Growing Tomatoes in the World of Tomorrow

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By Joseph Anderson

Despite the enormity of the planet we live on, there is an even bigger space for life to develop and exist right under our very noses. I’m not talking about the 4th dimension or the faerie realm; I am talking about the phyllosphere. You see, microorganisms can live inside the leaves, stems, and roots of plants. Overall, the surface area of the phyllosphere is twice as large as the surface area of all land on earth (Zimmerman and Vitousek 2012). Humans have billions of bacteria living inside our bodies to help us live. Plants have this same relationship, but with microbial fungi! Within these plants lie a diversity of life that is a new frontier in science.

Nematode

A picture of a Nematode. Most of them have a similar morphology.
http://enrichla.org/wp-content/uploads/2012/06/nematodes.jpg

As we start to look into this microcosm of life, we find that these fungi are far from dormant; many of these fungi play important symbiotic roles with the plants.  Nematodes are a soil microorganism found abundantly in most terrestrial soils. Although most of them survive by eating fungi, many of them are considered agricultural pests (Sikora et al. 2008). We have recently found several species of endophytic fungi that decrease the plant damage caused by nematodes (Sikora et al. 2008, Singh et al 2013). These recent studies have been proving the efficacy of using the beneficial endophytes as a natural biocontrol method for agriculture.

Endophytes get even cooler though. Since the discovery of endophytes, we have been looking at what roles they play in their plant hosts. What scientists are finding will surprise you. One scientist from Seattle has been looking at the endophytes that exist in plants that grow by hot springs. The soil in these environments can reach temperatures in excess of 100 degrees Fahrenheit, which is usually beyond the range at which most plants or fungi can survive (Tennesen, 2010). When the plant and its fungal partner are separated from each other, neither can survive in such extreme temperatures. So we know that this heat tolerance is an emergent property of the two species. That means that this trait only exists when the two different species are existing mutualistically. Dr. Rodriguez then took the endophyte from the extremely hot soil and inoculated a tomato plant with them. The tomato with the endophyte can now survive in soil temperatures over 140 degrees Fahrenheit. Later studied showed that this mutualism is actually a 3-way symbiosis (Rodriguez and Redman, 2008). For the endophyte to confer heat tolerance to the host plant, the endophyte needs to be infected with an RNA virus. Similar studies have looked at plants that live in coastal areas, where salt tolerance is something plants need to survive. For some species of plant, the salt tolerance is an emergent property of another fungal symbiosis (Rodriguez and Redman, 2008). The deeper we look into endophytes, the more complexity and interconnectedness we see.

I know this is completely mind blowing, but this rabbit hole keeps going. Are you ready to dive in? The same species of fungi that confers the heat or salt tolerance to the plants are found in other, low-stress environments. However, the endophytes that live in low-stress environments do not confer the same stress-tolerance that the high-stress endophytes do (Rodriguez and Redman, 2008). This means that the endophyte responds differently with its plant partner in different habitats. Basically, the fungus is helping the plant survive in whatever environment the symbiosis developed in. But wait, there’s more: the same species of fungi can be symbiotic, like we have been discussing, or it can be parasitic, depending on the plant species (Rodriguez and Redman, 2008). In some cases, like tomato plants, it can even come down to the variety of tomato. This is stumping scientists. What makes these endophytes choose mutualistic versus parasitic lifestyles? Dr. Rodriguez thinks it can come down to a single mutation.

During the last 5 massive extinctions that our planet has gone through, natural historians have noted an increase in fungal diversity immediately afterwards. Some environmental scientists theorize that we are going through the 6th major extinction event right now. How are we going to survive this time of extreme change on our planet? Paul Stamets, A forefront mycologist of our times, says that the creatures that ally with fungi survive catastrophes. For generations entire societies have experienced mycophobia, the irrational fear of mushrooms. It is time to transcend the mindset of fear and learn to love fungi, the same way we love the beauty of orchids or the comfort of your domesticated cat. We now know about the symbiotic systems flourishing between plants and fungi. Imagine the kind of emergent properties we could experience once we embrace our fungal partners for the good of the world.

Bibliography

Rodriguez, R. and Redman, R (2008, 19 June). More than 400 million years of evolution and some plants still can’t make it on their own: plant stress tolerance via fungal symbiosis. Journal of Experimental Botany 59(5), 1109-1114.

Sikora, R.A., Et al (2008, July). Mutualistic endophytic fungi and in-planta suppressiveness to plant parasitic nematodes. Biological Control, 46(1), 15-23.

Singh, U.B. Et al (2013, January). Can endophytic Arthrobotrys oligospora modulate accumulation of defence related biomolecules and induced systemic resistance in tomato (Lycopersicon esculentum Mill.) against root knot disease caused by Meloidogyne incognita. Applied Soil Ecology 63, 45-56.

Tennessen, M. (2010, May). More Food From Fungi? Scientific American, 302(5), 27-28.

Zimmerman, N. B. and Vitousek, P.M. (2012, 12 June). Fungal endophyte communities reflect environmental structuring across a Hawaiian landscape. PNAS, 109(32), 13022-13027.

Symbiotic Relationships and Fungus Examples

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By: Tanya Lynch

I’m sorry to say, but Science Fiction has lead us astray. Because of television shows like Stargate SG1 and characters like Teal’c (who carries a symbiote Goa’uld) many believe a symbiotic relationship is a relationship in which both parties involved benefit. Alas, this is not always true. In fact, symbiotic simply means a close, prolonged association between two or more different organisms of different species.

symbiosis chart

Source: http://www.yorku.ca/tnoel/march5/Symbiosis.html

There are actually five different types of symbiotic relationships: Parasitism, commensalism, mutualism, neutralism, and competition. Of these, parasitism and mutualism are the most common relationships formed by fungi.

Parasitism is when one species negatively affects the second species in the relationship. A tapeworm can live in a parasitic symbiotic relationship with a cow. The flea is a parasite on the dog. Corn smut, Ustilago maydis, is a prime example of fungal parasitism of corn.

Ms Maize
Photos: Corn – Dieter Spears/iStockphoto.com. Corn Smut Spores – APS Press. Fake Facebook status created with www.statusclone.com

Ustilago maydis, creates tumor-like growths mixed within the kernels of corn on the stalk. This fungus turns a cheery yellow ear of corn into a deformed and grey mass.  These grey tumor-masses are actually kernels that have been infected by U. maydis and filled with teliospores (a type of reproductive cell). In all but central Mexico, this fungus is considered a bothersome disease, but there it is quite the culinary delicacy (The American Phytopathological Society, 2006).

