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Introduction to Truffles and Their Use 20190812 2019-08-12 Introduction to truffles and their use – Truffle – Evolution Christina Wedén – Phylogeny Uppsala University – The species concept August 2019 – False and true truffles – Sexual and asexual reproduction What is a truffle? What is a fungus? • Mycology historically part of botany • Still part of botany as a scientific subject • Eukaryotic cells (true nucleus with chromosomes) • Common ancestor with animals • Unicellular (yeasts) and multicellular (mycelial) fungi 1 2019-08-12 Carl von Linné 1707-1778 Elias Fries 1794-1878 The flower king The father of mycology Life on Earth Eukaryotes... Phaeophyta Chrysophyta Xanthophyta EUKARYOTA Oomycota ... Fungi Dinoflagellata Ciliata Apicomplexa Animalia Rhodophyta Myxomycota Chlorophyta Entamoeba Cyanophyta Amoeba Kinetoplastida !-proteo Euglenophyta Trichomonads Microsporidia EUBACTERIA Diplomonads ARCHAEBACTERIA Redrawn and adjusted after M. L. Sogin, Current Opinions in Genetics and Development 1: 457 (1991) by Anders Backlund Från Baldauf et al., The Deep Roots of Eukaryotes Science 300: 1703-1706 (2003). 2 2019-08-12 biology - chemistry Yeast cells and human cells • Biosynthesis of substances in organisms, and share a major part of their cell biology. in different parts of organisms, to enable life …Although major differences in life forms! • Primary metabolites (substances) carry out the basic functions of the cell • Specialized metabolites (”secondary metabolites”) enable survival of the organism and of the species Two examples of specialized metabolites The fungal cell from fungi are: • penicillin produced by Penicillium species (moulds) to fight bacteria in competition for subtrate • sulphur containing volatiles produced by truffle species to attract spore vectors 3 2019-08-12 • Hypha = fungal cell • Hypha grow linked to each • Linked hypha grow out to form a branched mycelia • Fruiting bodies are the part of the fungus that Fungi grow as mycelia we often see, but only consists of around 1% – a 3D network of elongated of the total fungal biomass – 99% is vegetative fungal cells (hyphae) mycelia. deeplivingproject.com.au Source: Source: 4 2019-08-12 What is a truffle? • Although a single hyphal strand is not visable to the naked eye, densely growing mycelia can • Fungal species which have hypogeous be seen e.g. under bark, growing on the (subterranean) fruiting bodies kitchen compost or as mycelial tufts growing • Fruiting body = stucture carrying the spores out from mycorrhizal roots (shown below). which are the result of the fungus’ sexual reproduction • The fruiting body is made up of vegetative mycelia and spore tissue Vegetative mycelia Vegetative mycelia = white ”veins” = white tissue Mature spores Mature spores = brown tissue = brown tissue Outer peridium • The truffle life form has evolved multiple times in different groups of fungi Inner peridium Gleba Elaphomyces granulatus, the deer truffle (hjorttryffel) • Truffles are not an evolutionary distinct group, but instead belong to diverse fungal families Peridium • The truffle life form is thought to be an adaption to e.g. drought Gleba Tuber mesentericum, the Bagnoli truffle and Tuber aestivum, the Burgundy truffle (bagnolitryffel) (bourgognetryffel) 5 286 Phytochem Rev (2010) 9:279–301 species of macrofungi are known to science, but vectors, and to make them unearth and eat the truffles, 286 Phytochem Rev (2010) 9:279–301 estimations show that this may well be only a tenth of thereby spreading the spores. Depending on the species of macrofungi are known to science, but vectors, and to make them unearth and eat the truffles, the actual number of all living macromycete species animal vector’s lifestyle and ecology, these spores estimations show that this may well be only a tenth of thereby spreading the spores. Depending on the the actual number of all living macromycete species animal vector’s lifestyle and ecology, these spores (Hawksworth 2001). For fungi in the widest sense, the may be deposited in the shape of fecal pellets close to (Hawksworth 2001). For fungi in the widest sense, the may be deposited in the shape of fecal pellets close to total number of species on Earth has been estimated to the roots of mycorrhizal trees (Trappe and Castellano total number of species on Earth has been estimated to the roots of mycorrhizal trees (Trappe and Castellano 2019-08-12 1.1 million (Mueller et al. 2007). Within the fungal 1991). Truffles also show high levels of endemism 1.1 million (Mueller et al. 2007). Within the fungal 1991). Truffles also show high levels of endemism kingdom lies a vast, and to a large extent, still (Mueller et al. 2007), which may be explained by unexplored potential for future pharmaceutical drugs. mammalian fecal pellets being their main mean of kingdom lies a vast, and