Fungi in art

Early depiction of mushrooms in the Codex Magliabechiano from the mid-16 century (facsimile from Zelian Nutall, 1903). The image is thought to depict the long tradition of consuming entheogenic (psycoactive) mushrooms in Mesoamerica. The man is approached by a God of the Atzec underworld.

Fungi have had an enormous influence in the arts since centuries, being used as inspiration or subject matter, with artists using fungal materials as working matter since more recent times. Presence of fungi in the arts abound from across the globe (with well documented examples from Europe, the Americas, Southeast Asia, Siberia) and among different cultures since at least 1500 BCE, as for example in 'Mushroom Stones' in Mesoamerica (current Guatemala, Honduras, El Salvador, Mexico, and other countries). The use or inspiration of fungi in the arts is extremely various and prolific due to the highly different morphologies, forms, effects, properties, ecologies, biogeography (distribution), and lifestyles of fungi. Fungi can be multicellular or single-celled, and are an incredibly diverse kingdom of life belonging to the eukaryotes (nucleated or 'higher' organisms). Traditionally, mushrooms have been the predominant form of fungi in the arts, with other fungal forms (mycelia, hyphae, spores, moulds, yeast, or lichens) being neglected or ignored. Virtually all areas of the arts have been infiltrated by fungi. These areas include literature and fiction, paintings and the graphic arts, and since more recent times contemporary art, music, photography, comic books, sculpture, video games, dance, cuisine, architecture, fashion, and design. Contemporary artists experimenting with fungi often work within the realm of the BioArts.

Artists working with fungi are mostly representing (describing), showcasing, transforming, or utilizing them. Various ways and approaches of working with fungi are a spectrum which goes from the descriptive (representation, showcase) to the more active use of fungi as working matter (transformation, utilization). The distinction between these art practices and approaches are not clear-cut: Artists representing mushrooms might do so allegorically or using them as narrative, rhetorical, stylistic, or stage element; artists showcasing fungi might transform them (e.g. for preservation); artists using fungi as transformative agent might showcase and utilize parts of the fungal life cycles for their work. Contemporary artworks using fungi often explore the topic of transformation, decay, renewal, sustainability, circularity of matter. Artists working with living fungi (rather than their representation or deactivated parts thereof) occasionally use them as subject for time-laspe photography. Occasionally, artworks use fungi conceptually (that is, to communicate the capabilities and potential of fungi), metaphorically (that is, to convey an idea, process, or concept), or experimentally (that is, to develop new 'fungal art' techniques).

Artworks representing, showcasing, transforming, and utilizing fungi. Clockwise from upper left: (1) Mushroom sculptures likely inspired by the use of Psilocybe mushrooms, and thought to be involved in religious or spiritual rituals; depicted are pre-Colombian statues (ca. 1000 BCE-500 AD). (2) A glass sculpture by mycologist William Dillon Weston representing the phytopathogen Botrytis cinerea (ca. 1940). (3) The sculpture 'Champi(gn)ons' (2017) by artist Vmeer made with parasol mushrooms (Macrolepiota procera, metal, shellac, 30 x 20 x 8 cm); showcasing mushrooms requires their transformation and preservation. (4) The 'MY-CO SPACE' building prototype is a inhabitable structure utilizing the mycelium of the fungus Fomes fomentarius.

Beside working with and manipulating fungi, artists are passively (indirectly) influenced by them. For 'indirect influence of fungi' it is meant the depiction or description of the effect of fungi; the creation of art upon influence of fungi-derived substances, like hallucinogenic psylocibin derivatives or products of fungal fermentation (like alcohol); and, in several cases, both, like in Charles Bukowski's literature or in Hieronymus Bosch's visionary paintings. Cases where fungi have played an unbeknown role into the process of artistic creation (for example, because the fungal properties or effects had not yet been described, understood, or manipulated), could also be considered an indirect influence of fungi in the art.

Traditionally, mushrooms have been the main subject for depiction in the arts; more recently, artists experiment with further forms of fungi, like mycelia, hyphae, spores, moulds, yeast, or lichens. As of 2022, there are still relatively few examples of direct representation, showcase, transformation, or utilization of these latter forms in the arts; those examples mostly, but not always, fall within the contemporary arts. Often, artists focus on one major form of fungi (mushrooms, mycelia, hypha, spores, yeasts, molds, lichens) in the fungal lifestyle. There are few examples in which an artwork depict several forms across the fungus reproductive cycle. Similarly, an artform rarely encompasses multiple fungal forms due to the extremely varied morphologies, properties, phylogeny and lifestyles of fungi (except for the disciplines of non-fiction literature, documentary, participative art). Conversely, the enormous plasticity of fungi enables artists to work with different fungal forms to create very diverse artworks.

In terms of which arts and genres have been infiltrated by which form of fungi, artists mostly works with mushrooms in the graphic arts, literature, cinema, architecture, and more recently contemporary art; forms like mycelia and hyphae are increasingly used in the arts, and in particular in contemporary art, fashion, design, and architecture due to their plasticity, elusiveness, and their numerous functions, effects, and uses; spores or moulds are mostly used in artistic representations or fictional settings of infection or decay, often in literature or the visual arts. In the Western art, representation of fungi (mushrooms) have been historically saturated with negative associations, whereas many examples of mushroom depictions or descriptions in various Asian arts and Folk arts are more favourable towards fungi. Reflecting these representations of mushroom, Western cultures have been referred to as mycophobes (fear, loathing, or hostility towards mushroom), a term first coined as 'fungophobia' by British mycologist William Delisle Hay in his 1887 book An Elementary Text-Book of British Fungi, whereas Asian cultures have been generally described as mycophiles.

Painting of Slavic folktale character Baba Yaga with mushrooms on the forest floor. The fly agaric (Amanita muscaria) has entered folklore in numerous occasions due to its particular appearance and properties.

Increasingly, these paradigms are being shifted, with more mushrooms, spores, moulds, and yeasts being depicted positively, or without negative connotation, in the arts (as for example, in spore prints, in the musical theatre show The Mould That Changed the World, in Beatrix Potter's mushroom illustrations and children drawings, or in the contemporary art exhibition Fermenting Futures exploring the metabolic capabilities of yeasts, just to name a few). Given the surge of artistic experimentation with fungi in recent years, what some referred to as a 'fungal renaissance,' more and more exhibitions, festivals, events, group activities, and citizen science projects are entirely devoted to fungi. These events often aim at presenting, discussing, growing, and experimenting with multiple forms of fungi as well as with different artforms, and are often visited or contributed to by a growing community of (amateur) mycologists, mycophyles, and experts. Since 2021, the annual Fungi Film Festival invites, selects, curates and recognizes movies about fungi and lichens in all categories or genres, including: experimental movies, shorts, fantasy and science fiction movies, comedies, music videos, animated movies, stop motion, documentaries.

Fungi permeate the arts extensively and in different artforms and occasionally in very variegated artworks due to their intrinsic or external (attributed) qualities, properties, and applications, such as: the mysteriousness of mushroom appearance, the grotesque shape of many mushrooms, the invisibility of many fungal forms, the plasticity of mycelial and hyphal growth, the hallucinogenic effects of certain mushrooms, the toxicity of some species, the infectiouos potential of pathogenic fungi, including phytopathogenic or enthomopathogenic species, the sustainability of biomaterials from fungi for circular economy, the popular interest in fungi since recent years, and further attributes or properties of fungi. Some stories or artworks use fungi allegorically, or fungi are explored, interpreted or explained as component or inspiration of mythology and folklore. The visible parts of some fungi (mostly: mushrooms), and in particular those mushrooms with a distinctive, flashy, or flaring appearance like the fly agaric (Amanita muscaria), have significantly contributed to folklore.

A group of toads sit on top of a ring of mushrooms having tea in the drawing The Toads' Tea Party (ca. 1905) by writer, illustrator, natural scientist, and conservationist Beatrix Potter (1866-1943).

Often, artistic experimentations with fungi involve scientists, researchers, ecologists, designers, architects, and others tinkerers in multi-disciplinary approaches. Direct applications of fungi in architecture, design, and fashion often starts with artistic experimentations with fungi. Artists and scientists jointly defined a framework for fruitful collaborations between (fungal) science and the arts. Further important aspects of Fungi in Art relate to preservation of artworks from fungal decay and contamination, as well as initiatives fostering and supporting works able to stimulate dialogues between mycologists (fungal researchers), artists, and society (as for example from the Massee Art Grant by the British Mycological Society or works encouraged and supported by the Fungi Foundation). There are very few examples of musea entirely devoted to fungi (one example being the Museo del Hongo in Chile, South America).

The generally low visibility of fungi (other than mushrooms) in the arts can be correlated with the general knowledge and research on fungi, both of which lag behind in comparison with other life science disciplines: Mycology was named as a natural science discipline of its own in 1836 only, the fungi kingdom Funga was defined in 1969 only, and even today conservation efforts on fungal biodiversity lag behind in comparison to those of species in other kingdoms of life like animals and plants. Currently, in the International Union for Conservation of Nature's Red List of Threatened Species, only over 500 fungi are included, in comparison to over 58,000 plants and 12,000 insects.

Mushrooms in art

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See also: Mushrooms in art and Ethnomycology

Mushrooms are the visible part of fungi. Mushrooms are also called fruiting bodies, as they are the reproductive parts of fungi. Those fungi which do not form fruiting bodies (so called "asexual" species) are called Fungi Imperfecti (in Latin, Deuteromycota, literally meaning 'Fungi without womb'), a now obsolete term. Early natualists and mycologists described under Fungi Imperfecti all those fungi which could not be classified within established groups based on the appearance and morphology of mushrooms. Among Fungi Imperfecti are those Ascomycete or Basidiomycete fungi whose mushrooms have not been yet described. Yeasts and mould are (historically) considered Fungi Imperfecti. The mycobiont of lichens (the fungal partner of the algae symbiosis), also do not form fruiting bodies.

Mushrooms are the spore-bearing structures of fungi. Spores, the reproductive and dispersal unit of fungi, can germinate into hyphae, entangling into mycelium, which can produce fruiting bodies at the right conditions (e.g. in moist, wood soil in Norther Hemisphere during Autumn). Those are the mushrooms. The other forms and lifestyles of fungi are largely invisible, a reason why mushrooms are often represented or showcased, but rarely transformed and utilized, in the classical arts. In many languages, fungus and mushroom translate with the same word, like in the Italian 'fungo' (pl. 'funghi'), the German 'Pilz' (pl. 'Pilze'), the French 'champignon' (pl. 'champignons'), or the Bahasa Indonesia 'jamur' (pl. 'jamur jamur'). Mushrooms (let alone fungi) are itself hugely diverse. Fungi are considered a kingdom in itself (Funga), at the same phylognetic level as animals, plants, bacteria, archaea, and protists. Despite they can form multicellular structures (like mushrooms, mycelia, and hyphae), fungi are considered microorganisms. Occasionally, the term microfungi is used to refer to those fungi which do not form visible structures or form microscopic sporangia (spore-bearing fungal structures).

Scenes of mushroom hunting are a recurrent theme in the graphic arts. Depicted here is Mushroom picking (ca. 1860) by realist Polish artist Franciszek Kostrzewski (1826-1911).

Artists have often represented or described mushrooms as decorative, naturalistic, or symbolic element. In the graphic arts, architecture, sculpture, and literature, artists mostly represented or showcased mushrooms. Currently, contemporary artists are increasingly including mushrooms in their arts. Early depictions of fungi are petroglyphs from the Bronze Age (ca. 3300-1200 AD) near the Pegtymel River in Chukotka, Siberia ('Pegtymel petroglyphs'), representing people with mushroom-shaped head.

Given the mysterious, seasonal, sudden, and at times inexplicable appearance of mushrooms, as well as the hallucinogenic or toxic effects of some species, their depiction in ethnic, classic and modern art (around 1860–1970) is often associated in Western art with the macabre, ambiguous, dangerous, mystic, obscene, disgusting, alien, or curious in paintings, illustrations, and works of fiction and literature. British author Sir Arthur Conan Doyle wrote in his novel Sir Nigel:

"The fields were spotted with monstrous fungi of a size and color never matched before—scarlet and mauve and liver and black. It was as though the sick earth had burst into foul pustules; mildew and lichen mottled the walls, and with that filthy crop Death sprang also from the water-soaked earth."

In Asian or Folk Art, fungi are represented in a generally more positive or mystic way than in Western art.

Graphic arts

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Visual artists representing mushrooms have been very prolific throughout history. Whereas examples before the 15th century are rare, examples abound from European visual arts from the 1500 onwards including periods as the Renaissance, the Baroque, Flemish, and Romantic periods. The Registry of Mushrooms in Works of Art from the North American Mycological Association curates an extensive virtual collection of mushrooms in the visual arts.

While in the graphic arts mushrooms are mostly represented or showcased, they can occasionally be utilized to produce art. The 'Shaggy ink cap' and the Coprinus comatus and the 'Common ink cap' Coprinus atramentaria mushrooms produce spores by deliquescing (liquefying, or melting) their cap into a black ink, which can be used in drawing, illustration, and calligraphy. Protocols to produce the 'mushrooms ink' can be found online.

Prehistoric art

Mushrooms, and in particular fungi with psychoactive properties, have influenced cultures across the globe since millennia as entheogens. Early human civilizations carved mushroom figures, as in petroglyphs representing mushroom-headed people discovered near the Pegtymel River (Siberia). Mushrooms have been found in art traditions around the world, including in western and non-western works. Ranging throughout those cultures, works of art that depict mushrooms can be found in ancient and contemporary times. Often, symbolic associations can also be given to the mushrooms depicted in the works of art. For instance, in Mayan culture, mushroom stones have been found that depict faces in a dreamlike or trance-like expression, which could signify the importance of mushrooms giving hallucinations or trances. Another example of mushrooms in Mayan culture deals with their codices, some of which might have depicted hallucinogenic mushrooms. Other examples of mushroom usage in art from various cultures include the Pegtymel petroglyphs of Russia and Japanese Netsuke figurines.