Another example of a parasitic relationship is that of the genus Cordyceps (of which there are many species) and a poor insect host (E.B. Mains, 1958). In this most unfortunate relationship a spore will land in some fashion on a fly and germinate, then stromata (a visible clavate or sometimes branched structure coming out of insect body segments) will form outside of the body. These structures contain the sexual components of the fungi which will release spores when mature. Prior to the release of spores, a particular species in this family, Ophiocordyceps unilateralis, will use neurotoxins to control the movement of ants to climb to the highest part of a grass and bite down to hold on (Harry C. Evans, 2011). This allows the spores to eventually be released in the most prime conditions and location for eventual germination on another unsuspecting ant.

Things are much prettier on the mutualism side of things. The concept of mycorrhizal associations between a fungal partner and a plant partner is the mutualistic symbiosis most commonly referred to when talking about fungi.

Doug Fir

Photos: Fir Seedling – Ingrid Barrentine, Northwest Guardian. Microscope Ectomycorrhizae – http://sciweb.nybg.org/science2/hcol/mycorrhizae2.asp.html Fake Facebook status created with www.statusclone.com

In a mycorrhizal partnership a fungal partner will take hold onto the roots of some plant partner and each side will benefit in some way from the association. For instance, the fungus growing around (or in some cases in) the roots of the plant allows for greater surface area of those roots. This greater surface area allows for greater access to water. The fungal partner also helps to shuttle certain nutrients into the plant’s roots (The New York Botanical Garden, 2003). In return, the plant shares some of the carbon created during photosynthesis.

A symbiotic relationship isn’t only a beneficial relationship for both partners. As described above in the fungal examples, there are different types of symbiotic relationships that aren’t positive at all. So before you write your sci-fi novel or television series scripts, be sure to remember these awesome fungi and take into account the diversity of symbiotic relationships and what they look like!

Cordyceps

Sources:

The American Phytopathological Society. Common smut of corn. (2006). Retrieved from http://www.apsnet.org/edcenter/intropp/lessons/fungi/Basidiomycetes/Pages/CornSmut.aspx

The New York Botanical Garden. Hidden Partners: Mycorrhizal Fungi and Plants (2003). Retrieved from http://sciweb.nybg.org/science2/hcol/mycorrhizae2.asp.html

E. B. Mains. North American Entomogenous Species of Cordyceps. Mycologia , Vol. 50, No. 2 (Mar. – Apr., 1958) , pp. 169-222

Evans, H. C., Elliot, S., & Hughes, D. Hidden Diversity Behind the Zombie-Ant Fungus Ophiocordyceps unilateralis: Four New Species Described from Carpenter Ants in Minas Gerais, Brazil (March 2, 2011). Retrieved from http://www.plosone.org/article/info:doi/10.1371/journal.pone.0017024

 

Fungi Flashlights for Fairies (Bioluminescent Fungi)

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By Shelby Burnham

Aristotle (384-322) reported a mysterious light, distinct from fire, emanating light from decaying wood. Pliny the Elder (23-79 AD) also mentioned feasting on a glowing, sweet fungus found on trees in France (Kimbrough, L). Glow in the dark mushrooms are unique to the fungal kingdom for their mysterious “cold light”; bioluminescent fungi will continue to intrigue generations to come.

The most common reports about bioluminescence are “glowing wood” in forests at night. The most abundant and diverse bioluminescence are found in tropical rain forests, they are typically small and grow on leaf litter or rotting wood (Herring, P). Some countries are great places to hunt for bioluminescent fungi, these countries include: Brazil, Dominican Republic, Jamaica, Japan, Malaysia and Puerto Rico (Pescovitz, D). In these countries, scientists look not up at the stars but down to see a starry filled ground. The temperate forests in North America also have bioluminescent fungi. Two examples of bioluminescent fungi that grow in temperate forests of North American are: Omphaloutus olearius (the jack-o-lantern mushroom), and Armillaria mellea (the honey mushroom). O. olearius caps and gills glow in the dark but only the mycelium of the A. mellea glows in the dark (Kimbrough, L). All bioluminescent fungi emit light 24 hours a day but are only luminescent at night, preferably on a new moon night (Herring, P).

Photo by: National Geographic / Darlyne A. Murawski

Scientists suggest that mushrooms illuminate different organs for different reasons. Some evidence shows that certain nocturnal insects and mammals are attracted to the glowing light and therefore assist the fungi in spore dispersal. There is controversy over the idea that illuminating is more effective. Most bioluminescent species are found in the water, they use their luminescent tricks for communication, hunting and camouflage but their molecular structure is very different than a mushroom (Untamed Science). In the article called “Freaky Fungi Glow in the Dark” from the University of Sao Paulo, the authors state that there are 85,000 described species in the Fungi Kingdom, and only 65 species in the ent

ire Fungi Kingdom are known to be luminescent. All of the 65 luminescent fungi species are mushrooms that have thin-walled, white spores, and all are white rot fungi capable of digesting both cellulose and lignin in plant debris. All luminescent fungi are in the Basidiomycota phylum, and 70% of them are Mycenas. At the San Francisco State University, professor and avid mycologist Dr. Dennis Desjardin has discovered over 200 new species of fungi, and nearly 1/4 of all the luminescent fungi (Desjardin, D).

How does bioluminescence work? Bioluminescence are the result of a molecule called luciferin coming in contact with water and oxygen. When water and oxygen are prese

nt, the molecule luciferin is activated and thus becomes a luciferase enzyme. This reaction produces energy, and this energy excites the electrons in the luciferin molecule (Hastings, J). This means, the electrons get a little more energy than normal. When the molecules return to their normal state the extra energy gets converted into light. All the energy that bioluminescent mushrooms produces get turned into light, this light is more efficient than a flashlig

ht because they do not produce heat, just pure light. This is where they get their nickname from “Cold Light Mushrooms” (Untamed Science).

A quote from Dr. Dennis Desjardin at the San Francisco University: “There are a few species in temperate habitats that are luminescent, particularly Omphalotus species (jack-o-lantern mushrooms), but they are not very spectacular and it requires patience (sitting for a long time in the dark to let your eyes adjust). The tropics are the best place to see them. Brazil, Southeast Asia and Australia are particularly good places with a variety of luminescent mushroom species. Pick a new moon night during the rainy season, go out at night when it is completely dark, and roam in the forest looking for points of yellowish green light. Donʼt forget, however, that you cannot see a thing in the pitch dark, so youʼll walk into trees, streams, etc, and that there are lots of creatures that are nocturnal, like poisonous snakes, jaguars, necrotic spiders, etc. that need to be avoided! Itʼs lots of fun!” (Kimbrough, L).

