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In: Mushroomology
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1.1 Diversity of edible and medicinal mushrooms across continents. In sub-panel a, the total number of edible and mushroom species, the number of edible species (E) and the number of medicinal species (M) for each continent (Africa, Asia, Europe, North America, South America, and Oceania) are provided separately. In sub-panel b, the vertical orange bars represent the numbers of unique species for each continent and their shared species with other continents, categorized below by gray dots and vertical lines. The horizontal blue bars display the total number of species for each continent. 8

1.2 The number of edible and medicinal mushrooms within families of Basidiomycota and Ascomycota as well as all described species of each family. The relationships among families were shown in an artificial family-level phylogenetic tree. Some genera with edible and medicinal mushrooms are retained their generic names due to their uncertain attributions at the family level. In the panel, the column of all represents the total number of described species of the corresponding clades, and that of edible & medicinal represents the total number of edible and medicinal mushrooms of the corresponding clades, while those of edible and medicinal represent the number of edible and medicinal mushrooms, respectively. The orders to which the included families as well as some genera belong are labelled in the purplish and greenish boxes. 10

2.1 Generalized schemes of the three major breeding systems within the fungal subkingdom Dikarya. Schematized are filamentous species that develop fruiting bodies for sexual spore production. Haploid nuclei are indicated by small circles within cells, with distinct genotypes (hereby explicitly referring to the mating type information present in heterokaryotic mycelia) distinguished by filled and open circles, respectively. For the sake of simplicity, the scheme on the left only considers life cycle control in unifactorial heterothallics by a single mat locus with bipolar meiotic segregation. In such cases, the typical meiotic tetrad consists of two nuclei of each parental mating type which are individually enclosed in the four different meiospores. Bifactorial heterothallics (not illustrated), on the other hand, may contain in their typical tetrads either two nuclei of each parental mating type, or two nuclei of two new recombinant parental mating types, or one nucleus per each parental mating type and each new recombinant mating type. In pseudo-homothallics with a single mat locus control, the sexual spores are heterokaryotic and contain nuclei of both parental mating types, as shown in the diagram on the right. In pseudo-homothallics with two mat loci (not illustrated), the spores can either carry both parental mating types or both recombinant mating types. 61

2.2 Life cycle of the model ascomycete Neurospora crassa. The heterothallic species has haploid mycelia of two alternate mating types mat A (= mat1-1) and mat a (= mat1-2) which can undergo the same developmental processes, as indicated by two mat-specific life cycles (the left one for matA and the right one for mata). The haploid, dikaryotic and diploid stages in the life cycles are indicated by light grey, white and dark grey shading. Main environmental factors that positively (black letters with white background) or negatively (white letters with grey background) influence steps in the life cycles are indicated. Each mating type can form haploid female (nucleus-receiving) sexual structures, the ascogonia, that are enveloped by sterile hyphae, thus forming the protoperithecia. These are fertilized by mitotic spores or vegetative hyphae of the opposite mating type with the help of a specialized female cell, the trychogyne. Fertilization leads to dikaryon formation and the generation of ascogenous hyphae that are a prerequisite for ascus and ascospore formation inside the growing haploid fruiting body (perithecium). Note that the dikaryotic stage in ascomycetes is very short. In the young dikaryotic ascus, karyogamy takes place between haploid nuclei of different mating types, followed by the two meiotic division and one post-meiotic mitosis, after which eight haploid ascospores develop in the ascus. Note that information here is limited to environmental factors, mating-type loci and pheromones/pheromone receptors, because description of the many known genetic determinants is beyond the purpose of this review. 74