Paintings, tapestries, and illustrations

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“The mycologist may liken himself to a pioneer wandering through a land filled with alternatively beautiful and fantastic shapes. A land of delicate pencilings and glowing colors where mystery sits enthroned and wise men become its worshipper.”

— Mary Elizabeth Banning, Fungi of Maryland

 

Artists, painters, illustrators, naturalists and scientists have depicted mushrooms in their artworks since centuries. Artistic representations of mushrooms might give an indirect insight into the (historical) attitudes of different cultures across time towards them. In general, more mushrooms are present in artworks from cultures considered to be mycophiles (e.g. Italy, Asian cultures) than in cultures considered mycophobes. Representation of edible species like Caesar's mushroom (Amanita caesarea) and the King bolete (Boletus edulis) is more common than depiction of toxic ones. Mushrooms abound in Italian, Flemish, Germanic, and Dutch Baroque landscapes and still lifes. Landscape paintings involving mushrooms occasionally depict mushroom or truffle hunting. The Registry of Mushrooms in Works of Art collects and describe tapestries depicting mushrooms, in particular from the Gothic and Early Renaissance (1300-1500), the High Renaissance (1500-1600), and the Flemish Baroque (ca. 1600–1750). Whereas historical British artworks tend to be considered to be influenced by a 'mycophobe' attitude, in the 19th century Victorian Fairy Paintings depicting imaginary scenes involving fairies and other fantastic creatures often featured mushrooms. A great number of Victorian era illustrators and children-book authors depicted mushrooms in their artworks, including Beatrix Potter, Hilda Boswell, Molly Brett, Arthur Rackham, Charles Robinson, and Cicely Mary Barker.

Illustration of the fungus Dumontinia tuberosa by physician, mycologist, and illustrator Charles Tulasne (1816-1884) in the book Selecta Fungorum Carpologia (1861-65). During the Victorian era, numerous scientists drew accurate illustrations of fungi, blurring the border between mycology and the arts. (Name of the original work: Peziza tuberosa parasit on Anemone nemorosa)

The Registry of Mushrooms in Works of Art from the North American Mycological Association curates hudreds of artworks featuring mushrooms (mostly paintings and illustrations from Western artists).

Notable examples of visual artists depicting mushrooms and how they contributed to both mycology and the arts are:

• Lewis David von Schweinitz (1780–1834): illustrations of over 1000 fungal species which along with his contribution to mycology earned him the title of "Father of North American Mycology." Hundreds of his paintings have been digitized by the Academy of Natural Sciences of Drexel (Philadelphia)
• Charles Tulasne (1816–1884): illustrations of fungi in the book Selecta Fungorum Carpologia (1861–65). In addition to assisting his brother with the classification and study of fungi, Charles Tulasne collaborated with Louis on numerous scientific publications. He is known for his excellent illustrations, particularly in the three-volume Selecta Fungorum Carpologia. Regarding the artistic quality of his work, Charles Tulasne is sometimes referred to as "The Audubon of Fungi".
• Mary Elizabeth Banning (1822-1903):

Fungi in art
Born	(1822-04-06)6 April 1822 Talbot County, Maryland
Died	28 February 1903(1903-02-28) (aged 80) Winchester, Virginia
Nationality	American
Scientific career
Fields	Mycology

Mary Elizabeth Banning (6 April 1822 – 28 February 1903) was an American mycologist (fungi biologist) and botanical illustrator from Maryland.

She formally described 23 previously unknown species of fungi, publishing their type descriptions in the Botanical Gazette and Charles Peck's "Annual Report of the New York State Botanist".

Career

Mary Banning is best known as the author of The Fungi of Maryland, an unpublished manuscript containing scientific descriptions, mycological anecdotes, and 174 13" by 15" watercolor paintings of fungal species. The New York State Museum describes these paintings as "extraordinary...a blend of science and folk art, scientifically accurate and lovely to look at." Banning's manuscript took twenty years to complete (1868–1888). Although the Maryland State Archives, citing Stegman, asserts that "At this point in time, no one had written a book on American fungi," and Haines states, "In 1868 there were no books from which to learn about American fungi," neither assertion is technically correct: Schweinitz's Synopsis Fungorum Carolinæ Superioris was published in 1822. Nevertheless, had Banning's opus been published, it would certainly have been the first illustrated and popularly accessible fungal flora of the southern United States.

She was inducted into the Maryland Women's Hall of Fame in 1994.

Challenges as a female mycologist

Banning eventually became the "leading mycologist in her region". Despite this, she was a woman, and she had no formal higher education. Consequently, with the exception of her mentor Peck, she found herself largely ostracized by the educated male scientific establishment of the day. Her private letters reveal her deep dissatisfaction with this state of affairs. Unable to obtain funding and having to care for her invalid mother and sister, Banning "incurred increasing financial problems."

Conversely, her mycological pursuits led to several awkward encounters with fungus-fearing locals, many of which she recounts as asides in her scientific publications. On one occasion, she rode six miles in a crowded "public conveyance" holding a basket of Phallus duplicatus — a particularly foul-smelling fungus with the common name "netted stinkhorn." By the end of the ride, Banning states that "the smell had increased to such an extent that the flies nearly devoured me, in their eagerness to get at the fungus". The other passengers maintained a stony silence for the duration of the trip. On another occasion, a man approached her, asking if she had found any "frog stools" that day. When she replied that she had not, he answered, "And it's a blessed thing you can't find 'em!...Pison [sic] things...Better let frog stools alone! That's my advice to everyone." The man then walked away, muttering about Banning: "Poor thing. Crazy, certain sure. Clean gone mad!" On yet another occasion, Banning had employed three young boys to collect mushrooms. When they brought their finds to the hotel where she was staying and asked where they might find the 'frog stool lady,' the waiter replied, "Off with you! Have you gone crazy? Who ever heard tell of a frog-stool lady?"

In the preface to her unpublished manuscript, Banning discusses the origin of her project in terms recalling then-prevalent natural theology:

"My first idea of drawing and painting the Fungi of Maryland had for its object educational training in a mission school.... I confess to a smile at my choice of a subject, feeling that for once I had stepped from the sublime to the ridiculous. :::Yet I feel satisfied with my undertaking, believing that the study of Natural Science in any of its departments has a refining influence—that when used in its truest highest sense it is the Divinely appointed means of teaching faith as well as cultivating the minds and morals."

Family life

Banning was born in 1822 in Talbot County on Maryland's eastern shore. She was the daughter of Robert Banning and Mary Macky, and was the youngest of her father's eight children (six from a previous marriage). The Bannings were a well-established Maryland family: Mary Elizabeth's grandfather was a representative at Maryland's ratification of the federal constitution, and her father was a military captain, Collector of the Port of Oxford, and Member of the Maryland House of Delegates.

In 1845, when Banning was 23, her father died. In 1855, Banning, her mother, and her sisters moved to Baltimore. By 1860, her mother and a sister had become chronically ill, and Banning became their caretaker. Throughout, she maintained an interest in natural history, finally gravitating to the study of fungi. With her own money, she bought a microscope and started to amass a scientific library and private herbarium. She also initiated a correspondence with Charles Horton Peck, a New York State Museum scientist who, by then, was well on his way to becoming "the dean of American mycologists".

Final years and legacy

Amanita banningiana, named for Mary Elizabeth Banning

By the end of the 1880s, Mary's immediate family had died, and she found herself nearly penniless, with fading eyesight and growing rheumatism. She moved into a boarding-home in Winchester, Virginia. In 1889, she ended work on her manuscript, dedicating it to Peck, with whom she had corresponded for 30 years but never met. In 1890, she shipped the manuscript to Peck at the New York State Museum, writing, "In parting from it I feel like taking leave of a beloved friend with whom I have spent many pleasant hours. Circumstances impel me to put it in a safe place." Peck placed the manuscript in a drawer, where it would remain for the next 91 years. Banning died 13 years later, in 1903. She left her remaining money to the St. John's Orphanage for Boys.

In 1981, John Haines, a mycologist attached to the New York State museum as associate scientist, discovered Banning's manuscript. The museum organized the watercolors into the exhibit "Each a Glory Bright", which has been loaned to museums around the United States. Plates from the work can be viewed at the New York State Museum's page for this collection. Mary Banning is also the namesake of the provisional species Amanita Banningiana, the "Mary Banning Slender Caesar."

Partial bibliography

• Banning, Mary E. "Notes on Fungi." Botanical Gazette 5, No. 1 (January 1880): 5-10. https://archive.org/details/botanicalgazette56hano (accessed Aug 1, 2013)
• Banning, M. E. (1881). "New Species of Fungi Found in Maryland. Agaricus (Tricholoma) cellaris". Botanical Gazette. 6 (1): 165–167. doi:10.1086/325440. JSTOR 2556568. S2CID 85395983.
• Banning, Mary E. "Maryland Fungi. I." Botanical Gazette 6, No. 4 (April 1881): 200-202. https://archive.org/details/botanicalgazette56hano (accessed Aug 1, 2013)
• Banning, Mary E. "Maryland Fungi. II." Botanical Gazette 6, No. 5 (May 1881): 210-215.https://archive.org/details/botanicalgazette56hano (accessed Aug 1, 2013)
• Banning, Mary E. (1882). "Preservative for Fungi". Bulletin of the Torrey Botanical Club. 9 (12): 153. JSTOR 2476196.
• Banning, Mary E. (1882). "The Tuckahoe". Bulletin of the Torrey Botanical Club. 9 (10): 125–126. doi:10.2307/2476029. JSTOR 2476029.
• Banning, Mary E. The fungi of Maryland. Unpublished manuscript in the possession of the New York State Museum. Plates accessible at http://www.nysm.nysed.gov/treasures/explore.cfm?coll=29 (Accessed Aug 1, 2013)

References

• Ernst Häckel (1834–1919): illustrations of basidiomycota, ascomycota, lichens in Kunstformen der Natur (1904).

Kunstformen der Natur (Art Forms in Nature)

Author	Ernst Haeckel
Country	Germany
Language	German
Subject	Natural history, biology, art
Genre	Non-fiction
Publication date	Published in sets between 1899 and 1904; collectively in two volumes in 1904

The 8th print, Discomedusae. The center and bottom-center images are Desmonema annasethe; the tentacles reminded Haeckel of his late wife's long flowing hair.

Kunstformen der Natur (known in English as Art Forms in Nature) is a book of lithographic and halftone prints by German biologist Ernst Haeckel.

Publication

Originally published in sets of ten between 1899 and 1904 and collectively in two volumes in 1904, it consists of 100 prints of various organisms, many of which were first described by Haeckel himself. Over the course of his career, over 1000 prints were produced based on Haeckel's sketches and watercolors; many of the best of these were chosen for Kunstformen der Natur, translated from sketch to print by lithographer Adolf Giltsch.

A second edition of Kunstformen, containing only 30 prints, was produced in 1914.

Themes

According to Haeckel scholar Olaf Breidbach, the work was "not just a book of illustrations but also the summation of his view of the world." The over-riding themes of the Kunstformen plates are symmetry and level of organization. The subjects were selected to embody these to the full, from the scale patterns of boxfishes to the spirals of ammonites to the perfect symmetries of jellies and microorganisms, while images composing each plate are arranged for maximum visual impact.

Among the notable prints are numerous radiolarians, which Haeckel helped to popularize among amateur microscopists; at least one example is found in almost every set of 10. Cnidaria also feature prominently throughout the book, including sea anemones as well as Siphonophorae, Semaeostomeae, and other medusae. The first set included Desmonema annasethe (now Cyanea annasethe), a particularly striking jellyfish that Haeckel observed and described shortly after the death of his wife Anna Sethe.

Influence

Kunstformen der Natur was influential in early 20th-century art, architecture, and design, bridging the gap between science and art. In particular, many artists associated with Art Nouveau were influenced by Haeckel's images, including René Binet, Karl Blossfeldt, Hans Christiansen, and Émile Gallé. One prominent example is the Amsterdam Commodities Exchange designed by Hendrik Petrus Berlage: it was in part inspired by Kunstformen illustrations.

Gallery of prints

Haeckel's original classifications appear in italics.

• 
  Sea anemone (Actiniae)
• 
  Anthomedusa (Anthomedusae)
• 
  Siphonophore (Siphonophorae)
• 
  Diatoms (Diatomea)
• 
  Antelope (Antilopina)
• 
  Arachnid (Arachnida)
• 
  Ascidian (Ascidiae)
• 
  Frog (Batrachia)
• 
  Bryozoa
• 
  Turtle (Chelonia)
• 
  Ciliate (Ciliata)
• 
  Annelid (Chaetopoda)
• 
  Bat (Chiroptera)
• 
  Conifer (Coniferae)
• 
  Copepod (Copepoda)
• 
  Box jellyfish (Cubomedusae)
• 
  Decapods (Decapoda)
• 
  Cephalopod (Gamochonia)
• 
  Fern (Filicinae)
• 
  Red Algae (Florideae)
• 
  Lichen (Lichenes)
• 
  Lizard (Lacertilia)
• 
  Moss (Muscinae)
• 
  Nepenthes (Nepenthaceae)
• 
  Nudibranch (Nudibranchia)
• 
  Trachylina (Narcomedusae)
• 
  Orchid (Orchidae)
• 
  Boxfish (Ostraciontes)
• 
  Prosobranchia (obsolete classification)
• 
  Siphonophorae
• 
  Polycystine (Spumellaria)
• 
  Radiolarian (Stephoidea)
• 
  Moth (Tineida)
• 
  Hummingbird (Trochilidae)
• 
  Tubularia (Tubulariae)
• 
• 
  Cirripedia, Kunstformen der Natur, (1904).