This is a video called the “Science of Glowing Mushrooms”

References

Denk, J. (n.d.). Fire fox mushroom (honey mushroom, armillaria mellea). Retrieved from http:// morelmushroomhunting.com/newsletter_july_2009.htm

Desjardin, D. E. (2008). Freaky fungi glow in the dark. LiveScience & National Science Foundation,

Hastings, J. W. (1996). Chemistries and colors of bioluminescent reactions. Elsevier Science,

Herring, P. J. (1994). Luminous fungi. Institute of Oceanographic Sciences, Deacon Laboratory, Brook Road, Wormley, 8(4), 18.

Kimbrough, L. (2013). Why bioluminescent fungi glow in the dark.

Murawski, D. A. (n.d.). Jack-o-lantern mushrooms (omphalotus olearius) glowing green at night.. Retrieved from http://www.artflakes.com/en/products/jack-o-lantern- mushrooms-omphalotus-olearius-glowing-green-at-night

Pescovit, D. (2009). Glow-in-the-dark mushrooms.

Science of Glowing Mushrooms [Web Video]. Retrieved from http://www.youtube.com/user/ UntamedScience?feature=watch

Zombie Ant Fungi

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By Savannah Richard

Not much attention is given to non-agricultural related insect pathogens and parasitic fungi, making these fungi all the more mysterious and fascinating. Ophiocordyceps unilateralis is a fungal pathogen that parasitizes its host; in this case Camponotus leonardi ants and other close relatives are the victims. Ophiocordyceps unilateralis is in the Ascomycota phylum and was previously in the Cordyceps family but has recently diverged into its own genus (Pontopiddan et al 2009). This fungus can be found in most tropical environments from South America, Africa to Eastern Asia.

Ants become infected when they are on the forest floor foraging and spores, laying on leaves or other dead ants, attach to the exoskeleton. These spores penetrate the ant’s exoskeleton to Zombie antseat away at the soft tissue inside. During this process of feeding off the ants tissue the fungus produces compounds that affect the ant’s brain and subsequently change the ant’s behavior. The exact cause is still unknown but the infected ants crawl up a tree and attach themselves using their mandibles to the underside of a leaf. Mycelia then grow through the bottom exoskeleton of the ant onto the leaf to help secure the ant (Anderson et al 2012). Ants have also been found to position themselves facing north almost exactly 25cm from the forest floor; in high humidity areas. These conditions are ideal for fungal growth and are attributed to the fungal pathogens control of the ant’s behavior, hence the nickname zombie fungi (Andersen et al 2009).

After the ant has attached itself to the leaf and has died, the fungus begins the last part of its life cycle. A wiry stalk grows out of the ant’s head that has perithecia (spore bearing surface) at the ends, as shown in this video: http://www.youtube.com/watch?v=XuKjBIBBAL8. The fungus then drops the spores back onto the forest floor and waits for another host to pass through to infect. The entire life cycle of Ophiocordyceps unilateralis takes around 4-10 days. This fungus is also known to attack large groups of ants and almost wipe out entire colonies (Anderson et al 2009). This leaves behind several ant corpses on the underneath of leaves which scientists have coined as “ant graveyards”. Ants have adapted ways of sensing graveyards and avoiding these areas in the forest. With this developed sensitivity to Ophiocordyceps unilateralis ants have also been shown to carry out other infected ants into the forest away from the rest of the colony (Pontoppidan et al 2009).

All Cordyceps species including Ophiocordyceps species are thought to have medicinal properties. Cordyceps species produce a bio-metabolite called cordycepin, which has anti-inflammatory and anti-tumor properties. Cordyceps also contain many nutritional compounds such as B1, B2, B12, and K vitamins. In traditional Chinese medicine Cordyceps were used to aide in bronchitis, asthma, respiratory and cardiovascular diseases. Pharmaceutical companies have been able to isolate cordycepin and turn it into powder form for mass production. Culturing Cordyceps is now being experimented with to some degree of success and may expand in availability sometime in the near future (Tuli and Sandhu 2013).

References:

Andersen, S., Ferrari, M., Evans, H., Elliot, S., Boomsma, J., Hughes D., (May 2012)

Disease Dynamics in a Specialized Parasite of Ant Societies. Plos One. Retrieved from: http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0036352

Andersen, S., Gerritsma, S., Yusah, K., Mayntz, D., Hywel-Jones, N., Billen, J., Boomsma, J., Hughes, D. (September 2009) The Life of a Dead Ant: The Expression of an Adaptive Extended Phenotype. The American Naturalist, Volume 174, No.3 Retrieved from http://www.jstor.org/stable/10.1086/603640

BBC Worldwide. Cordyceps: attack of the killer fungi- Planet Earth Attenborough BBC wildlife. ( November 2008) Retrieved from: http://www.youtube.com/watch?v=XuKjBIBBAL8.

Pontoppidan, M., Himaman, W., Hywel-Jones, N.,  Boomsma, J., Hughes, D., (March 2009) Graveyards on the Move: The Spatio-Temporal Distribution of Dead Ophiocordyceps-Infected Ants. Plos One. Retrieved from: http://www.plosone.org/

Tuli, H., Sandhu, S., (February 2006) Pharmacological and Therapeutic potential of Cordyceps, with special reference to cordycepin. 3 Biotech, volume 4, issue . Retrieved from: http://link.springer.com/article/10.1007/s13205-013-0121-9

Neurospora crassa – the Ultimate Genetic Puppet

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By: Nils Nelson

Since the rise of evolutionary theory stemming from Darwin in the 1840s the mystery of the exact processes of inheritance and evolution has begun to unfold. Though it was not until 1970 that the first organism’s genome was sequenced, there was a trail of key developments in proving genes existence and purpose that resulted in the now immense field of genetics and our understanding of biology today.

In 1941 George Beadle and Edward Tatum published a paper on the relationship of genes to enzymes in the fungus Neurospora crassa first reported in French bakeries in 1843 (Beadle, Tatum. 1941). This became known as the one gene-one hypothesis, which later became the one gene-one polypeptide theory. This discovery became the foundation for solving genetic code and brought the fields of biochemistry and genetics together. Beadle and Tatum proved their hypothesis by first mutating sexual spores of the fungus with x-rays and ultraviolet light then isolating mutated spores. Next they selectively bred compatible strains with mutants then grow the resulting genetic offspring on nutrient deprived agar growth medium. Carefully they added the precursors for a specific nutrient and track when an enzyme was not present to convert the pre product molecules. This led them to be able to narrow down the defect of a single gene to the absence of a single enzyme. Today there is a current project attempting to produce a knockout mutant strain of N. crassa mutant for each gene in the genome.