2.3 Reciprocal courtship and fertilization of Neurosora crassa between two strains of different mating type is controlled by mating-type-specific gene expression of pheromone and pheromone receptor genes. Each figure part shows a scheme of the interaction between two different mating types, with one acting as female partner, generating ascogonial coil and protoperithecium with trychogyne, and the other one acting as male partner, generating a microconidium. Macroconidia and vegetative cells can also act as male partner, but are not included here for clarity. During fertilization, a “male” nucleus from the microconidium is translocated through the trychogyne to the “female” ascogonium. Below the schemes, a detailed cellular view of the interaction between trychogyne and conidium is given, including details of mating type-specific gene expression. A. Courtship between a female-acting mat1-1 strain and a male-acting mat1-2 strain. As shown at the top, a female-acting mat1-1 strain forms protoperithecia with ascogonia (ascogonial coils) at their base (shown in white), from which a trichogyne (also shown in white) grows and searches for a suitable attracting male cell for courtship (e.g. a microconidium shown in grey acting as spermatium) of a male-acting mat1-2 strain and which finally takes up the cell nucleus from it during fertilization. B. Courtship between a female-acting mat1-2 strain and a male-acting mat1-1 strain. The situation is reciprocal to the one shown in A. with the mat1v2 strain being the female-acting partner (with ascogonial coil and trichogyne shown in grey) and the mat1-1 strain being the male-acting partner (with a microconidium acting as spermatium shown in white). In the detailed scheme of the enlarged cells in A. below, mat1-1 is located in the nucleus of the female trichogyne and mat1-2 in the nucleus of the spermatium. In B., mat1-2 is located in nucleus of the female trichogyne and mat1-1 in the nucleus of the spermatium. The nuclei with crucial chromosomal segments are highlighted in grey. The structures of the alternate mat1 idiomorphs on chromosome 1 (Chr 1) are indicated in the respective mat1-1 and mat1-2 nuclei with genes being either shown in white (mat1-1) or in grey (mat1-2) while conserved neighbouring genes are shown in black (including an ATPase gene ar-2 needed in N. crassa for sexual development, Randall and Metzenberg 1998; for other genes please see text). Pheromone receptor gene pre-1 (on Chr 3) and pheromone gene ccg-4 (on Chr 1) and their products regulated by mat1-1 are also shown in white. Pheromone receptor gene pre-2 (on Chr 7) and pheromone gene mfa-1 (on Chr 5) and their products regulated by mat1-2 are also shown in grey. The cell walls as the margins of the cells are presented by the outer thick black lines, under which the protoplast membrane is located. Expressed pheromone receptors integrate into the membrane to which the cognate pheromones must bind for pheromone signaling transfer. In each individual trichogyne-microconidium combination, the essential protein products are selectively expressed so that courtship and fertilization can take place between a female trichogyne and a male cell of the opposite mating type. Gene fmf-1 on Chr 1 shown as grey-white striped is a Schizosaccharomyces pombe Ste11-like gene for an HMG-box TF that is required in both female- and male-acting cells for regulation of pheromone signalling and fertilization. 76

2.4 Life cycle of a tetrapolar heterothallic Agaricomycete with nuclear phases (haploid, dikaryotic, diploid) under complex network control of development by the two mat loci and additional environmental factors as known in C. cinerea. Heterokaryosis with different alleles in both mat loci, HD and PR, is necessary to form a fertile dikaryon after mycelial fusion of two compatible self-sterile homokaryons. General environmental factors with influences on mycelial growth of homokaryons and dikaryon and fruiting body induction and maturation are nutrients, temperature, humidity and conditions of aeration; increase in CO2 blocks mycelial growth and (normal) development. On the dikaryon, dark and blue light phases, in close interaction with mat genes of both loci, determine the onset and progress of fruiting or, dependent on colony age and the light intensity, alternatively the disintegration of a dikaryon into its homokaryotic components (e.g. in C. cinerea via light-induction of oidiophores in aerial mycelium and mitotic haploid oidia produced at their tips, Polak et al. 1997; deduced from work of S. commune, increased mycelial aeration interrupts HD-regulated nuclear pairing and subsequently PR-induced gene expression, Schuurs et al. 1998). Fruiting body development at various subsequent stages is further controlled by dark/light conditions and CO2 concentrations, karyogamy to proceed in the basidia needs blue light and compatible PR genes, while meiosis is blocked by nitrogen. 93