See also

• On Growth and Form

References

• Breidbach, Olaf. Visions of Nature: The Art and Science of Ernst Haeckel. Prestel Verlag: Munich, 2006.

External links

• Marine Biological Laboratory Library - An exhibition of material on Haeckel, including background on many Kunstformen der Natur plates.
• University Art Gallery, University of Massachusetts Dartmouth - An Ernst Haeckel exhibition from 2005 pairing prints from Kunstformen der Natur with modern sculptures.
• Kurt Stüber's Biolib - An online version of Kunstformen der Natur with scanned images of the 100 plates, accompanying descriptions, table of contents and supplemental pages (in German).
• Kunstformen der Natur (PDF)

• Beatrix Potter (1866-1943): Curious as to how fungi reproduced, Potter began microscopic drawings of fungus spores (the agarics) and in 1895 developed a theory of their germination. Through the connections of her uncle Sir Henry Enfield Roscoe, a chemist and vice-chancellor of the University of London, she consulted with botanists at Kew Gardens, convincing George Massee of her ability to germinate spores and her theory of hybridisation. She did not believe in the theory of symbiosis proposed by Simon Schwendener, the German mycologist, as previously thought; instead, she proposed a more independent process of reproduction.
• Violetta White Delafield (1875–1949): creations of around 600 illustrations of fungi. Delafield created hundreds of annotated watercolors of fungi and plants, noted for their level of detail; she made a note of the collection location, a detailed specimen description and analysed the cellular structure of the fungus with the help of a microscope. Her extensive illustrations are particularly significant as fungal specimens tend to deteriorate soon after collection and would often change their colour and form. Delafield's significant collection of specimen was left to the Fungal Herbarium at the New York Botanical Garden, her papers and research materials on mycology and horticulture are held with the Delafield family papers by the University of Princeton. A selection of her work was exhibited in 2019 at Bard College as part of the ‘Fruiting Bodies’ exhibition and has been preserved in a digital collection.

Photography

A countless number of amateur and professional photographs of mushrooms abound on the internet and on social media. Non-fiction books about fungi, especially those involving identification of fungi, includes photographs of fungal species and their fruiting bodies. The book by Scott Chimileski and Roberto Kolter Life at the Edge of Sight: A Photographic Exploration of the Microbial World showcases 'the invisible world waiting in plain sight,' including fungi.

Since 2005, the North American Mycological Association (NAMA) organises an annual Photography Art Contest on mushrooms and fungi.

Literature

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Mushrooms in fiction

See also: Category:Fictional fungi

"Ye elves of hills [...] whose pastime is to make midnight mushrumps."

— William Shakespeare, The Tempest Act 5, Scene 1 (ca. 1610)

Work of literary fiction involving mushrooms and fungi are often linked to infection, decay, toxicity, mystery, and fantasy, ambiguity, and thus have mostly a negative connotation. There are a few examples of mushrooms depicted or involved in a positive way, some examples being the children book The Wonderful Flight to the Mushroom Planet by Eleanor Cameron or the science-fiction novel Omnivore by Piers Anthony. In line with the assumption that Russian society is traditionally more affine to mushrooms than Western society, a scene of mushroom foraging in Leo Tolstoy's Anna Karenina is associated with love, family, and a sense of commonality. During the Victorian era, fungi started to acquire a more playful, childish, or jolly role in works of literary fiction. The author, artist, illustrator, and mycologist Beatrix Potter created meticulous and accurate illustrations of mushrooms, including in her children-book series of Peter Rabbit. Several renowned authors have used fungi as plot device. These include Percy Shelly, Lord Alfred Tennyson, Sir Arthur Conan Doyle, D.H. Lawrence, H.G. Wells, Ray Bradbury, and more.

Fungi have been a common trope in the science fiction, horror, supernatural, fantasy and crime fiction genres. Fungi have a long tradition in science fiction. In Ray Bradbury's Come into My Cellar, mushrooms are alien invasors threatening society. The short story is one of the rare examples in which several forms of fungi appears (spores and mushrooms): In the story, an alien form of spores from fungi lands on Earth and compels humans, and kids in particular, to grow mushrooms and infect more persons, thus using humans as a medium of propagation of fungi through mind control.

Fungi have occasionally appeared in the murder mystery literature due to their toxicity. Crime and detective writer Agatha Christie has repeatedly used mushrooms as murder weapon in her crime fiction. The use of (toxic) mushrooms in fiction do not often reflect reality, either because a misidentified species is used (for example, a non-toxic one), because the preparation or intake of the toxic is wrong (for example, when not enough toxin is present), or because the progress of poisoning is unrealistic (for example, if the toxin kills too quickly).

A non-exhaustive list of fictional stories involving mushrooms is given below:

• Anna Karenina (1878), a realist novel by Leo Tolstoy
• Journey to the Center of the Earth (1864), a science fiction novel by Jules Verne
• Alice's Adventures in Wonderland (1865), a children novel by Lewis Carroll
• The Purple Pileus (1896), a fantasy short story by H. G. Wells
• The Voice in the Night (1907), a horror short story by William Hope Hogson
• The Thumb Mark of St Peter (1928), a murder mystery short story by Agatha Christie
• The Documents in the Case (1930), a murder mystery story by Dorothy L. Sayer
• The Puff-Ball Menace (1933), a supernatural story by John Wyndham
• Come into My Cellar (1962), a science fiction short story by Ray Bradbury
• Ambergris trilogy (2001, 2006, 2009), a fantasy collection by Jeff VanderMeer

The 'Bad Bug Bookclub' by the Manchester Metropolitan University, a regular in-person and online book club organised by Joanna Verran, reads and discusses litrary works on microorganisms, including fungi; books with fictional fungi include The Girl with all the Gifts by M.R. Carey, Trouble with Lichen by John Wyndham, and others.

The quarterly periodical FUNGI Magazine runs a regular feature called ''Bookshelf Fungi'' reviewing fiction and non-fiction books on fungi.

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Mushrooms in poetry

Had nature any outcast face
Could she a son contemn
Had nature an Iscariot
That mushroom – it is him

— Emily Dickinson, The Mushroom (1896)

Here is the handful
of shadow I have brought back to you:
this decay, this hope, this mouth–
full of dirt, this poetry

— Margaret Atwood, Mushrooms (1981)

In poetry, as in literature, fungi are historically associated with negative feelings or sentiments, although, together with the rising popularity of fungi, this trend might hold less true in recent years. The poem The Mushroom (1896) by Emily Dickinson is unsympathetic towards mushrooms. American author of weird horror and supernatural fiction H. P. Lovercraft created a collection of cosmic horror sonnets with fungi as subject called Fungi from Yuggoth (1929–30).

Margaret Atwood's poem Mushrooms (1981) explores the topics of the life cycle and nature. The poem by Neil Gaiman, The Mushrooms Hunter, is a poem touching, through the lens of mushroom hunting throughout history, on the topics of womanhood, human creation, and destruction. The poem was written for 'Universe in Verse,' a festival combining science with poetry, and won the Rhysling Award for best long poem in 2017. The poem features in a short animated video with the voice-over of Amanda Palmer.

Storytelling, oral tradition, myth, and folklore

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See also: Ethnomycology

"I am … a mushroom
On whom the dew of heaven drops now and then."

— John Ford, The Broken Heart (1633)

Through storytelling and oral tradition, fungi have had an conspicuous influence in mythology (defind as the personification of deities and elementals) and folklore (defined as popular traditions and beliefs) and religions across various latitudes, civilizations, and historical epochs. The entheogen (psychoactive) properties of certain fungi has arguably contributed to the involvement of fungi in myth and folklore.

The distinction between literature, folklore, and myth is not always clear cut, and occasionally open to interpretations. Some writers argue that fungi have inspired numerous myths, and vice versa that many myths can be re-interpreted through the lens of fungal ecology. In her essay Jesus if a Fungal God, author Sophie Strand writes:

"As we learn more about fungi, let us embrace that they have always been here. Beneath our feet. And inside our most popular myths."

A gnome carries the fly agaric (Amanita muscaria) on a German Christmas greeting card (ca. 1900) saying 'Viel Glück im neuen Jahre!' (All the best for the new year!). The fly agaric has inspired countless folklore tales and entered mainstream mushroom culture.

The painting Judas Hangs Himself (ca. 1890) by James Tissot. Naming of mushrooms have been inspired by folklore. The common naming of the 'jelly ear' fungus (Auricularia auricula-judae) is 'Jew's ear,' derived from the belief that Juda Iscariot hanged himself on a elder tree, where the fungus often grows.

Occasionally, the involvement of fungi in myth and folklore is driven by allegory, cultural practices, or popular interpretations. For example, given the cultural relevance and prevalence of fermented (alcoholic) beverages throughout history, there are numerous deities associated with wine and beer, which can be regarded as an indirect effect of fungi in the arts.

Fungi (yeasts) play a conspicuous role in several religions, for example through fermentation (e.g. wine) and leavening (e.g. bread). In the Parable of the Leaven, one of the Parables of Jesus, the growth of the Kingdom of God is akin to the leavening of bread through yeast. According ton Matthew 13:33 (and, similarly, to Luke 13:20-21):

“He told them still another parable: 'The kingdom of heaven is like yeast that a woman took and mixed into about sixty pounds[a] of flour until it worked all through the dough.'”

However, yeast is associated with corruption in other passages of the New Testament, as in Luke 12:1:

“Jesus began to speak first to his disciples, saying: 'Be on your guard against the yeast of the Pharisees, which is hypocrisy.'”

Mead, the alcoholic beverage from fermentation of honey, has also influenced mythology. Mead was produced in ancient times throughout Europe, Africa, and Asia, and has played an important role in the mythology of some peoples. In Norse mythology, for example, the Mead of Poetry, crafted from the blood of Kvasir, would turn anyone who drank it into a poet or scholar.

Some scholars argue that the Egyptian God of the afterlife Osiris is a personification of entheogenic mushrooms. As one of the evidences they provide, they indicate how Egyptian Crowns are shaped like primordia of Psilocybe cubensis mushrooms. The Egyptian tale known as Cheops and the Magicians illustrates growth of mushrooms on barley.

In the Chinese classic tale The Mountain and the Sea, the soul of a young woman becomes a mushroom as a symbol of immortality.

In Lithuanian and Baltic mythology, fungi are considered the fingers of Velnias, the God of the underworld, reaching up from the underground to feed the poor. Baltic and Ugric religions had mushrooms elements, including a "Mother of Mushrooms" (Sěņu māte). The tale The War of the Mushrooms is a popular folk tale in several Slavic cultures. After the outbreak of the Ukraine-Russia war in 2022, an exhibition at the Ukrainian Museum in New York revisited the classic story in light of current events. The supernatural being Baba Yaga in Slavik folklore is often associated with mushrooms. In some Russian tales it often appears a villainous wizard called Mukhomor, literally 'poison mushroom,' which is asusmed to be derived from the fly agaric.

The fly agaric (Amanita muscaria) is a mushroom with characteristic red cap and white dots, and has greatly infiltrated folklore with mainstream popularity. According to several interpretations, the legendary figure of Santa Claus is heavily influenced by the fly agaric; anecdotal evidences include the use by Saami shamans in the Lapland region, who would visit the homes of people by reindeer-drawn sleds and enter through the chimney when the entrance door was stuck by snowfalls; the fondness of reindeers in eating fly agaric mushrooms; the belief by Saami people that whoever eats an Amanita muscaria will resemble it, becoming among other plump and reddish; and the sense of flying that consumption of fly agaric might induce.

See also: Amanita muscaria § Cultural depictions

The stinkhorn Phallus indusiatus (also called 'veiled lady' among many different common names) has entered folklore across many cultures, probably due to its peculiar shape. In French, P. indusiatus is commonly called 'le satyre voilé' ('the veiled satyr,' from the male nature spirit in Greek mythology).

Mushrooms have been used as an instrument for anti-Semitic discrimination or propaganda over the centuries. As Jews have a long tradition of eating mushrooms, which are considered Kosher in Jewish dietary law, mushrooms have been referred to as "Jew's Meat". The connotation was dispregiative, especially during the Middle Ages, when mushrooms where considered as toxic and disgusting. In the infamous children-book Der Giftpilz during Nazi Germany, Jews are depicted as poisonous and difficult to distinguish from 'Gentiles'.

Non-fictional books about fungi

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See also: List of books about mushrooms

There is a conspicuous corpus of literature on mushrooms, including foraging, identifying, growing, and cultivating fungi. Although these books are non-fictional, the works are often excellent examples of storytelling and tinkering, and are a fundamental source for fungi enthusiasts and the growing do-it-yourself community. These works are not only an important source but also a way for artists to converge and experiment with fungi. The authors are often pioneers and contribute to the increased popularity, popularisation, and development of mycology, fungal ecology, mycoremediation, fungal conservation, biocontrol, medicinal fungi, mushroom gathering and identification, and fungal research. The book The Mushroom at the End of the World by Chinese-American anthropologist Anna Lowenhaupt Tsing on matsutake mushrooms offers insights into the cultural relevance and the significance of fungi for modern society, circularity, and decay. Other books proposed speculative or disputed theories on the cultural influence of fungi throughout history, like The Sacred Mushroom and the Cross by John Allegro, and were received critically by fellow mycologists.

The online book club 'MycoBookClub' discusses monthly a selection of mostly non-fiction books on fungi on Twitter.

Cinema, TV shows, and motion pictures

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The experimental short movie Beneath by Beth Walker (UK), presented at 2022 Fungi Film Festival.

The comedy short movie Shroom Mates by Rosie Windsor (UK), presented at 2022 Fungi Film Festival.