This was not the end of the road for scientific insights provided by N. crassa. The ability to determine the phenotype of all four products of individual meiosis granted insight into the function of genetic recombination in all eukaryotes and became principle evidence for the adaptive function of recombination and out crossing (Murray, 1960). This fungus gave us the first evidence for gene conversion which is a correction mechanism during genetic combination of combatable partners. Another genetic protection system first reported in N. crassa was repeat-induced suppression in which haploid genomes are scanned for duplicate genes, and if found, are polarized by a released mutagen which acts to silence or methylate a genetic element. Studies on N. crassa additionally proved that fungi were eukaryotic which developed our understanding metabolic pathways in all eukaryotes (Davis 2000). Lastly, on the scientific glory list from this fungus is that it was involved with the discovery of mitochondria in 1953. Today, N. crassa is still studied extensively for discoveries in genetics, epigenetics, circadian rhythms, and molecular processes. It is also used in education (Dunlap 1999; Dunlap 2000; Dunlap 2008). This fungus has earned the title of a model organism due to its history of study and usefulness. Some of these useful attributes include a largely haploid life cycle, fast life cycle (10 days), genetic simplicity, wide distribution, and availability of strains. Some of my favorites include “frost”, “cum”, “it pokes along”, and “microcycle blastoconidiation”. The entire genome for N. crassa was sequenced in 2003.

hphae neuro Petri Neuro Neuro

 

Photos: Fungal Genetics Stock Center. http://www.fgsc.net/cultureimages/morpholpix.htm

Neuro life cycle

Source: Wikipedia

wild Neurospora

Jacobson David. Stanford University. http://www.fgsc.net/Neurospora/sectionB4.htm

N. crassa is a fungi in the class ascomycota. It reproduces both asexually via structures called conidia and sexually via structures called asci. N. crassa ascospores are found to only germinate upon a heat treatment of atleast 60C. This has to do with its life strategy in ecology. N. crassa grows primarily in circum boreal regions, although it has recently been found in temperate region. It occurs on wood and vegetation after forest fires where it appears as a red to orange dust or cotton appearance. Although in lab N. crassa has been studied exensively, much is still unknown about its functions in nature. Due of its relationship with heat it became a bane for bakeries and remains a common contaminant today. N. crassa remains one of the most important organisms of genetic study to date in addition to a slew of other findings and developments in biology. Not only did this fungus vastly increase our understanding of fungi it provided us with key insights into the functions of higher eukaryotes such as ourselves. It is amazing to me how open ended the possibilities for us to increase our biological understanding with fungi. We have the ability now to deeply investigate and study these organisms with the advent of sterile environments, gene sequencing, staining, computing etc. Further inquiry into this comparatively untouched field will become increasingly significant to progress in biology.

Sources:

Beadle, G. W., and E. L. Tatum, 1941 Genetic Control of Biochemical Reactions in Neurospora. Proc Natl Acad Sci U S A 27: 499-506.

Davis, R., 2000 Neurospora: Contributions of a Model Organism. Oxford University Press, New York.

Davis, R., 2000 Neurospora: Contributions of a Model Organism. Oxford University Press, New York.

Dunlap, J. C., 1999 Molecular bases for circadian clocks. Cell 96: 271-290.

Dunlap, J. C., 2008 Salad days in the rhythms trade. Genetics 178: 1-13.

Murray, N. E. 1960. Complementation and recombination between Methionine-2 alleles in Neurospora crassa. Heredity 15:207–217.

Websites:

Gavric Olivera. Images of Neurospora Morphological Mutants in Culture. Fungal Genetics Stock Center. http://www.fgsc.net/cultureimages/morpholpix.htm

Neurospora Crassa Life Cycle. Wikipedia. http://en.wikipedia.org/wiki/File:Neurospora_crassa_life_cycle.jpg

Jacobson David. Stanford University.  http://www.fgsc.net/Neurospora/sectionB4.htm

The wonders of Fomes fomentarius

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By Mary Perkins

The fungus Fomes fomentarius (“tinder fungus”), though easy to pass without notice, has an array of wondrous qualities that should be recognized. These range from wood degraders, fire starters, clothing, weaponry, and medicinal uses. It has a long history of being used all around the globe, even dating back to the 5,000 year old Iceman, Otzi.

Lifestyle of the Fungi

Fomes fomentarius is a dark gray hoof-shaped (Lincoff, 1991) saprotrophic (lives on dead organic matter) fungus that grows on dead wood or wounded trees, mainly hardwoods, “. . . particularly fond of birch trees, although other trees, including firs, can be the host,” (Stamets, 2002). The most important role this polypore (the fertile tissue is composed of many pores rather than gills) fungus plays in ecology is the break-down of wood. It has the capability of degrading lignin, as a white rot. Wood is a very hard substance that only certain fungi can break down, Fomes fomentarius being one of them. They help rid the forest floor of litter, acting as nature’s recycler. Imagine how high a pile trees can reach if they were not decomposed. Not only do they clean up the forest floor, they help soften wood to make it possible for insects and nesters, such as birds and squirrels, to inhabit the trees (Bunnell & Houde. 2010).

Uses of Fomes fomentarius

Humans have had an intimate relationship with Fomes fomentarius for many years. They have utilized this fungus for medicinal purposes, carrying embers, fire starters, weaponry, and even clothing. The Okanagan-Colville natives used the fungus to make antimicrobial teas and poultices to treat infections and arthritis (Stamets, 2002). It was used to cauterize wounds by Laplanders and the Cree to treat frostbite (Rogers, 2011). The entire fruiting body can be hollowed out to carry embers while travelling from one camp tfomes hato another. The inside of the fungus can be dried and is easy to light with only a spark. The fungus was also used to discharge guns, from the spark of the flint, to the fungus, to the gunpowder (Stamets. 2002). When the fruiting body is smashed it becomes felt-like, usable for clothing materials, such as this fashionable hat, (see photo).

Evidence of the use of the tinder fungus has been found dating as far back as the Iceman, Otzi. A 5,000 year old mummy was found, preserved in ice, with his clothing and tools. Among these objects was Fomes fomentarius along with flints, as part of his, “fire-making kit,” (Moore et. al, 2013). It is amazing to think that the same use of the same fungus has been in practice for so many years, covering a vast area and peoples. We are similar beyond borders.