to a large extent, still (Mueller et al. 2007), which may be explained by In the section Fungi in Linnaeus’ Materia Medica spore dispersal (Fig. 5). (1749), the genus Lycoperdon is mentioned as an unexplored potential for future pharmaceutical drugs. mammalian fecal pellets being their main mean of aphrodisiac and remedy against flatulence. Most In the section Fungi in Linnaeus’ Materia Medica spore dispersal (Fig. 5). probably this passage refers to what today is known (1749), the genus Lycoperdon is mentioned as an as the deer truffle, Elaphomyces granulatus, earlier Basidiomycetes have basidiospores. The known under the name Boletus cervini. In more So called false truffles belong to the basidiomycetes recent sources the deer truffle has been reputed for its basidiumaphrodisiacis a structure which andholds remedyand against flatulence. Most abilities to induce oestrus manifestation in cows dischargeprobably the spores this. passage refers to what today is known (Klintberg 1998), forming a somewhat unsuspected link to the Linnaean indication. The edible truffles of as the deer truffle, Elaphomyces granulatus, earlier the genus Tuber have historically been used as an aphrodisiac by both men and women and in modern known under the name Boletus cervini. In more science small amounts of the steroid androstenol has recent sources the deer truffle has been reputed for its been found in mature specimens of the black truffle Tuber melanosporum. Androstenol is a major com- abilities to induce oestrus manifestation in cows ponent of the boar pheromone, excreted in the saliva (Klintberg 1998), forming a somewhat unsuspected during mating (Claus et al. 1981). Truffles are an intricate fungal life form occurring link to the Linnaean indication. The edible truffles of in both the two major groups of macromycetes: asco- and basidiomycetes (see Fig. 4). Only within the Ascomycetesthe genushave ascosporesTuber . haveThe ascus historicallyis been used as an ascomycetes, species that develop truffle fruit bodies a sporeaphrodisiacsack which encloses by boththe spores menand and women and in modern are estimated to have evolved at least fifteen times in open to discharge them at spore maturity. 286 multiple families (Læssøe and Hansen 2007). Thus, Phytochem Revscience (2010) small 9:279–301 amounts of the steroid androstenol has from a modern phylogenetic perspective, truffles are regarded as a life form rather than a specific group of been found in mature specimens of the black truffle species of macrofungifungi. Despite are this, known all truffle species, to science, regardless of but vectors, and to make them unearthTuber and eat melanosporum the truffles, . Androstenol is a major com- phylogenetic placement, live in mycorrhizal symbio- estimations showsis that with thisvascular may plants. well Due to theirbe onlyhypogeous a tenthnature of thereby spreading the spores. Dependingponent of the on boar the pheromone, excreted in the saliva and subsequent loss of active spore dispersal, all the actual numbertruffle of species all living are also dependant macromycete on an animal vector species animal vector’s lifestyle and ecology,during these mating spores (Claus et al. 1981). (Hawksworth 2001for). spore For dispersal. fungi The in evolution the widest of a hypogeous sense, fruit the may be deposited in the shape of fecalTruffles pellets closeare an to intricate fungal life form occurring body might be an adaptation to dry climatic condi- total number of speciestions, but on the Earth relationship has between been the estimated hypogeous Bohlin to et al.the 2010 roots of mycorrhizal trees (Trappein both and the Castellano two major groups of macromycetes: asco- fungus, the tree and the animal vector seem to be 1.1 million (Muellerimperative et for al. the2007 life and). distribution Within of this the life form fungalFig. 4 Phylogeny1991 of). fungi Trufflesphylogeny of asco- also and basidiomy- show high levelsand basidiomycetes of endemism (see Fig. 4). Only within the cetes with focus on taxa with hypogeous fruiting bodies, kingdom lies asuggesting vast, and a more sophisticatedto a large fungal extent,niche against stilltruffles, based(Mueller on a combination et of al. several2007 published), studies which mayascomycetes, be explained species by that develop truffle fruit bodies vertebrates. The dispersal vectors may be mice, (Læssøe and Hansen 2007; Hibbet et al. 2007; Celio et al. unexplored potentialsquirrels, for deer future or wild pharmaceuticalboars (Trappe and Castellano drugs.2006). Taxamammalian labeled in blue with • indicate fecal evolutionary pellets groups being theirare estimated main mean to have of evolved at least fifteen times in 1991). Consequently the truffles, i.e. the hypogeous that have developed truffles. The most well known truffle-
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