Adaptations of literary fiction into motion pictures follow similar tropes present in science fiction, horror, supernatural, and crime fiction genres. Examples of adaptations include the 2016 British post-apocalyptic science fiction horror movie The Girl with All the Gifts, based on the novel with the same title, and the 1963 Japanese horror film Matango (マタンゴ) directed by Ishirō Honda, partially based on William Hope Hodgson's short story The Voice in the Night (1907).

The 2000 documentary Pegtymel by director Andrei Golovnev shows the daily life of Chukchi indigenous reindeer herders on the Pegtymel River (Siberia), where the eponymous petroglyphs (representing mushroom-headed people) are located, including their consumption of hallucinogenic fly agaric mushrooms. The commercially successful 2019 documentary Fantastic Fungi released on Netflix includes interviews with renowned mycologist Paul Stamets and presents the intriguing world of fungi from a medical and ecological perspective with the use of narration, time-lapse photography, and interviews. The documentary covers fungi and not only mushrooms. The documentary The Mushroom Speaks (2001) by Marion Neumann covers topics such as decay, bioremediation, and symbiosis by following scientists, experts, and fungal pioneers.

Recently, new film festivals devoted entirely to fungi have been created and solicit submissions on all aspects of fungal biology from both amateur and professional filmmakers. Screening ar online or at specific venues. Most notably are the Fungi Film Festival (since 2021) co-created by Radical Mycology author Peter McCoy and the UK Fungus Day Film Festival (since 2022) by the British Mycological Society. Topics and themes often present at the 2022 Fungi Film Festival are personification of mushrooms, experimental/conceptual representation of fungal forms, and utilization of mushrooms for their (hallucinogenic) properties.

Performative arts (theatre, comedy, dance, performance art)

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The American stand-up comedian and satirist Bill Hicks drew inspiration from Terence McKenna's 'Stoned Ape Theory' in his 1993 show Revelation.

Comic books and video games

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In the Belgian comic franchise The Smurfs, the characters with the same name inhabit houses resembling mushrooms. American fantasy and science fiction comic book artist Frank Frazetta illustrated the cover image of the 1964 edition of the novel The Secret People (1935) by John Beynon (pseudonym of John Wyndham), in which fictive 'little people' inhabit areas with giant mushrooms. Dave Gibbon's comic strip Come into My Cellar is based on Ray Bradbury's short story with the same name.

Game designer Shigeru Miyamoto acknowledged Lewis Carroll's Alice in Wonderland as direct influence for the 'super mushroom' in developing Nintendo's Super Mario video game. The celebrated video game franchise The Last of Us is set in post-apocalyptic US, after a mutant fungus wiped off humanity, turning infected into zombies. Further video games where mushrooms appear as healt-boosting collectibles or poisonous mushrooms are Skyrim (2011), Stardew Valley (2016), and Zelda: Breth of the Wild (2017).

Music

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"Hoc cantum dedicatum est, qui fungus amant"

"This song is dedicated to those who love mushrooms."

— Lepo Sumera, Mushroom Cantata

Mushrooms have an influence in music as subject, cultural reference, or as medium for music creation. Numerous musicians, bands, composers, and lyricists mentioned or drew inspiration from fungi. A music gerne called Fungi from the British Virgin Islands is defined as a mixture of many styles and instruments. The Czech composer and mycologist Václav Hálek (1937-2014) claimed to have created numerous musical works inspired by fungi. American composer John Cage (1912-1992) was an enthusiastic amateur mycologist and co-founder the New York Mycological Society. Music can be created utilizing fungi, as in the process of bio-sonification.

Music inspired by mushrooms

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Numerous musicians, bands, composers, and lyricists mentioned or drew inspiration from fungi, like the Israeli psychedelic trance band Infected Mushroom, the US heavy metal band Mushroomhead, Russian romantic composer Modest Mussorgsky's (1839-1881) song Gathering Mushrooms, Igor Stravinsky's (1882-1971) How the Mushrooms went to War, and many more. In Women Gathering Mushrooms, the musicologist Louis Sarno (1954-2017) recorded women from the Central Africa Mbenga pygmy tribe of the Aka (also Biaka, Bayaka, Babenzele) singing while collecting mushrooms, resulting in a polyphonic composition. According to mycologist and author Merlin Sheldrake, the activity of the gatherers above ground mirrors the fungal life below ground, as "mycelium is polyphony in bodily form." Icelandic avant-guard musician Björk's 2022 album Fossora (including tracks such as Mycelia, Sorrowful Soil, and Fungal City) is referred to as her "mushroom album." 'Fossora' is the feminine declination of the Latin fossore, meaning "she who digs."

Music created with mushrooms

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Fungi are occasionally a direct medium for the creation of music. With the use of sonification and synthezisers, musicians and bioartists are able to create sounds and music by converting mushrooms bioelectric signals. Some artists creating music by sonificating mushrooms note that different mushrooms produce different sounds: for example, Ganoderma lucidum produces melodic sounds, while Pleurotus ostreatus produces constant sounds. The 'Nanotopia Midnight Mushroom Music' is a radio station devoted to streaming mushroom-generated music. The rap artists 'FungiFlows' composes lyrics inspired by fungi and mushrooms while wearing a fly-agaric-shaped hat. The Czech composer and mycologist Václav Hálek (1937-2014) stated to have composed over 1,500 symphonies inspired by fungi, including the composition called Mycosymphony.

Architecture and sculptures

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In architecture and sculpture, mushrooms are mostly represented or showcased. Mushrooms are carved in buildings or depicted in sculptures or potteries, like pre-Columbian pottery mushrooms from Mesoamerica.

The Porter's Lodge pavilion at the entrance of Park Güell features a lookout tower with a mushroom-shaped dome.

At the entrance of Park Güell by Catalan modernist architect Antoni Gaudí (1852-1926), the Porter's Lodge pavilion features a lookout tower with a mushroom-shaped dome, probably inspired by Amanita muscaria or by stinkhorns.

The mycologist William Dillon Weston (1899-1953) created glass sculptures of microfungi, mostly plant pathogens, to fight bouts of insomnia. The artworks represent either magnified fungi (usually up to 400X times for fungi; up to 1200X for spores) or real-size plants affected by fungi (like in Ustilago maydis and Phytophthora infestans) and are made of transparent or opaque glass, although coloured glass was used when needed. The sculptures are mostly between 5–20 cm in size and often do not have a base and stand on the mycelium. Almost a hundred glass sculptures are conserved at the Whipple Museum in Cambridge (UK). Fungi represented are among others species from the genera Alternaria, Botrytis, Penicillium, Cordyceps, Sclerotinia, Fusarium, Puccinia as well as spores (ascospores, basidiospores). The other known example of glass sculptures representing (among others) fungi is the Blaschka Glass Flowers at Harvard Museum in Cambridge, Massachusetts (US).

The sculpture Triple Mycomorph by Bernard Reynolds (1915–1997) at Christchurch Mansion holds resemblance with the stinkhorn mushroom Phallus indusiatus.

Mushrooms are occasionally showcased by artists who collect, manipulate, preserve, and exhibit them, as in the 'Mind The Fungi' exhibition (2019-2020) at Futurium in Berlin (Germany).

Culinary arts

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Fungi enter cuisine mostly as fruiting bodies (mushrooms). Mushrooms are a source of protein in diet and a staple in many cultures and cuisines, and a common ingredient in many recipes worldwide. Most mushrooms sold commercially are the button mushroom (Agaricus bisporus), commonly know as champignons. Many mushrooms, including some coveted in haute cuisine, like truffles and boletus, cannot be cultivated and need to be harvested. Due to their dietary properties and their suitability as meat substitute, mushrooms can be considered a novel culinary trend, including the cultivation and consumption of species which only recently became popular in cooking, like Cordyceps.

Many fungi are considered delicacies in cuisine and gastronomy. Truffles, which are occasionally confused with tubers (storage organs in plants, like potatoes), are subterranean fruiting bodies (that is, mushrooms which grow below ground) of certain fungi belonging to the genera Tuber, Geopora, Peziza, Choiromyces, and others. Truffles have developed a distinctive aroma as spore-dispersion strategy: Instead of relying on wind and other mechanical means, truffles attract animals which eat them and carry their spores to new locations after defecation.

The North American Mycological Association (NAMA) hosts a series of resources to encourage all aspects of 'mycophagy.'

Infection of maize corn with the plant pathogen Ustilago maydis leads to a tumor in the plant, which is consumed as a delicacy in Mexico (called huitlacoche).

Corn smut (Ustilago maydis) is a plant pathogen infecting maize and teosinte. The delicacy is actually the gall (a tumor-like growth on the infected plant) induced during the fungus infective cycle, rather than a fungus fruiting body. The fungus infection gives the plant a 'mushroomy' taste, and it is used in Mexico as the delicacy huitlacoche, eaten usually as a filling in quesadillas.

"Cuitlacoche" redirects here. For the genus of birds otherwise known as "Cuitlacoche", see Toxostoma.

Corn smut
Ustilago maydis diploid teleospores
Scientific classification
Domain:	Eukaryota
Kingdom:	Fungi
Division:	Basidiomycota
Class:	Ustilaginomycetes
Order:	Ustilaginales
Family:	Ustilaginaceae
Genus:	Ustilago
Species:	U. maydis
Binomial name
Ustilago maydis (DC.) Corda

Corn smut
Huitlacoche
Common names	huitlacoche (Mexico), blister smut of maize, boil smut of maize, common smut of maize, corn truffle
Causal agents	Ustilago maydis
Hosts	maize and teosinte
EPPO Code	USTIMA
Distribution	Worldwide, where corn is grown

Corn smut is a plant disease caused by the pathogenic fungus Ustilago maydis. One of several cereal crop pathogens called smut, the fungus forms galls on all above-ground parts of corn species such as maize and teosinte. The infected corn is edible; in Mexico, it is considered a delicacy, called huitlacoche, often eaten as a filling in quesadillas and other tortilla-based dishes, as well as in soups.

Etymology

In Mexico, corn smut is known as huitlacoche (Spanish pronunciation: [(ɡ)witlaˈkotʃe], sometimes spelled cuitlacoche). This word entered Spanish in Mexico from Classical Nahuatl, though the Nahuatl words from which huitlacoche is derived are debated. In modern Nahuatl, the word for huitlacoche is cuitlacochin (Nahuatl pronunciation: [kʷit͡ɬɑˈkot͡ʃin]), and some sources deem cuitlacochi to be the classical form.

Some sources wrongly give the etymology as coming from the Nahuatl words cuitlatl [ˈkʷit͡ɬɑt͡ɬ] ("excrement" or "rear-end", actually meaning "excrescence") and cochtli [ˈkot͡ʃt͡ɬi] ("sleeping", from cochi "to sleep"), thus giving a combined mismeaning of "sleeping/hibernating excrement", but actually meaning "sleeping excrescence", referring to the fact that the fungus grows between the kernels and impedes them from developing, thus they remain "sleeping".

A second group of sources deem the word to mean "raven's excrement". These sources appear to be combining the word cuitlacoche for "thrasher" with cuitla, meaning "excrement", actually meaning "excrescence". However, the avian meaning of cuitlacoche derives from the Nahuatl word "song" cuīcatl [ˈkʷiːkɑt͡ɬ], itself from the verb "to sing" cuīca [ˈkʷiːkɑ]. This root then clashes with this reconstruction's second claim that the segment cuitla- comes from cuitla ("excrement").

One source derives the meaning as "corn excrescence", using cuītla again and "maize" tlaōlli [t͡ɬɑˈoːlːi]. This requires the linguistically unlikely evolution of tlaōlli "maize" into tlacoche.

In Peru, it is known as chumo or pacho.

Taxonomy

U. maydis is the best known and studied of the Ustilaginomycetes, a sub class of basidiomycota, and so is often used as the exemplar species when talking about its entire class.

Characteristics

The fungus infects all parts of the host plant by invading the ovaries of its host. The infection causes the corn kernels to swell up into tumor-like galls, whose tissues, texture, and developmental pattern are mushroom-like. The galls grow to 4 to 5 inches in diameter. These galls are made up of hypertrophied cells of the infected plant, along with resulting fungal threads, and blue-black spores. These dark-colored spores give the cob a burned, scorched appearance; this is the origin of the generic name Ustilago, from the Latin word ustilare (to burn).

This ear of corn has been infected with Ustilago maydis.

Biology

Life cycle

Ustilago maydis haploid sporidia

When grown in the lab on very simple media, it behaves like baker's yeast, forming single cells called sporidia. These cells multiply by budding off daughter cells. When two compatible sporidia meet on the surface of the plant, however, they switch to a different mode of growth. First, they produce one or another pheromone, and begin producing one or the other type of pheromone receptor - this depends on mating type a or b, as determined by alleles at two unlinked mating loci. If this signalling is successful they then send out conjugation tubes to find each other, after which they fuse and make a hypha to enter the maize plant. Hyphae growing in the plant are dikaryotic; they possess two haploid nuclei per hyphal compartment. In contrast to sporidia, the dikaryotic phase of U. maydis only occurs during successful infection of a maize plant, and cannot be maintained in the laboratory.

Proliferation of the fungus inside the plant leads to disease symptoms such as chlorosis, anthocyanin formation, reduced growth, and the appearance of tumors harboring the developing teliospores. These teliospores help to overwinter the pathogen into the next season. They survive in the soil.

Mature tumors release spores that are dispersed by rain and wind. Under appropriate conditions, a metabasidium is formed in which meiosis occurs. Resulting haploid nuclei migrate into elongated single cells. These cells detach from the metabasidium to become the sporidia, thus completing the life cycle.