The tinder fungus plays many roles. It is important to forest ecology as well as beneficial to humans. It is a decomposer and a homemaker. It can be weaponized or create warmth and healing. It may not be a pretty fungus but there is more than meets the eye.

References:

Arora, D. 1986. Mushrooms demystified. New York: Random House

Bunnell, F. L., & Houde, I. (2010). Down wood and biodiversity – implications to forest practices. Environmental Reviews, 18(1), 397-421. doi:10.1139/A10-019

Lincoff, G., Knopf, A. 1991.The Audubon society field guide to North American mushrooms. New York: Knopf

Moore, D., Robson, G.D., Trinci, A.P.J. 2013. 21st century guidebook to fungi. New York: Cambridge University Press

Rogers, R. 2011. Fungal pharmacy. Berkeley, CA: North Atlantic Books

Stamets, P. 2002. MycoMedicinals An informal treatise on mushrooms. Hong Kong: Colorcraft Ltd.

Vetrovsky, T., Voriskova, J., Snajdr, J., Gabriel, J., and Baldrian, P. (2011) Ecology of coarse wood decomposition of saprotrophic fungus Fomes fomentarius. Biodegradation, 22(4), 709-718 doi: 10.1007/s10532-010-9390-8

www.mesiah.edu

Albatrellus ovinus

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By Marcus Goodman

Few things are pursued with such a broad spectrum of enthusiasm as gaining an understanding of the Kingdom Fungi. Loathed by as many as are enthralled by them, fungi in their fruiting form have an effect on some which can border on the maniacal. Some are intrigued by kingdom Fungi’s complexities which have enabled similar morphological traits to converge as well as separate across phyla. Others are more entranced by the mystical, spiritual and medical benefits. A quality alleged by some of the kingdom Fungi. Some just love the pretty colors; while others have more gastronomical hopes in mind. The latter is where my interest seems to be most prevalent, and it was in this pursuit I first encountered Albatrellus ovinus (Shaeff).

Untitled

Photo by Author H. Crisp

I was enjoying an extended season as a budding mycophagist in the Quinault Lake region of Washington State over Thanksgiving weekend. This region hadn’t yet experienced the fungi-fruit killing frosts experienced by most of western Washington earlier in November. Consequently, many fungi-fruiting bodies were in prime form for the picking—Hydnum repandum, Hydnum umbilicatum, Tricholoma magnivelare, Sparassis crispa, Hericium abietis, Craterellus tubaeformis, were just a few edible species present or abundant. I was on my way home from visiting a Western Red Cedar bog, a pretty rare biological community found in the Quinault area of the temperate rainforest, when I first noticed large white-buff-yellow fruiting bodies in the vicinity of old growth Tsuga heterophylla. At first, glance I almost walked right on by, potentially mistaking them for similar-looking Tricholomas I’ve had difficulty identifying this year. One thing I’ve learned with mushrooms: don’t assume anything, and if something catches your eye, don’t hesitate to investigate—you never know what you’ll find.

Here’s what I observed:

  • Habitat:  Old growth Tsuga heterophylla dominated forest. 85% canopy Gaultheria shallon, Polystichum munitum, Vaccinium ovatum dominant shrub/scrub Polytrichum spp., Kindbergia oregano, Holocomium splendens dominant groundcover
  • Growth Habit:   Scattered to gregarious
  • Substrate: Humicolus. Minimal soils present. Specimens found in isolated soil/hummus. Potentially decomposed wood present
  • General Characteristics:  Cap:8-16 cm.white-to-buff-to-light yellow, convex to plane with decurved to uplifeted margin. Cap surface dry, smooth, slightly scaled towards center. Margin even and regular. Cap flesh white, firm, dry-to-slightly moist, no apparent bruising, taste or smell.
  • Gills: Decurrent, pores, white-to-buff, smooth, minute
  • Stem: 0.5-2.5 cm thick, 3-8 cm long. White-to-buff. Central, equal to slightly clavate surface smooth at base to porous at decurrent pores. No mycelium observed at base.

You can imagine, as someone intent on maximizing the culinary benefits of my mycelial friends, I was more than intrigued at this finding. If it wasn’t for a full foraging bag of other known delectables, I would have gathered some for the table. As it was, I gathered none—planning on coming back in the morning for further investigation and observation. This decision was one I would come to regret.

All best laid plans are usually just that. Plans. The reality was an obscured series of days, blurred by a food coma that rendered all hopes of activities more than 20 minutes beyond a suitable napping location simply futile. How many mushroom foraging hopes have been shattered by the ubiquitous turkey on the days following Thanksgiving?

During rare moments of lucidity, I was able, through semi-conscious keying sessions, to use the standard mushroom literature for the Pacific Northwest: Mushrooms Demystified (Arora, 1986) and Mushrooms of the Pacific Northwest (Trudell, Ammirati, 2009) to narrow the possibilities to genus Albatrellus, but without an actual specimen, the specific epithet proved elusive. Arora—the obligate mycophagist—declares (ovinus and associates) them edible when cooked well, but includes the disclaimer, “Large quantities can have laxative effects.” Ever creative with his descriptive terms, Arora uses—“okraceous”—for the flavor/texture/consistency. As usual, Trudell and Ammirati are silent on the matter. Although their general conclusion is our coastal specimens are A. avellaneus (Though not supported by Arora’s description) with A. ovinus apparently not occurring in our region. This inability to solidify the identity was further enabled as a result of the disproportionate napping activity.

Albetrellus

Raija Tuomainen

However, after the weekend was over and I was back at home, I couldn’t stop thinking about the pale, firm, slightly moist, fleshy consistency of the one that got away. Absence definitely makes the heart grow fonder. What to do? Internet search time. Not a big fan of this, but there are a few locations with worthwhile and reliable information:

http://www.mushroomexpert.com/albatrellus_ovinus.html

http://www.mycobank.org/Biolomics.aspx?Table=Mycobank&MycoBankNr_=292351

Like all things in the fungal kingdom, nothing is ever easy. However, if this was “Mission Difficult,” anyone could do it and there’d be less mushrooms in the woods, right? It turns out, A. ovinus is a highly regarded mushroom in the Scandinavian region—specifically Finland. This excited me, since I come from Finnish decent. Unfortunately, I can’t roll my rr’s. Because of this my grandma refused to teach me any Finnish beyond numbers 1-10, and some miscellaneous body parts I won’t mention here. All I could do was look at the pictures and dream. I finally gathered questionable, boiler plate mushroom guidance from an array of sources not worth citing:

  • Don’t consume A. ovinus with other mushrooms
  • Cook A. ovinus well
  • May cause a laxative effect
  • Don’t eat raw
  • May contain phenolic compounds (Dekermendjian, et al, 1997)
  • Becomes bitter with age
  • Slimy when cooked
  • Blah, blah, blah

I know I should always treat mushrooms with respect, especially one that may contain phenolic compounds, which is a chemical compound found in plants and some fungi, but the above comments seem to contradict the apparent delectability of the genus in Scandinavia and Northern Europe. As I sit here and gaze dreamily at all of the Googled images of this mushroom, many about to, or being eaten, I can’t help but want to know more. By next fall I’ll either have to learn Finnish, or give Albatrellus ovinus a taste test. So it goes with the kingdom fungi.

