Spore-derived isolates from a single basidiocarp of bioluminescent Omphalotus olivascens reveal multifaceted phenotypic and physiological variations”
Cool paper showing the variability in luminescence, growth rate, pigmentation, chemical composition and production of specific compounds with antibiotic and anticancer properties among 47 haploid, monokaryotic strains derived from the spores of a single Omphalotus olivascens mushroom.
“Enabling community-based metrology for wood-degrading fungi”
What is a globally available, highly standardized, economical basis for wood-decay fungal growth medium you may ask? Pringles™ ! In this paper, Open Fung’s Rolando Perez and co-authors share Pringles™ -based growth substrate and measurement methods that reduce reductions in reduced variability in comparing fungal growth between labs, another example showing that developing fungal applications can be remarkably accessible.
“Transformation of compatible mating-type genes in monokaryons triggers fruiting body development by activating mating pathways in Pleurotus eryngii.
Fruitbody formation in many fungi is dependent on having different genes at one or two specific mating loci in a heterokaryotic individual – one with nuclei contributed by two or more parents. In this fascinating paper, the authors transform monokaryotic, haploid Pleurotus eryngii, the king oyster mushroom, with compatible mating genes, allowing the monokaryons to form fruitbodies, but not basidiospores. Has interesting implications for mushroom breeding and understanding the genetics of basidiomycete mating.