Host/pathogen conflict

Plants have evolved efficient defense systems against pathogenic microbes. A rapid plant defense reaction after pathogen attack is the oxidative burst, which involves the production of reactive oxygen species at the site of the attempted invasion. As a pathogen, U. maydis can respond to such an oxidative burst by an oxidative stress response, regulated by gene YAP1. This response protects U. maydis from the host attack, and is necessary for the pathogen's virulence. Furthermore, U. maydis has a well-established recombinational DNA repair system. This repair system involves a homolog of Rad51 that has a very similar sequence and size to its mammalian counterparts. This system also involves a protein, Rec2 that is more distantly related to Rad51, and Brh2 protein that is a streamlined version of the mammalian Breast Cancer 2 (BRCA2) protein. When any of these proteins is inactivated, sensitivity of U. maydis to DNA damaging agents is increased. Also mitotic recombination becomes deficient, mutation frequency increases and meiosis fails to complete. These observations suggest that recombinational repair during mitosis and meiosis in U. maydis may assist the pathogen in surviving DNA damage arising from the host's oxidative defensive response to infection, as well as from other DNA damaging agents.

Proteome

U. maydis is known to produce four Gα proteins, and one each of Gβ and Gγ.

Management

Losses from corn smut can vary greatly, however annual yield losses rarely exceed 2% when resistant cultivars are planted. This disease can have a large economic impact on sweet corn, specifically when smut galls replace the kernels. There are many ways to control and manage corn smut; however, corn smut cannot be controlled by any common fungicide at this time, as Ustilago maydis infects individual corn kernels instead of infecting the entire cob, like head smut. Some beneficial ways to contain corn smut include resistant corn plants, crop rotation, and avoiding mechanical injury to the plant. A mechanical injury can cause the corn to become easily accessible to Ustilago maydis, enhancing infection. Additionally, clearing the planting area of debris can help control corn smut, as the teliospores from corn smut overwinter in debris. This is not the best practice, though, because corn smut can also overwinter in the soil; crop rotation is recommended. Lastly, as excess nitrogen in the soil augments infection rate, using fertilizer with low nitrogen levels, or just limiting the amount of nitrogen in the soil proves to be another way to control corn smut.

Ear of corn infected with Ustilago maydis

Environment

Although not all the conditions that favor growth of Ustilago maydis are known, there are certain environments where corn smut seems to thrive, depending on both abiotic and biotic factors. Hot and dry weather during pollination followed by a heavy rainy season appear to improve the pathogenicity of corn smut. Furthermore, excess manure (and therefore nitrogen) in the soil also increases pathogenicity. Not only do these abiotic factors increase infectability, they also increase disease spread. High winds and heavy rain also increase disease spread as the spores of corn smut can be more easily transmitted. Other biotic factors largely have to do with the extent by which humans interact with the corn and corn smut. If corn debris is not cleared at the end of the season, the spores can overwinter in the corn fragments and live to infect another generation. Finally, humans wounding the corn (with shears or other tools of the like) present the opportunity for corn smut to easily enter the plant.

Non-culinary uses

Model organism

The yeast-like growth of U. maydis makes it an appealing model organism for research, although its relevance in nature is unknown. The fungus is exceptionally well-suited for genetic modification. This allows researchers to study the interaction between the fungus and its host with relative ease. The availability of the entire genome is another advantage of this fungus as a model organism.

U. maydis is not only used to study plant disease, but it also is used to study plant genetics. In 1996, a study on U. maydis genetics led to the discovery of synthesis-dependent strand annealing, a method of homologous recombination used in DNA repair. Other studies in the fungus have also investigated the role of the cytoskeleton in polarized growth. It is largely due to work with U. maydis that the function of the breast-cancer gene BRCA2 is now known. The fungus is mostly studied as model organism for host pathogen interaction and delivery of effectors protein.

Industrial biotechnology

Ustilago maydis is able to produce a broad range of valuable chemicals such as ustilagic acid, itaconic acid, malic acid, and hydroxyparaconic acid. With this ability it is gaining more and more relevance for industrial applications.

Culinary use

See also: List of delicacies

Smut feeds on the corn plant and decreases the yield. Smut-infected crops are often destroyed, although some farmers use them to prepare silage. However, the immature infected galls are still edible, and in Mexico they are highly esteemed as a delicacy. It is known as huitlacoche, and sold for a significantly higher price than uninfected corn. The consumption of corn smut in Mexico originated directly from Aztec cuisine. For culinary use, the galls are harvested while still immature — fully mature galls are dry and almost entirely spore-filled. The immature galls, gathered two to three weeks after an ear of corn is infected, still retain moisture and, when cooked, have a flavor described as mushroom-like, sweet, savory, woody, and earthy. Flavor compounds include sotolon and vanillin, as well as the sugar glucose.

Huitlacoche is a source of the essential amino acid lysine, which the body requires but cannot manufacture. It also contains levels of beta-glucans similar to, and protein content equal or superior to, most edible fungi.

The fungus has had difficulty entering into the American and European diets as most farmers see it as blight, despite attempts by government and high-profile chefs to introduce it. In the mid-1990s, due to demand created by high-end restaurants, Pennsylvania and Florida farms were allowed by the United States Department of Agriculture (USDA) to intentionally infect corn with huitlacoche. Most observers consider the program to have had little impact, although the initiative is still in progress. The cursory show of interest is significant because the USDA has spent a considerable amount of time and money trying to eradicate corn smut in the United States. Moreover, in 1989, the James Beard Foundation held a high-profile huitlacoche dinner, prepared by Josefina Howard, chef at Rosa Mexicano restaurant. This dinner tried to get Americans to eat more of it by renaming it the "Mexican truffle" and it is often compared to truffles in food articles describing its taste and texture.

Native American tribes in North America ate corn smut as well. The Hidatsa tribe of North Dakota's practice of preparing and eating corn smut is described vividly in Buffalo Bird Woman's Garden.

Native Americans of the American Southwest, including the Zuni people, have used corn smut in an attempt to induce labor. It has similar medicinal effects to ergot, but weaker, due to the presence of the chemical ustilagine.

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  Huitlacoche corn taco
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  Quesadilla de huitlacoche, as it is often served in central Mexico
• 
  Huitlacoche for sale in the produce department of a Soriana store in Mexico

Recipes of Mexico

A simple Mexican-style succotash can be made from chorizo, onions, garlic, serrano peppers, huitlacoche, and shrimp with salsa taquera. The mild, earthy flavors of the huitlacoche blend nicely with the fats of the chorizo and bond to mellow out the heat from the peppers and salsa.

Another Maya favorite on the Riviera Maya (Cancun to Tulum) is to add huitlacoche to omelettes. Its earthy flavors bond with the fats that cook the eggs to mellow the flavors into a truffle-like taste.

Huitlacoche is also popular in quesadillas with Mexican cheese, sautéed onions, and tomatoes.

The blueish color transforms into the recognizable black color only with heat. Any dish with huitlacoche must include a slow simmer of the fungus until it becomes black, which also removes most of the starch of the corn, and what is left is a black oily paste.

Availability

In Mexico, huitlacoche is mostly consumed fresh and can be purchased at restaurants or street or farmer's markets throughout the country and, to a much lesser extent, can also be purchased as a canned good in some markets and via the internet. Farmers in the countryside spread the spores around intentionally to create more of the fungus. In some parts of the country, they call the fungus "hongo de maiz", i.e. "maize fungus".

Nutritional value

When corn smut grows on a corn cob, it changes the nutritional worth of the corn it affects. Corn smut contains more proteins than the uninfected grains normally do. The amino acid lysine, of which corn contains very little, abounds in corn smut.

See also

• Edible mushroom – Edible fungi fruit bodies
• Medicinal fungi – Fungi that can be used to develop medicationsPages displaying short descriptions of redirect targets

References

• McGee, Harold (2004). On Food and Cooking (revised ed.). Scribner. p. 349 "Huitlacoche, or Corn Smut". ISBN 978-0-684-80001-1.

External links

• U. maydis Genome at the Broad Institute
• MUMDB giving easy access to U. maydis genes

The 'Shaggy ink cap' mushroom Coprinus comatus produces spores by deliquescing (liquefying, or melting) its cap into a black ink. Both the mushroom and the black ink of C. comatus and Coprinopsis atramentaria (the 'Common ink cap') are edible, but adverse effects might be felt if consumed together with alhocol. For this reason, C. atramentaria is also called 'tippler's bane.'

Contemporary arts

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Contemporary artworks involving fungi usually handle or utilize mycelia, yeasts, and other fungal forms rather than mushrooms. The video and light artist Philipp Frank creates so called 'projection mapping' by casting light effcts on mushrooms growing in nature in the 'Funky Funghy' project.

Social games (board games, card games)

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Plant pathology scientist Lisa Vaillancourt at the University of Kentucky developed a 'Fungal Mating Game' based on standard card decks as educational tool for students to better understand the process and concept of fungal mating using as example mating of Saccharomyces cerevisiae (baker's yeast), Neurospora crassa, Ustilago maydis, and Schizophyllum commune. The game can be played both collaboratively and competitively.

Counterculture

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See also: Lysergic acid diethylamide § Counterculture

And you've just had some kind of mushroom
And your mind is moving low
Go ask Alice
I think she'll know

— Jefferson Airplane, White Rabbit

Mushroom-inspired counterculture in the years 1960-70 coincided with the popularity of hallucination-inducing substances present in certain fungi (e.g. Psilocybe species) and the widespread use of LSD in certain social and cultural circles, ensuing policy backlashes, and a general restriction towards the diffusion, popularity, and knowledge about psilocybin and its derivatives. Advocates and pioneers in the research on the effects of LSD and other psycoactive substances (including certain mushrooms) like the Swiss chemist Albert Hofmann advocated for caution and against a ban on such psychoactive substances, arguing in favour of research elucidating the effects and therapeutic potentials of fungi-derived psycoactive substances. Current research on psychoactive mushrooms shows promises for the treatment of mental-health ailments like chronic depression and anxiety.

A 'mushroom counterculture' has been often fuelled by eccentric, unorthodox, and unfalsifiable hyphotheses and interpretations of the influence of (hallucinogenic) mushrooms in culture developments, as in the 'Stoned Ape Theory' by Terence McKenna (arguining that human conscience evolved following consumption of psycoactive mushrooms by our primate ancestors) or as in the book The Sacred Mushroom and the Cross by John Allegro (arguing that Christianity arose as a cult venerating and practising ingestion of psycoactive mushrooms). In his book Food of the Gods, Terence McKenna presents the hypothesis that art itself (along with human language and consciousness) arose from consumption of psychoactive musrooms; McKenna hyphotesis has been controversial, receiving negative appraisal as well as enjoying a cult following.

Several Mushroom festivals around the world, like the Telluride Mushroom Festival in Telluride, Colorado, the Radical Mycology Convergence in Mulino, Oregon, feature and foster a community of practitioners involved with aspects of mushroom foraging, cultivation, cuisine, art, and local culture. Mycological Societies, groups, and associations often organises local event aimed at educating and engaging with the public, like the Transylvania Mushroom Camp in Romania.

Mycelia or hyphae in art

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When a fungal spore lands on fertile ground, it germinates into a hypha (plural: hyphea), a filament of cells exploring and growing into the surrounding substrate. Hypha are metabolic active, often septate (polynucleate) threads of cells, which entangle into dense fungal masses called mycelium (plural: mycelia). Both hyphea and mycelia are referred to as the vegetative parts of the fungus. The fungal vegetative stage involves growth and expansion, usually isodiametrically (that is, in all directions, also referred to as concentric or radial growth), into the soil. Upon favourable conditions, the hyphae can develop into specialized structures, like mushrooms, to complete the fungus reproductive cycle. Further specialized hyphal structures are lamellae (gills below mushroom caps), sporocarps (spore-holding structures), haustoria (root-like structures penetrating deep into a substrate, like in the case of enthomopathogenic fungi infecting ants), sclerotia (hardened myclieum rich in food reserves allowing the fungus to go into dormancy), Hartig net (a network of hyphae in mycorrhizal fungi surrounding symbiotic plant cells), and many more. Fungi which are not known to form mushrooms and are grouped into the Fungi Imperfecti, also called Deuteromycota (from the Latin, literally translated into 'Fungi without womb'). Among them are obligate endophytic fungi which do not form fruiting bodies. Yeasts, moulds, and lichens do not form mushrooms and are historically grouped into the Fungi Imperfecti. The mycelial and hyphal growth phase of fungi is also referred to as the anamorph phase (asexual stage) and, by extension, the hyphae and mycelia are referred to as anamorph structures; the sexual reproductive stage (mushroom) is called teleomorph (from the Latin and Greek, 'teleo' = end, goal, purpose, and 'morph' = form).

A glass sculpture by mycologist William Weston respresenting the phytopathogen Botrytis cinerea (ca. 1940) showing the magnified hyphae and conidiophores of the fungus.

Many fungi do not form hyphae and mycelia, but grow, reproduce, and disperse by other means. These fungi include yeasts (reproducing by budding or fission), the mycobiont of lichens (dispersing by 'propagules'), or moulds (which form hypahe and mycelia but reproduce asexually by forming so-called 'asexual spores' or conidia). By asexual reproduction it is meant the formation of spores or propagation particles without exchange of genetic material between two organisms (although genetic mutations might occur in the process); by sexual reproduction it is meant the transmission and recombination of genetic material between two organisms. In fungal sexual reproduction, there are several mechanisms of genetic exchange; the notion of binary gender does not apply for the sexual reproduction of fungi. Mating can occur in fungi which do not form mushrooms. In both fungi which do form and which do not form mushrooms, hyphae are the stage of fungal mating: 'compatible' hyphae grow towards each other and fuse their plasma membrane to exchange nucleic (genetic) material.