References:

Arora, D. 1986. Mushrooms Demystified, Second Edition. Random House, Inc. New York.

Dekermendjian, K., Shan, R., Nielsen, M., Stadler, M., Sterner, O., Witt, M.R. 1997. The affinity to the brain dopamine D1 receptor in vitro of triprenyl phenols isolated from the fruit bodies of Albatrellus ovinus. European Journal of Medicinal Chemistry. Vol. 32, Issue 4, pgs. 351-356.

http://www.mushroomexpert.com/albatrellus_ovinus.html

http://www.mycobank.org/Biolomics.aspx?Table=Mycobank&MycoBankNr_=292351

Trudell, S., Ammirati, J.2009. Mushrooms of the Pacific Northwest. Timber Press, Inc. Portland OR.

Vrkoc, J., Budesinsky, M., Dolejs, L.1977. Phenolic meroterpenoids from the basidomycete Albatrellus ovinus. Phytochemistry.Vol 16, Issue, 9, pgs 14091411.

HONEY BEE MURDER! Suspect number one? Nosema ceranae!

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By Kaitlyn Texley

We’ve all heard it said, without bees humanity would cease to exist four years after. Thanks to a fungus we may be approaching this apocalyptic prediction. Colony collapse is nothing new. We’ve read about it, seen it on TV and heard conspiracy theories about the multitudes of causes. Recent research into what is actually causing the rapid collapse revealed that it is actually the parasitic microsporidia Nosema ceranae.  Even weirder, bees exposed to fungicides and pesticides the rates of fungal infection, or nosemosis, are even higher (Pettis, et al 2013).

The spores of the N. Ceranae enter through the bee’s mouth and travel into the mid-gut. In bees, the mid-gut is where the digestive enzymes used to break down pollen reside. Here the microsporidia’s spores grow, reproduce and create more spores. The microsporidia feeds off of the cells within the bee’s mid gut causing damage and weakening the bee. As it does this, the microsporidia also inhibits the genes responsible for the regeneration of the intestinal tissue (Dussaubat et al, 2012). So not only is the mid gut being digested and consumed by the fungi, the bee’s body can’t heal any of the destruction.

The bee’s body responds by generating antioxidant enzymes and sending them to the site of the Nosema ceranae infection (Dussaubat et al, 2012).  This seems to do little for the poor bee. The microsporidia continues to ravage the intestinal tract eventually travelling to the rectum and out with the bee’s poop. Healthy worker bees then clean up the poop and become infected by the microsporidia themselves.

The effects are really terrible. The little worker bee can have 30-50 million spores within their tiny little body (Nosema ceranae and nosema disease of honeybees, 2010). Of course this has an impact on their well-being. They become tired and are unable to produce “brood food” because of the total takeover by the fungus. This food is meant to be eaten by the baby bees, however in a severe case of Nosema ceranae infection, there may not be many babies. If the queen becomes infected her ovaries stop working and she is unable to produce eggs, and thus, babies (Nosema ceranae and nosema disease of honeybees, 2010).

How does a hive get exposed to the microsporidia? Sometimes honey-bees will rob other hives if they can’t get enough food. If they rob a hive that has honey containing Nosema ceranae spores they’re screwed (Nosema ceranae and nosema disease of honeybees, 2010). Apiaries can also increase the likelihood of infection by combining two dwindling colonies, both of which are, unbeknownst to the apiary, infested with Nosema ceranae (Nosema ceranae and nosema disease of honeybees, 2010).

Bee killer
Figure 1.  Nosema ceranae life cycle.

The severity of the colony’s infection coincides with the seasons.  During the colder spring months when the weather is sporadic, the bees are stuck inside of their hives. This is also when baby making begins, the worker bees are continually cleaning out contaminated combs for new baby bees and being exposed to the microsporidia. This is when the hive become highly infested and usually, if the infection is bad enough, when the colony itself collapses. If the colony does not collapse and they make it to summer, they are offered a new chance. The drones and worker bees (the adults affected by the fungus) leave the hive and die. The baby bees emerge microsporidia-free and the combs of the hive are cleaner as brooding has decreased. It’s a new lease on the colony’s life.

If the coming fall is wet, good conditions for microsporidia, the bees have a higher chance of contracting Nosema ceranae again. If the fall is dry, the colony may be safe from the infection for the following year.

It’s important for farmers, apiaries and scientists to learn as much as possible about Nosema ceranae so they can prevent the contraction of it. Although not all fungus are bad, Nosema ceranae is dependent on the bee’s mid-gut to reproduce and thrive. Unfortunately this is not only bad news for the little honey bee that’s infected, it’s bad news for the entire colony, and in fact the entire human population. If colonies keep collapsing, we may not only have a shortage of honey, we may have a shortage of nearly 1/3 of the planet’s food supply (Yang, 2006). That is not a small amount. In fact, that is a very large amount. You would have to say goodbye to blueberries, almonds and many other fruits, vegetables and nuts.

So although you may not care very much about Nosema ceranae and what it’s doing to a bee’s mid-gut, you will care when the repercussions of this widespread infection effect the food choices at your local grocery store. With a reduction of pesticides and fungicides the susceptibility for an infection to happen and spread will decrease (Pettis, et al, 2013). Along with knowledge of how the microsporidia spreads farmers, apiaries and scientists may be able to reduce the spread of Nosema ceranae and keep our food where it belongs, on our plates!

References:

Moisset, Beatriz, Buchmann, Stephen. Bee Basics; An Introduction to Our Native Bees. (2011). Retrieved January 28, 2014, from http://www.fs.usda.gov/Internet/FSE_DOCUMENTS/stelprdb5306468.pdf

Dussaubat, Claudia et al. Gut Pathology and Responses to the Microsporidium Nosema ceranae in the Honey Bee Apis mellifera. (2012). PLOSone. DOI: 10.1371/journal.pone0037017

Pettis, J., Lichtenberg,E., Andree, M., Stitzinger, J., Rose, R., vanEnglesdorp, D. (2013). Crop Pollination Exposes Honey Pees to Pesticides Which Alters Their Susceptibility to the Gut Pathogen Nosema ceranae. PLOSone. DOI: 10.1371/journal.pone.0070182

Nosema ceranae and nosema disease of honeybees. (2010). Retrieved January 26, 2014, from http://www.biosecurity.govt.nz/pests/nosema-ceranae.

Yang, Sarah. Pollinators Help One-Third of World’s Crop Production, Says New Study. (2006) UC Berkeley News. Retrieved January 28, 2014, from http://berkeley.edu/news/media/releases/2006/10/25_pollinator.shtml

Photo: http://scientificbeekeeping.com/the-nosema-twins-part-1/

Aspergillus niger: not your everyday mold

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By Jasmine Zamora

When many people think of fungi, they first envision beautiful fruiting bodies that grow out of the damp grass and soil.  However, there is a much darker side of the kingdom fungi that can cause sickness and death.  The members of the Aspergillus genus do not grow on recently rained-on terrain, nor do they need a specific temperature to grow in.  These are molds, most familiarly Aspergillus niger, can in some cases cause disease in humans, animals and plants.Aspergillus

Aspergillus niger isn’t as notoriously dangerous as Aspergillus fumigatus, which is the most prevalent airborne fungal pathogen (Latgé 1999).  However, Aspergillus niger contains toxins that can make people with weak immune systems become very sick and can sometimes result in death.  These toxins can be inhaled by humans, most commonly people who work around plants or peat, and can cause a lung disease called Aspergillosis, which has infected over 300,000 people worldwide (Keir 2013).  Aspergillus niger is not one of those that are so deadly, but can definitely cause sickness and allergic reactions.  Aspergillus niger is an asexual saprophytic fungus that can grow on dead leaves, stored grain, compost piles and other decaying vegetation.  A. niger is a very thermotolerant fungus that can thrive in freezing conditions and very hot weather (Metzger 2008).  It produces its spores on an asexual structure called the conidium.  The spores can be inhaled when simply working with anything A. niger has colonized.

Aspergillus niger contains several toxins, some harmless and others harmful to certain people.  The main toxins it contains are malformin C, and ochratoxin A.  A. niger can be as beneficial as it is harmful, though.  Through fermentation, it can produce useful enzymes that can be used in the production of corn syrup, Beano, wine and cider (Rajkumar 2010).

Fortunately, most people can handle the inhalation of a moderate amount of A. niger spores.  Aspergillus spores are in the air we breathe almost everywhere we go.  It is those who suffer from leukemia, HIV or AIDS, severe fungal allergies and other immune deficiencies that could become very sick to the intake of A. niger spores (Bartholomew 2009).  There has actually been a case where a 70 year old man had to have his foot amputated because there was a “painful black ‘gangrenous appearing’ mass on his foot.  “Tissue samples showed not only branching hyphae, but dark pigmented fungal fruiting heads with double sterigmata in which Aspergillus niger was identified” (EPA.gov).  This may be a case of a sickly man that was just unfortunate enough to come into contact with A. niger, but the so-called “harmless” fungi can effect healthy people in rare cases.  Otomycosis, an ear infection that can be very painful, can be caused by Aspergillus niger.  Allergic reactions can be severe when an individual that is very allergic to fungi.  “When inhaled, A. niger can cause hypersensitivity reactions such as asthma and allergic alveolitis” (EPA.gov).  This is uncommon, but nearly fatal if the wrong person became infected.

A. niger can also affect plants, such as onions and tomatoes. Small animals, such as mice and chicks were fed moldy soybeans with A. niger on them, and the subjects subsequently died after digestion (EPA.gov). Onions are a common plant that A. niger likes to inhabit, causing spoilage and can then result in economic loss in farming communities.  Mangoes, grapes and tomatoes are also victims of the pathogen, as well as 34 other genera of plants.  Lastly, A. niger can cause the rotting of wood and other hard surfaces. Wood surfaces have been found softening or deteriorating because of the contamination of A. niger. There have also been reports of A. niger effecting very random substrates such as polyvinyl acetate, polyester-type polyurethanes, and even English style crumpets.

Aspergillus niger does not discriminate when it comes to what it wmoldants to contaminate.  However, this very durable fungus is nothing to be worried about, unless you have a weak immune system or have a sensitive allergy to fungi, or if you are a mouse.  There are also remedies that can be taken if one contracts Otomycosis or another common fungal infection.  There are many other more harmful fungi that are in the same family (A. fumigatus) to be worried about, so be sure you don’t inhale too many A. niger spores, and don’t eat heavily molded fruits that are covered in a black substance, and you should be fine.

 Works Cited

Anonymous. 1997. Aspergillus niger Final Risk Assessment. Biotechnology program under toxic substances control act (TSCA) U.S. Environmental Protection Agency U.S.A. 3171.

Corrigan. (October 15, 2009). Aspergillus niger. In Encyclopedia of Life. Retrieved January 28, 2014, from http://eol.org/pages/2920814/details.

Keir, G. J., Garfield, B., Hansell, D. M., Loebinger, M. R., Wilson, R., Renzoni, E. A., & Maher, T. M. (2013). Cyclical caspofungin for chronic pulmonary aspergillosis in sarcoidosis. Thorax, thoraxjnl-2013.

Latgé, J. P. (1999). Aspergillus fumigatus and aspergillosis. Clinical microbiology reviews, 12(2), 310-350.

Metzger, B. (November 16, 2008). Aspergillus niger . In Mushroom Observer. Retrieved January 28, 2014, from http://mushroomobserver.org/name/show_name/15030?_js=on&_new=true.

Paul, R., Singh, V., Tyagi, R., Singh, A., & Dubey, D. (2010). Micro-Elements work for the growth and total soluble protein production in Aspergillus niger at different concentrations. Journal of Pure and Applied Microbiology, 4, 293-296.

Sutton, D. A. 2005. Aspergillus niger.  Retrieved February 10, 2014. from:http://www.doctorfungus.org.

SCIMAT.  Aspergillus niger fungus. Retrieved February 10, 2014. from: http://viemxoang.vn/wp-content/uploads/2011/10/aspergillus_niger_fungus_aspergillus_niger_fungus_ah963a.jpg.