Hyphae are the most metabolically active structures of fungi, secreting high amounts of digestive enzymes in the surrounding environment to consume the growth substratum, as well as bioactive metabolites, including substances used in modern medicine (antibiotic and antimicrobial drugs). The tip of hyphae, called Spitzenkörper (from the German, meaning 'pointed body') is the most active hyphal part, where germination, growth, branching, and secretion take place. The life-saving antibiotic penicillin and its derivatives originate from moulds of the genus Penicillium. Hyphae and mycelia grow by extension and branching, and fungi forming those structures are often referred to as 'filamentous fungi'.

Mycelia and hyphae have seldomly been represented, showcased, transformed, or utilized in the traditional arts due to their invisible, ignored, and overlooked lifestyle and appearance. Depictions of mycelia and hyphae in the graphic arts are very rare. The mycelium of certain fungi, like those of the polypore fungus Fomes fomentarius which is sometimes referred to as Amadou, has been reported though history as biomaterial. More recently, hyphae and mycelia are used as working matter and transformed into contemporary artworks, or used as biomaterial for objects, textiles and constructions. Mycelium is investigated in cuisine as innovative food or as source of meat-alternatives like so-called 'mycoproteins,' enjoying increasing visibility, marketing, commercialization, and endorsement from celebrities. In literature and fiction, hyphae and mycelia are considered (if at all) for their intrinsic properties of decomposition, contamination, and decay. The filamentous, prolific, and fast growth of hyphae and mycelia (like moulds) in suitable conditions and growth media often makes these fungal forms good subject of time-lapse photography. Indirectly, psychoactive substances present in certain fungi have inspired works of art, like in the triptych by Hieronymus Bosch, The Garden of Earthly Delights, with curious and visionally imagery allegedly inspired by ergotism poisoning caused by the sclerotia (hardened mycelium) of the phytopathogenic fungus Claviceps purpurea.

Graphic arts

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Particular from The Temptation of St. Anthony by Matthias Grünewald (1512-1516) showing a sufferer from ergotism, an ailment also called St. Anthony's Fire caused by ergotamine, a fungal toxin present in the plant pathogenic fungus Claviceps purpurea. Because the effect of fungi, rather than fungi itself, influenced the creation of this and similar artworks, this can be considered an indirect influence of fungi in the arts.

See also: Temptation of Saint Anthony in visual arts

The German Renaissance painter Matthias Grünewald (c. 1470–1528) depicted in The Temptation of St. Anthony (1512-1516) a sufferer from ergotism, also referred to as St. Anthony's Fire. Ergotism is caused by ingestion of sclerotia (hardened mycelium) of Claviceps purpurea, a fungal endophyte infecting rye and other plants. Ergotism is cause by consumption of rye and other food contaminated with the sclerotia of the fungus, as for example in flour. Bread from contaminated flour looks black due to the sclerotia. The mycelium contains the fungal alkaloid ergotamine, a potent neurotoxin which can cause convulsions, cramps, gangrene of the extremities, hallucinations, and further adverse and potentially lethal effects depending on dosage. Ergotamine is a precursor molecule in the synthesis of psychedelic drug lysergic acid diethylamide (LSD).

Literature

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See also: List of books about mushrooms and Category:Fictional fungi

Whereas non-fiction books about fungi often (if not always) include hyphae and mycelia, examples of hyphae and mycelia in literary fiction are much rarer in comparison to mushrooms and spores. When these fungal forms are included in work of fiction, they are often associated with elements of rot and decay.

Cinema, TV shows, and motion pictures

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Due to their fast, radial growth (also called isodiametric growth, that is, with same speed and size in all directions), as well as their biodegradation potential, mycelia and hyphae are often used as time-lapse photography to present filamentous growth and/or decay. The short movie Wrought (2022) by Joel Penner and Anna Sigrithur is a series of time-lapses exploring rot, fermentation and decay displaying moulds, yeasts, mushrooms, and further decomposers, with voice-over to describe the processes. The movie was awarded 'Best of the Fest' and 'Audience Choice' awards at the 2022 Fungi Film Festival.

Music

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Examples of hypha and mycelium influence in music are scarce. In her mushroom-inspired album Fossora (2022), Icelandic avant-guard musician Björk included tracks such as Mycelia and Fungal City.

Fungi might have a direct influence on music instruments. The process of wood spalting is often used as aestetic element, e.g. in the manufacturing of guitar bodies. The luthier Rachel Rosenkrantz experiments with fungi (mycelium) to create 'Mycocast,' a guitar body made of fungal biomass due to the acoustic properties of mycelium and its growth plasticity (e.g. the ability to take virtually any shape upon being casted in a desired form). Violins from wood infiltrated by mycelia of the fungus Xylaria polymorpha (vernacularly called 'dead man's fingers') produce sounds close to those from the Stradivarius violin. Researchers are investigating the use of fungi to the species Physisporinus vitreus and Xylaria longipes in controlled wood decay experiments to create wood with superior qualitites for music instruments.

In some cases, music generation using fungi is conceptual, as in Psychotropic house (2015) and Mycomorph lab (2016) of the Zooetic Pavillion by the Urbonas Studio based in Vilnus (Lithuania) and Cambridge (Massachusetts), in which a mycelial structure is designed to act as amplifier for sounds from nature mixd into loops.

Architecture, sculptures, and mycelium-based biomaterials

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See also: Mycelium-based materials

File:Figure 6 in Almpani-Lekka et al (2021), Fungal Biology and Biotechnology, A review on architecture with fungal biomaterials the desired and the feasible.jpg

The “Growing Pavilion” made (among others) with fungal mycelia exhibited at the 2019 Dutch Design Week in Eindhoven (The Netherlands). Left: external view; right: interior view. From Almpani-Lekka et al. (2021).

Mycelium is being investigated and developed by researchers and companies into a sustainable packaging solution as alternative to polystyrene. Mycelium as working matter in sculptures is attracting interest from artists working in the contemporary arts.

Early experimentations of artist with mycelia have been exhibited at the New York Museum of Modern Art. Experimentations with fungi as components – and not only as contaminant or degraders of buildings – have started around 1950. Current collaborations between scientists, artists, and society at large are investigating and developing mycelium-based structures as building materials. Use of fungi from the genera Ganoderma, Fomes, Trametes, Pycnoporus, or Perenniporia (and more) in architecture include applications such as concrete replacement, 3D printing, soundproof elements, insulation, biofiltration, and self-sustaining, self-repairing structures.

Beside the study of fungi for their beneficial application in architecture, risk assessments investigate the potential risk fungi can pose with regard to human and environmental health, including pathogenicity, mycotoxin production, insect attraction through volatile compounds, or invasiveness.

Fashion, design, and mycelium-based textiles

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Historically, ritual masks made of lingzhi mushroom (species from the genera Ganoderma) have been reported in Nepal and indigenous cultures in British Columbia. Currently, fungal mycelia are molded, or rather grown, into sculptures and bio-based materials for product design, including into everyday objects, to raise awareness about circular economy and the impact that petrol-based plastics have on the environment.

Artificial leather made from fungal mycelium. (A) Reishi™ from Mycoworks; (B) Mylea™ from Mycotech Lab; (C) Mylo™ from Bolt threads; (D) mycelium leather from VTT Technical Research Centre (Finland). From Vanderlook et al. (2021).

Biotechnology companies like Ecovative Design, MycoWorks, and others are developing mycelium-based materials which can be used in the textile industry. Luxury fashion brands like Adidas, Stella McCartney, and Hermès are introducing vegan alternatives to leather made from mycelium.

The tinder polypore Fomes fomentarius (materials derived from which are referred to as 'Amadou') has been used by ancestral cultures and civilizations due to its flammable, fibrous, and insect-repellent properties. Amadou was a precious resource to ancient people, allowing them to start a fire by catching sparks from flint struck against iron pyrites. Bits of fungus preserved in peat have been discovered at the Mesolithic site of Star Carr in the UK, modified presumably for this purpose. Remarkable evidence for its utility is provided by the discovery of the 5,000-year-old remains of "Ötzi the Iceman", who carried it on a cross-alpine excursion before his death and subsequent ice-entombment. Amadou has great water-absorbing abilities. It is used in fly fishing for drying out dry flies that have become wet. Another use is for forming a felt-like fabric used in the making of hats and other items. It can be used as a kind of artificial leather. Mycologist Paul Stamets famously wears a hat made of amadou.

Fungi have been used a biomaterial since many centuries, for example as fungus-based textiles. An early example of such "mycotextiles" comes from the early 20th century: a wall pocket originating from the Tlingit, an Indigenous Population from the Pacific Northwest (US) and displayed as historical artefact at the Dartmouth College's Hood Museum of Art, turned out to be made of mycelium from the tree-decaying agarikon fungus. Fungal mycelia are used as leather-like material (also known as pleather, artificial leather, or synthetic leather), including for high-end fashion design products.

Beside their use in clothing, fungus-based biomaterials are used in packaging and construction. There are several advantages and potentials of using fungus-based materials rather than commonly used ones. These include the smaller environmental impact compared with the use of animal products; vertical farming, able to decrease land use; the thread-like growth of mycelium, able to be molded into desirable shapes; use of growth substrate derived from agricultural wastes and the recycling of mycelium within the principles of circular economy; and mycelium as self-repairing structures.

Culinary arts

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See also: Fermentation in food processing

Mushrooms are traditionally the main form of fungi used for direct consumption in the culinary arts. The fermentative abilities of mould and yeasts have a direct influence in an enormous variety of food products, including beverages such as beer, wine, sake, kombucha, coffee, soy sauce, tofu, cheese, or chocolate just to name a few.

Recently, mycelium is increasingly being investigated as innovative food source. The Michelin-star restaurant The Alchemist in Copenhagen (Danemark) experiments with mycelium of fungi such as Aspergillus oryzae, Pletorus (oyster mushroom), and Brettanomyces with funding from the Good Food Institute, to create novel fungus-based dishes, including the creation of 'mycelium-based seafood' and the consumption of raw, fresh mycelium grown on a Petri dish with a nutrient-rich broth.

The US-based company Ecovative is creating fungus-based food as meat alternative, including 'mycelium-based bacon.' The US-based company Nature's Fynd is developing various kinds of food products, including meatless patties and cream cheese substitutes, using the so-called 'Fy' protein from Fusarium.

Contemporary arts

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"At this point, I stepped back and let the sculpture sculpt itself."

— Xiaojing Yan, Mythical Mushrooms: Hybrid Perspectives on Transcendental Matters

Hapha and mycelium gets increased attention in the contemporary art due to its growth and plasticity, and is occasionally the starting point for artworks in the contemporary art exploring the biological properties of degradation, decomposition, budding ('mushrooming'), and sporulation (spore formation and release).

An early form of BioArt is Agar art, where various microorganisms (including fungi) are grown on agar plates into desired shapes and colours. Thus, the agar substrate becomes a canvas for microbes, which are an analogue to the artist's colour repertoire (palette). In agar art, fungi (and other microorganisms, mostly bacteria) assume different appearances based on intrinsic characteristics of the fungus (species, morphology, fungal form, pigmentation), as well as external parameters (like inoculation technique, incubation time or temperature, nutrient growth medium, etc.). Microorganism can also be engineered to produce colour or effect which are not intrinsic to them or are not present in nature (e.g., they are mutant from the wild type), like for example bioluminescence. The American Society for Microbiology (ASM) holds an annual 'Agar Art Contest' which attracts considerable attention and elaborate agar artworks. An early 'agar artist' was physician, bacteriologist and Nobel Prize winner Alexander Fleming (1881-1955).

The Folk Stone Power Plant (2017), like the Mushroom Power Plant (2019), by Lithuanian artist duo Urbonas Studio, are physical installations based on 'mycoglomerates,' that is an interpretation and representations of vaguely-described microbial symbioses aimed at energy production alternative to fossil fuel. The Folk Stone Power Plant is a 'semi-fictional' alternative battery installed in Folkestone (UK) during the Folkestone Triennale, aiming at a reflection about symbioses (both in nature and between artists and scientists) and about unconventional power sources. The design is based on drawings from polymath and naturalist Alexander von Humboldt (1769-1859), while the microbial power source, hidden within the stone, mirrors the largely unnoticed, yet crucial, contribution of mycelial networks (that is, mycorrhiza) in ecology.

The Living Color Database (LCDB), freely available at www.color.bio, links organisms across the tree of life (in particular fungi, bacteria, and archaea) with their natural pigments, the molecules' chemistry, biosynthesis, and colour index data (HEX, RGB, and Pantone), and the corresponding scientific literature. From Sharma & Meyer (2022).

In her work Linghzi Girl (2020) by Chinese-Canadian artist Xiaojing Yan, female bust statues casted with the mycelium of lingzhi fungus (Ganoderma lingzhi) are exhibited and left to germinate. From the mycelium-based sculptures sprout mushrooms, eventually spreading, once ripe, a cocoa-powder dust of spores on the bust, after which the sculptures are preserved by desiccation to stop the fungal cycle and maintain the artwork. Artist Xiaojing Yan thus explains the audience's reaction to her work:

"The uncanny appearance of these busts seems frightening for many viewers. But a Chinese viewer would recognize the lingzhi and immediately become delighted by the discovery."

During an artist-in-residence project The colors of life (2021) at the Techische Universität Berlin (Germany), artist Sunanda Sharma focus on the biotechnology-relevant fungus Aspergillus niger, and visualises its black pigmentation through fungal melanin by means of video, photo, animation, and time-lapse footage. Within the same residence, the artist created an open source database The Living Color Database (LCDB), which is an online compendium of biological colors for scientists, artists, and designers. The Living Color Database, freely available at www.color.bio, links organisms across the tree of life (in particular fungi, bacteria, and archaea) with their natural pigments, the molecules' chemistry, biosynthesis, and colour index data (HEX, RGB, and Pantone), and the corresponding scientific literature. The Living Color Database comprises 445 entries from 110 unique pigments and 380 microbial species.