Prototaxites: The largest organism of its time!

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By Harmony Counsellor

Painting a picture:

Prototaxites     It’s the Devonian era; 450 m.y.a. Picture yourself looking over the vast fields of sprawling land – nothing to obscure your view. As you can see it, the land is like a prairie but wet, very wet. You see some mountains, some small land plants, some bugs roaming around the grass, and a giant mass. Your eyes stop moving – you stop breathing. What could it be? All you see is this giant veil that could swallow you up. You don’t know how to respond, but you can’t keep yourself from moving closer. It doesn’t seem to move. As you get closer, it gets clearer. This giant mass has turned out to be the shape of a Rocket Pop (mixed berry flavor). The size, as we can compare it to these days, is about as tall as a tree – 26 feet tall and about 3 feet wide. (Prototaxites, 2014)

What would you have thought if you were a there? The largest mass you have ever seen on the whole planet. While it looks so tame on the outside, do we dare explore? As science stands now, we wish we could explore. The remnants of this mass we call Prototaxites is but a fossil we cannot figure out. Throughout the last 150 years, (Hobbie, 2010) thoughts have wandered between it being a tree, fungal, algae, lichen, or even a liverwort. We can say so much about it, but the identity still remains a mystery.

History of Prototaxites:

CanadianArcheologist scientist J.W. Dawson first discovered Prototaxites in 1859 and described it as a tree under: Taxus. (“Prototaxites, a huge…”) He discovered the fossil within a conifer claiming the fungi was decomposing the tree. 14 years later, it was described it for the first time as Prototaxites roughly translating to ‘first yew’. (Prototaxites, 2014)

waterfallThe next accepted theory was one of rolled up liverworts. Graham (2010) suggested that as the soil degraded the liverworts rolled towards the bodies of water creating these ringed structures. By 2012, many authors have disproved this theory. (Hobbie, 2010; Kennedy, 2012; Boyce, 2010; Edwards, 2012)

WIERD FACTS: 

Though about 13 specimen have been found and identified (Taylor, 2010), Holland seems to have the highest population coming straight out of the river (“Prototaxites, a huge…”). The closest environment to the Devonian ages where protaxites was found is Lyman Glacier in Washington State!

But, what is it?

fossilBeing the largest organism in the period of existence (Prototaxites, 2014; Boyce, 2007; Hobbie, 2010), science is experimenting with everything they have. Since the time of its discovery we knew it was a giant land organism, but the time period is too old for any woody plants. Not woody, so what IS IT? The evidence of carbon isotope analysis supported by Boyce in 2007 suggests it is also not a vascular plant. If vascular plants are not an option, the only thing we have to go off of is Hobbie’s conclusion; ‘…land organism with the most recent anatomical work supporting a fungal interpretation’. (Hobbie, 2010) Anatomically speaking we have a giant mass that was neither a tree nor plant – but is it a fungus?

spore fossilWhat it was made of can also tell us what it ate. If you look at the other evidence, like biogeochemistry, it suggests we have an organism that consumes primary producers (cryptobiotic crusts, mosses, etc.).  Fact: Saprotrophic fungi are about 3% higher in Carbon 13 than their substrate. Simply put, you are what you eat. Carbon 13 is what plants use for photosynthesis and analyzing the amount of Carbon 13 will tell you what something is made of. As we look at our Prototaxites, it is depleted in its Carbon 13 and would have to consume autotrophic substrate (self feeding organisms). (Hobbie, 2010) Concluding that if it were a fungus it ate everything that photosynthesizes – other fungi and anything else in its way. If I were to make a judgment, I would say this is one mushroom not to be messed with.

Recent Evidence suggests…  

proto structurePhys.org suggests Graham in February 2010 may have proved the liverwort theory to be the most substantial evidence of its origins. But Hobbie published his paper in March 2010 stating the pattern of circles would not be prominent ‘circles’ but more roll like. With those two conflicting sets of evidence, we are almost left with an unknown organism – so what is the last theory? The last supporting evidence points to fungi. It seems to be made of rhizmorphs or hyphal networks (fungi’s version of a plant root). (Kennedy, 2012; Edwards, 2012) What about evidence pointing to algae, lichen and liverworts? Well Edwards (2012) is suggesting that the algae and lichen are part of the fungi’s tissue.  Lastly, what we do know is this: The largest organism in the Devonian time period was an eating monster! Isotope research concludes that prototaxites ate – ate anything that photosynthesizes. That doesn’t necessarily mean it was in the form of digestion like fungi, but could be in any form of eating. I picture a giant fungus with teeth, what about you?

Bibliography:

Anning, M. (2013, December 13). Awesome dead shit: Prototaxites. Retrieved from http://www.maryanningsrevenge.com/2013/12/awesome-dead-shit-prototaxites.html

Giant fossil prototaxites: Unraveling a 400-million-year-old mystery. (2010, February 10). Retrieved from http://phys.org/news185022458.html

Prototaxites . (2014, February 09). Retrieved from http://en.wikipedia.org/wiki/Prototaxites

Boyce, C. K. (2007, May). Devonian landscape heterogeneity recorded by a giant fungus. Retrieved from http://ctserver.uchicago.edu/pdfs/boyce/2007GeologyPrototaxites.pdf

Edwards, D. (2012). Selective feeding in an early devonian terrestrial ecosystem. Palios, 27, 509-522. doi: 10.2110/palo.2011.p11-094r

Graham, L. E. (2009, December 03). Structural, physiological, and stable carbon isotopic evidence that the enigmatic paleozoic fossil prototaxites formed from rolled liverwort mats. Retrieved from http://www.amjbot.org/content/97/2/268.full

Hobbie, E. A. (2010). Carbon sources for the palaeozoic giant fungus prototaxites inferredfrom modern analogues. . Royal Society Publishing, 277(No. 1691), pp. 2149-2156. Retrieved from http://www.jstor.org/stable/25706434.

Kennedy, K. L. (2012). Paleoenvironmental inferences from the classic lower devonian plant-bearing locality of the campbellton formation, new brunswick, canada.Society for Sedimentary geology, 27(6), 424-438. Retrieved from http://www.bioone.org/doi/full/10.2110/palo.2012.p12-004r

Prototaxites, a huge, 400 million years old, fungus? or an enormous lichen?. (n.d.). Retrieved from http://steurh.home.xs4all.nl/engprot/eprototx.html

Taylor, C. (2010, February 10). Prototaxites: A giant that never was?. Retrieved from http://coo.fieldofscience.com/2010/02/prototaxites-giant-that-never-was.html