Spores in art

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Fungal spores are the equivalent of seeds in plants. As the reproduction and dispersal unit of many fungi, spores guarantee a wide dispersal in nature, as well as genetic variability, increasing individual fitness in nature. Mating by sexual reproduction in fungi results in the formation of mushroom (also called fruiting bodies), which are the structures bearing spores. Many fungi can reproduce and form spores asexually, as in the formation of asci (singular: ascus) in Ascomycetes or of conidiophores in moulds. Many fungi do not form spores but reproduce by budding like yeasts; other fungi forms so-called 'vegetative spores,' which are specialized cells able to withstand unfavorable growth conditions, as in black yeasts. Lichens do not reproduce and disperse by sporulation, but by fragments called 'propagules.'

Examples of fungal spores in the arts are rare due to their invisibility and difficulties to treat and manipulate as working matter. Notable exceptions are so called 'spore prints,' or glass sculptures by mycologist William Dillon Weston (1899-1953) representing magnified microfungi and spores (ascospores, basidiospores). Often, fungal spores are employed as an agent of infection and decay in literature and the graphic arts, whereas recently they are increasingly used in the contemporary art in a positive or neutral way to reflect about processes of transformation, interaction, decay, circular economy, and sustainability.

Graphic arts

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See also: Spore print

So called 'spore prints' are created by pressing the underside of a mushroom to a flat, white or coloured surface, to allow the spores to be imprinted on the sheet. Since some mushrooms can be recognized based on the colour of their spores, spores prints are a diagnostic tool as well as an illustrative technique. Several artists used and modified the technique of spore printing for artistic purposes. Mycologist Sam Ristich exhibited several of his spore prints in an art gallery in Maine around 2005–2008. The North American Mycological Association (NAMA) created a 'how-to guide' for people interested in creating their own spore prints.

The artwork Auspicious Omen – Lingzhi Spore Painting by Chinese-Canadian artist Xiaojing Yan creates abstract compositions resembling traditional Chinese landscapes by fixing spores of the linghzi fungus with acrylic reagents. The linghzi mycelial sculptures by Xiaojing Yan, including Linghzi Girl (2020) and Far From Where You Divined (2017) are allowed to germinate into mushrooms during exhibition, creating a dust of spores raining down on the female busts, children, deers, and rabbits. The artworks are then desiccated for preservation, stopping the fungal growth and the metamorphosis of the sculptures. Artworks as such, including growth of the fungus, an incontrollable transformation of the art object, and several forms in the fungal life cycle, are rare.

Literature

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See also: Category:Fictional fungi

Whereas non-fictional books about fungi cover spores in the context of fungal spore formation, dispersal, harvesting, or germination, works of literary fiction involving spores are generally linked to infection and decay, and thus have mostly a negative connotation. In stories where mushrooms are perceived or represented as threat, spores fulfill the same role. In the short story Come into My Cellar, by Ray Bradbury, for example, spores are depicted as an alien invasion.

Comic books and video games

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The critically acclaimed and commercially successful video game franchise The Last of Us by Sony Computer Entertainment (Part I, released in 2013; downloadable content adds-on The Last of Us: Left Behind, released in 2014; Part II, released in 2020) is a post-apocalyptic, third-person action-adventure game set in North America in the near future, after a mutant fungus decimates humanity. The 'fungal apocalypse' is inspired by the effect ant-pathogenic fungi like Ophiocordyceps unilateralis have on their insect preys. The fictional fungus infects humans by means of spore inhalation, after which the persons turn into zombie-like creatures progressing into incremental stages of infection to fungus-like humans. The infected persons develop cannibalism and can transmit the fungal infection to other humans by biting. In advanced stages of infection, so-called clickers have their body transfigured by the fungus, with polypore-like bulging on their heads and faces; having lost the ability to see, they locate their preys by echolocation. An important part of the plot of The Last of Us game franchise revolves around vaccines against the fungal disease; as opposed to vaccination against viral and bacterial pathogens, research on vaccines for human fungal diseases lags behind, with currently no vaccine available against human fungal pathogens. The Last of Us Part II has been awarded best video game of 2020 by The Game Awards. A television adaptation by HBO starring among others Pedro Pascal as Joel, Bella Ramsey as Ellie, and Nick Offerman as Bill, is due in January 2023.

The comic strip by Dave Gibbon Come into My Cellar is based on Ray Bradbury's short story with the same name, where fungal spores are an alien entity taking over humanity by mind control, especially of children obsessed with growing mushrooms in their home basement. An adaptation into Italian appeared for the famous comic series Corto Maltese in 1992 with the name Vieni nella mia cantina.

Yeasts, moulds, or lichens in art

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Many fungi do not reproduce and disperse by spores. Instead, they live single-celled and reproduce by budding or fission as in yeasts, or live in a symbiosis with an algal or cyanobacterial partner as in lichens, which are obligate symbiotic organisms (that is, they cannot survive apart), growing very slowly, and dispersing by fragmentation into so-called 'propagules.' Moulds do form spores ('asexual spores') but no mushrooms, and grow into filaments (hyphae and mycelia) which thrive in moist environments and spoil food. Despite being unicellular, yeasts can reproduce sexually by mating. Yeast cells which are haploid (that is, having only one set of chromosomes) and belong to different 'mating types,' so-called a and α (alpha) mating types, recognize each other by means of pheromone signalling to fuse and exchange genetic material. The yeast Saccharomyces cerevisiae (from the Latin, translated into 'sugar-fungus of beer'; also called brewer's yeast or baker's yeast) is among the most well-studied organisms among others due to its fast growth, unicellularity, ability to reproduce both sexually and asexually (genetic tractability) and is considered a model organism in research and the life sciences. S. cervisiae is the first eukaryotic organism which genome (the entirety of its DNA or genetic material) was ever sequenced. Alcoholic fermentation producing beer, wine, and spirit is mostly in the hands of the yeast Saccharomyces cerevisiae. Also called also baker's yeast, S. cerevisiae is the leavening agent in bread. Despite growing mostly in a unicellular (single-celled) fashion, upon certain external stimuli like nutrient limitation or starvation (in particular toward carbon and/or nitrogen sources), yeasts can assume a filamentous growth forming chains of elongated cells. The morphological switch from unicellular to filamentous growth is crucial for pathogenic yeasts, like for example in the opportunistic fungus (that is, present as commensal in the human gut microbiota and only become pathogenic under certain conditions, like a weakened host immune system) Candida albicans responsible for vaginal infection. Moulds, like those which spoils food, are major natural producers of antibiotics, like penicillin. Industrial production of chemicals like citric acid, or enzymes like lipases, is also largely in the hands of filamentous fungi and moulds like Aspergillus niger. Blue cheese is cheese ripened by the Penicillium roqueforti and other filamentous fungi.

Lichens, a symbiosis between fungi and algae or bacteria, illustrated by German zoologist, naturalist, phylosopher, and illustrator Ernst Haeckel in Kunstformen der Natur (1904). Naturalists illustrating their observations often created remarkable work of arts.

Yeasts, moulds, and lichens did not enter into the arts very often. Despite being distributed worldwide and extremely common (e.g. being responsible for fermentation and leavening in the production of wine, beer, bread, and fermented products like coffee, soy sauce, tempeh, and many more), their direct influence in the arts remains modest. Indirectly, yeasts have had a conspicuous influence in the art, as fermentation has contributed enormously to different cultures around the globe and across time; in La traviata (1853) by Italian opera composer Giuseppe Verdi, for example, one of the best-known opera melodies is 'Libiamo ne' lieti calici' (in English, translated into "Let's drink from the joyful cups"), which is but one of numerous brindisi (toast) hymn. Other testimonies of the indirect effect of yeasts in the arts are the numerous deities and myths are associated with wine and beer. The field of ethnomycology focuses more on the influence of psychoactive fungi on human culture rather than on aspects such as medicine, food production practices, or cultural influence in the arts. Yeasts and moulds are often an agent of decay and contamination in the arts, whereas recently they are increasingly used in the contemporary art in a positive or neutral way to reflect about processes of transformation, interaction, decay, circular economy, and sustainability. Time-lapses photography is a tool often deployed by artists to accelerate and depict fungal growth in the arts, in particular growth of filamentous fungi (mould, hypahae or mycelia). Aside from various illustrations, lichens are very seldomly represented in the arts to their slow growth as well as their frailty towards maniputation.

Notable examples of yeasts, moulds or lichens in the arts include:

• Ernst Häckel illustrations of lichens in Kunstformen der Natur (1904)
• Chemical compounds from some lichens are used as dying substances (this is also true for compounds derived from mushrooms)
• In the science fiction novel Trouble with Lichen (1960) by John Wyndham, a chemical extract from a lichen is able to slow down the aging process, with a profound influence on society
• In Stephen King's horror short story Gray Matter (1973), a recluse man living with his son drinks a 'foul beer' and slowly transforms into an inhuman blob-like abomination that craves warm beer and shun light, and transmutes into a fungus-like fictional creature
• The short movie Who's Who in Mycology (2016) by Marie Dvoráková, a comedy which won numerous awards at international film festivals, involves 'a young trombone player [...] trying to open an impossible bottle of wine [...] and some mold gets in his way
• The novel Lichenwald (2019) by Ellen King Rice, author of 'Mushroom Thrillers' is a crime story involving lichens, dementia, and manipulations
• The Dutch textile artist Lizan Freijsen created the Fungal Wall for the microbe museum Micropia, together with TextielMuseum Tilburg, a wall-sized tapestry by tufting resembling mould growth
• In so-called 'mold paintings,' surfaces of buildings or sculptures are intentionally overgrown with moulds to create visually appealing effects
• The contemporary artist Kathleen Ryan creates oversized, composite sculptures of rotting fruits, like in the Bad Fruit series
• The short movie Wrought (2022) by Joel Penner and Anna Sigrithur is a series of time-lapses exploring rot, fermentation and decay displaying moulds, yeasts, mushrooms, and further decomposers,

Performative arts (theatre, comedy, dance, performance art)

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The musical theatre show The Mould That Changed the World is a show running both in the US (in Washington, D.C. and Atlanta, Georgia) and the UK (in Edinburgh and Glasgow, Scotland) which centers around the life and legacy of Alexander Fleming, the Scottish discoverer of the antibiotic penicillin and 1945 Nobel Prize winner in Physiology or Medicine. Alexander Fleming discovered in 1928 during his work as bacteriologist that bacteria growing on a Petri dish were inhibited by a mould contamination, namely from a fungus of the genus Penicillium, from which the antibiotic name 'penicillin' derives. The story involves jumps in time to highlight the legacy of the discovery of antibiotics and is partly set during the Great War, when Alexander Fleming served as private, as well as the personification of some characters (e.g. Mother Earth). The musical has been developed for educational purposes to raise awareness against the tremendous, worldwide threat that the rise of antimicrobial resistance poses. The musical provides freely available teaching resources and has been developed with the participation of the British Society for Antimicrobial Chemotherapy (BSAC). The musical choir is composed of both professional singers and actors as well as health care professionals, lab technicians, and scientists, and is an example of an artistic project merging science and the arts.

The dance contest for scientists called 'Dance your Ph.D.' sponsored by the American Association for the Advancement of Science (AAAS) is an annual competition established in 2008 encouraging communication and education of complex scientific topics through interpretative dance. All scientific fields and area of research are covered (biology, chemistry, physics, and social science) and several contestant entries involved fungi, including some winners. The 2014 winner was plant pathologist and aerial acrobat Uma Nagendra from University of Georgia (Athens) with Plant-Soil Feedbacks After Severe Tornado Damage, a trapeze-circus dance representing the effect of extreme environmental events (like tornadoes) on tree seedlings and the positive effect those events can have with regard to withstanding phytopathogenic fungi. The 2022 winner was Lithuanian scientist Povilas Šimonis from Vilnius University with Electroporation of Yeast Cells, a dance illustrating the effect of electroporation (a method involving pulses of electricity to deactivate cells, or make them more porous and prone to acquire extracellular DNA, a crucial step in genetic engineering) on yeasts.

Contemporary arts

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In the contemporary arts, works involving fungi are often interactive and/or performative and tend to transform and utilize fungi rather than merely represent and showcase them. In her work Myconnect (2013), bioartist Saša Spačal invites the audience to interact with the artwork, involving Shiitake (Lentinula edodes) or Oyster mushrooms (from the genus Pleurotus), which takes the form of a capsule connecting the human with the fungus on a sensory level. Bioartists use yeasts to provoke a reflection on genetic engineering. Slovenian intermedia artist Maja Smrekar’s created yoghurt using a genetically-modified yeast with a gene from the artist herself in Maya Yoghurt (2012).

The blogger and feminist Zoe Stavri baked in 2015 sourdough bread using yeast she isolated from her own vaginal yeast infection using a Dildo, which she then mixed with flour and water and let leaven, and finally ate. The activity, which she documented both on her blog posts and on social media, tagging it with the hashtag #cuntsourdough, caused a lot of discussion on social media, including repulsion, hate messages, and food safety concerns, as the practice did not involve axenic isolation of the leavening yeast; however, during baking, microorganisms present in dough are most probably heat deactivated and thus harmless. As the activist herself noted: "people have been making and eating sourdough [with wild yeasts] for millennia." People had experimented before with microorganisms from the vaginal microbiota to create food and incite a reflection on topic of food fermentation and female bodily autonomy and self-determination. In 2015, the Medical Doctor/PhD student Cecilia Westbrook fermented milk into yogurt using as starter culture from her own vaginal secretion, which she then ate. As opposed to Zoe Stavri's vaginal sourdough bread, the homemade yogurt relied on the fermentation properties of lactic acid baceria (e.g. lactobacilli), rather than yeasts (fungi); and, unlike bread, yogurt is a culture of living microorganisms. The praxis is thus considered a food hazard by the US Food and Drug Administration.

The exhibition Fermenting Futures (2022) by bioartists Alex May and Anna Dumitriu in collaboration with the University of Natural Resources and Life Sciences (BOKU) is an artwork which wants to make the audience reflect about the role of yeast biotechnology to confront global issues of contemporary society by culturing and showcasing fermentation flasks of Pichia pastoris for the bioconversion of carbon dioxide into the biodegradable plastics. The artwork The Bioarchaeology of Yeast recreates by moulding the biodeterioration marks left by certain yeasts, like black yeasts, on work of art and sculptures, and displays them as aestic objects, reflecting on the process of erosion; the installation Culture used CRISPR technology to confer to a non-fermenting strain of Pichia pastoris the ability to ferment and work as leavening agent as the baker's yeast.

In 'Fungal Dot Painting,' fungal conidia are inoculated into agar droplets, deposited on a black (acrylic glass) surface, and incubated to allow fungal growth. The results resemble pointilism. From Grunwald et al. (2021).

In 'Etched Fungal Art,' an acrylic glass surface modified by etching (lathing or printmaking) is poured over with a suspension of fungal conidia in an agar-based substrate, and then incubated to permit fungal growth into the etched channels. From Grunwald et al. (2021).

A team of artists and researchers developed novel art techniques using the model (that is, widely studied in laboratory research) mould Aspergillus nidulans. The artist-scientist team described the development of two new techniques: 'Fungal Dot Painting' and 'Etched Fungal Art.' In Fungal Dot Painting, akin to pointilism where small dots unite to compose an image, fungal conidia are inoculated into agar droplets which are then deposited on a dark surface of black acrylic glass for contrast, and incubated at the desired condition to allow fungal growth. In 'Etched Fungal Art', an acrylic glass surface modified by etching (lathing or printmaking) is poured over with a suspension of fungal conidia in an agar-based substrate, and then incubated to permit fungal growth into the etched channels. Both artforms allow for temporal dynamism, insofar being composed of living fungal organisms they change and evolve over time.

Other (non-fungal) examples: the slime mould Physarum polycephalum

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Physarum polycephalum is a slime mould (myxomycete) and not a fungus; however, it grows in a similar fashion as filamentous fungi and it is increasingly used by bioartists. Due to its complex problem-solving abilities, the slime mould is used to mimic or investigate human behaviours.

Physarum polycephalum has been shown to exhibit characteristics similar to those seen in single-celled creatures and eusocial insects. For example, a team of Japanese and Hungarian researchers have shown P. polycephalum can solve the shortest path problem. When grown in a maze with oatmeal at two spots, P. polycephalum retracts from everywhere in the maze, except the shortest route connecting the two food sources.

When presented with more than two food sources, P. polycephalum apparently solves a more complicated transportation problem. With more than two sources, the amoeba also produces efficient networks. In a 2010 paper, oatflakes were dispersed to represent Tokyo and 36 surrounding towns. P. polycephalum created a network similar to the existing train system, and "with comparable efficiency, fault tolerance, and cost". Similar results have been shown based on road networks in the United Kingdom and the Iberian peninsula (i.e., Spain and Portugal).

Indirect influence of fungi in the arts

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Whereas most artforms involving fungi are directly influenced by them – that is, they tend to directly represent (describe), showcase, transform, or utilize fungi – there are numerous examples where fungi have (had) an indirect influence in the arts. This indirect influence of fungi in the arts can be broadly classified into three categories:

• The description of fungal effects (for example, in the numerous representation of the Temptation of Saint Anthony in visual arts)
• The creation of art under the effects of fungi (for example, in literary works partially influenced by the use of alcohol, as those from authors like Ernest Hemingway, Truman Capote, and Charles Bukowski)
• artworks where the role of fungi is unbeknown to the artist (e.g. in mythopoiesis related to alcoholic fermentation, resulting in numerous deities of wine and beer, before an understanding of the process)

Of notable example are the works of Early Netherlandish painter Hieronimus Bosch (ca. 1450–1516) and of transgressive fiction writer Charles Bukowski (1920-1994) insofar part of their artworks have been likely created under the influence of fungal substances while they also depict the effect of fungal metabolites.

Preservation of artworks against fungal decay

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Fungi and other microorganisms represent a treat to historical and cultural heritage, as they can colonize artworks (paintings, papers, sculptures, buildings, etc.) and damage them by means of mechanical, chemical, or aesthetic damage. Researchers, art historians, and art conservationists are developing techniques and approaches to minimise the deleterious microbial effect of fungi (and other microorganisms) on artworks.

A recent study investigated the surface of a Brazilian contemporary painting by Teresinha Soares and found fungal (mould) species such as Aspergillus flavus, Aspergillus niger and Penicillium citrinum. The fungi showed cellulolytic activity in vitro, as well as tropism towards certain pigments present on the painting, indicating specificity. In 2022, the ubiquitous, newly discovered yeast species Blastobotrys davincii was isolated from a worldwide survey of house dust. The yeast reproduces asexually, is xerophilic (tolerant to desiccation), and has been found also the surface of Leonardo da Vinci's Portrait of a Man in Red Chalk (self-portrait, c. 1512), from which the species name is derived.

Microorganisms like fungi are not only considered in the preservation of artworks do due their decaying and contaminating properties. Some microorganisms can contribute to preservation of artworks due to their biocontrol properties. Some contemporary artworks explore the topic to raise awareness and develop techniques and tools for bio-conservation of artworks, like the work Unruly objects (2021) by Anna Dimitriu et al.

An area of applied research focuses on limiting the growth, harm, and health hazard of mould growing inside buildings, often referred to as 'microbiology of the built environment.'

Further explorations, applications, and fostering of the 'fungal arts'

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Sustainability approaches

"Pilze sind Meister der Zersetzung aber auch Meister der Synthese."

"Fungi are masters of decomposition as well as masters of synthesis."

— Vera Meyer, Pilze zwischen Wissenschaft und Kunst (Fungi between science and art), BIOTOPIA, 2021

Several artistic explorations of fungi have as background, intention or goal the development of sustainable solutions to current environmental issues, or aim at raising awareness on these topics. These endeavors often involve a multi-disciplinary approach between artists and fungal practitioners, and transform or utilize fungi for the desired goal. Occasionally, a commercial outcome beyond the purely artistic approach or experimentation is striven for or achieved. Given the potential of fungi to both degrade biological wastes and to grow and form new biomass, these approaches fall often within the realm of circular economy. Patents to intellectually protect the technological developments are often filed.

'MY-CO SPACE' is a habitable prototype of a building made of mycelium. Mycelium of the tinder fungus Fomes fomentarius is grown in hexagonal frames with a plant-based substrate, desiccated, and assembled into the habitable structure. From the 'Engage with Fungi' project in Meyer and Pfeiffer (2022). Pictures from Wolfgang Günzel.

Examples of the use of fungi in sustainability approaches fall within production of fungus-based materials for personal use (vegan leather, house furniture) or as construction materials, or for alternative burial practices (bioremediation), just to name a few.

Fungus-derived material from mycelium are being developed to create artificial leather for high-end fashion products and hold promises to be a sustainable alternative to animal-derived leather.

The exhibition Fungal Futures (2016) by Italian designer Maurizio Montalti on display at the microbe museum Micropia explores the use of mycelium and its growth properties for sustainable production of mass consumer goods and everyday products.

South Korean artist Jae Rhim Lee developed with the Infinity Burial project (2011) a so-called 'Mushroom burial suit' infused with fungal spores to raise awareness on an promote an environmental friendly burial practice, as upon death the body is prepared with environmental pollutants for the sake of short-term preservation prior to burial or cremation. The same concept is developed by the Dutch start-up Loop based at TU Delft with the 'Living coffin.'

The habitable sculpture 'MY-CO SPACE' is a structure made of fungi and wood designed for and showed at the open-air exhibition tinyBE: living in a sculpture (2021), together with other seven habitable sculptures by international artists. Mycelium of the tinder fungus Fomes fomentarius are inoculated on hemp shives and cultivated in hexagonally shaped frames made of plywood. After about six weeks of growth, the fungus overgrows the plant biomass and connects the loose substrate particles, after which the fungus is inactivate by desiccation at 60 °C for two days. The panels are then assembled into the habitable sculpture made of fungal mycelium, called 'MY-CO SPACE.'

Citizen science, educational projects, resources

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More and more artists work with fungi in inter- and transdisciplinary projects to raise awareness about the potential of fungi in educational and citizen science (participative) projects, e.g. in Building Fungitopia (2022) by Isabel Correia or Fungitopia and the Nature of Growing (2017) by Marjan Groot. Numerous artworks, collectives, or collaborative projects aims at communicating the importance of fungi to the broader public, as well as at encouraging an empathic relationship with fungi.

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  'Walk & Talk' within the project 'Mind the Fungi' (2018) in Berlin Tegel (Germany). From Regine Rapp in '"On Mycohuman Performances: Fungi in current artistic research."

  The 'Mind the Fungi' project Walk & Talk is a series of open excursions to educate about fungi in nature
• The Fungi Foundation organizes a series of programs and hosts resources on its website with the goal of educate and organise expeditions and excursions
• The North American Mycological Association (NAMA) hosts The Fungus Files aimed at educating children at elementary-school level
• The Mycological Society of America (MSA) is a professional society of mycologists which hosts a series of education resources on its website
• The iNaturalist website is often used for sharing biodiversity observations on fungi and mushrooms (among other living beings)
• The Society for the Protection of Underground Networks (SPUN) is a scientific research organisation collaborating with local communities with the mission of protecting and harnessing underground mykorrhizal networks
• The New York Mycological Society (NYMS) organises a yearly Fungus Fest in New York and regular 'walks'

Fungi in traditional, oriental, or alternative medicine

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The Chinese poet Tao Yuanming (365-427) holding a lingzhi mushroom as depicted by Chen Hongshou (1598–1652)

Before the advent of modern medicine, fungi in their many forms, including their metabolic products and their properties, have been used for centuries in health and medicine. Alleged or inferred benefits of fungi in traditional, oriental, or alternative medicine include anesthesia, extension of life expectancy, antimicrobiosis, treatment of birth complicacies, and more. Beverages containing alcohol (the product of metabolism of yeasts like Saccharomyces cerevisiae) have been used throughout history as sedative or analgesic before the advent of modern aneshesia. The use of certain moulds and their antimicrobial properties has been documented throughout history before Alexander Fleming's discovery of penicillin and its subsequent industrial production. Documents show 'mould cures' for wounds and other ailments in cultures and civilization as different as the Ancient Egypt, indigenous people of Australia, the Jewish Talmud, and XVII century England.

 

The polypore Ganoderma lucidum (also referred to as reishi or mannetake in Japan, and as lingzhi in China) is considered a 'Mushroom of Immortality' in certain oriental cultures, although its beneficial medical properties have been disputed. The fungus has been depicted several times in the arts, as in the painting of Tao Yuanming holding a lingzhi mushroom by Chen Hongshou (1598-1652), a Chinese painter of the late Ming dynasty. Commercial production and distribution of lingzhi mushrooms is now a multi-million dollar business.

Concoctions from the fungus Claviceps purpurea were used in the Middle Ages to treat birth complicacies. The discovery of LSD by Swiss Chemist Albert Hofmann, a chemical similar to the psychedelic compound psilocybin naturally occurring in Psilocybe mushroom, originates from research on the derivates from ergot fungi and the knowledge of its historical use to treat birth complicacies.

Exhibitions on fungi

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Several exhibitions and musea increasingly dedicate space for fungi in the arts. Several relevant examples are below:

• 'The Art of Mushroom' in Serralves (Porto, Portugal); from June 2022 to June 2023
• 'El Museo del Hongo' (The Fungus Museum) in Santiago (Chile); permanent exhibition
• 'Nature’s Mysterious Networks: Mushrooms, Mycelia and Yeasts' at Groundwork Gallery in Norfolk (UK); from October to December 2021
• 'MUSHROOMS: The Art, Design and Future of Fungi' at Somerset House in London (UK); from July to September 2020; featured in Google Arts & Culture
• Micropia Museum in Amsterdam (The Netherlands), featuring several permanent exhibitions on fungi
• Beatrix Pottery Gallery, featuring among other her mushroom illustrations; permanent exhibition

Patrons of the 'fungal arts' and interest groups

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A series of nonprofit organizations, groups, societies, and circles encourage and support the 'fungal arts' and enhance the visibility of fungi within society in various ways:

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  The 'Fauna Flora Funga' initiative aims at increasing awareness and conservation efforts of fungi, alongside animals ('fauna')) and plants ('flora').

  The North American Mycological Association (NAMA) organises regular Visual Art Contests and Photography Contests
• The Fungi Foundation is the first non-governmental organisation dedicated to fungi and hosts a series of events, resources, and material on their website
• The microorganism museum Micropia curates a list of educational resources on fungi, fungi-inspired artworks, and fungi-themed exhibitions
• The 'Fauna Flora Funga' project is a global initiative aimed at increasing awareness of fungi, alongside animals ('fauna') and plants ('flora'), in particular regarding conservation efforts, and encourages signatories to endorse their statement. Signatories include Jane Goodall, Michael Pollan, Paul Stamets, Philipp Ball, Alan Rayner and many more.
• The British Mycological Society established a UK Fungus Day since 2013 and since 2023 supports artists 'from all disciplines whose work helps encourage all sectors of society to engage and learn about the wonderful world of fungi' with the Massee Art Grant
• The Royal Botanical Garden, Kew publishes regularly a State of The World's Plants and Fungi, which is an 'in-depth look at how we can protect and sustainably use the world's plants and fungi for the benefit of people and the planet.' The last report has been published in 2020.

See also

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• Microbial art
• Agar art
• BioArt
• Mushrooms in art
• Human interactions with fungi
• Mycelium-based materials

References

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