2.5 Mating-type loci matHD (in short HD, classically the A loci) and matPR (in short PR, classically the B loci) of C. cinerea. A. Simplified schematics of mat loci and their allelic interactions in formation of a fertile dikaryon from two compatible sterile homokaryons having different mat loci alleles. Gene in different HD and different PR alleles are marked in white and grey boxes, respectively. A black bar within HD1 genes and a white bar within HD2 genes indicate the sequence regions encoding the distinct homeodomains in the TF products of the HD locus. Ph and Ste3 designate pheromone precursor and pheromone receptor genes in the PR locus, respectively. The second number in names of genes symbolizes the respective alleles. Within the dikaryon, dashed lines with two-sided arrows mark for the ferile dikaryon compatible interactions between genes of the HD and genes of the PR locus. B. Top: The structures of the two best characterized and fully sequenced A alleles (A42 and A43) with three to five complete or partial paralogous HD1-HD2 gene pairs. Grey shading indicates regions of high homology between A42 and A43, non-shaded regions indicate levels of high sequence dissimilarity. The bipartite A locus is split into the Aα sublocus with the functional gene pair a and the Aβ sublocus with functional gene pairs b and d (positions and extensions of the Aα and Aβ subloci are marked by vertical black lines above the two maps). The internal 7 kb-long region of homologous sequence in between allows free recombination between Aα and Aβ. Positions and names of gene pairs (a to e pair) are indicated below the subloci by horizontal black and grey lines, where black means group with proven function and grey no shown function. Different alleles of gene pairs in A42 and A43 are characterized by different patterns in the respective gene boxes, with the arrows indicating the direction of transcription; identical patterns mark identical alleles. Combinations of HD1 and HD2 genes of allelic gene pairs shown by cloning and transformation to functionally interact are marked by dashed lines with two-sided arrows. Genes of conserved sequences (mip and β-fg) flanking upstream and downstream the alleles of the HD locus are presented by black boxes. B. Bottom: The structures of the two well characterized B alleles B24 and B43. Positions and names of paralogous groups of genes (Group 1 to Group 3) are indicated by horizontal black lines, different alleles of genes are marked by different patterns in the respective gene boxes. Ph and Ste3 genes of allelic groups shown by cloning and transformation to functionally interact are marked by dashed lines with two-sided arrows. Extra genes with grey bordered boxes were found by genome sequencing and are not yet characterized. Only the B43 allele is fully sequenced making it is possible that there are additional unidentified genes in B42. 95

4.1 Representatives of white-rot (A), brown-rot (B), grass-rot (C), plant symbiotic (D), insect fungi symbiotic (E), plant parasitic (F), and insect parasitic (F) fungi. 190

4.2 The life cycle of a basidiomycete fungus and its nuclear pairing and fusion. A, the life cycle of a basidiomycete fungus. B, in dikaryons, karyogamy occurs at the end of the life cycle immediately before the onset of meiosis, with the two parental nuclei remaining unfused and sharing the same cytoplasm throughout the life cycle in cells. 192

4.3 Schematic view of the plant lignocellulose structure and enzymes involved in their degradation. Polymeric cellulose microfibrils are tightly packed in ordered layers surrounded by branched hemicellulose chains and aromatic lignin heteropolymers. 203

4.4 Overview of laccase expression regulation in fungal cells. This figure summarizes the factors that induce differential laccase gene expression and the mechanisms of laccase gene regulation pathways reported to date. 207

5.1 A di-allele matrix. The constituent homokaryons of selected heterokaryons are recovered by protoplasting. The homokaryons are crossed in all possible combinations and evaluated for the trait of interest. In this way the breeding value of each homokaryon can be estimated in different genetic backgrounds and used to make a selection the parental homokaryons to generate segregating populations. 227

5.2 A breeding scheme to introduce QTL of donor lines into two acceptor lines. This approach intents to avoid inbreeding depression and is described in details in the “Outline of an effective QTL analysis and breeding strategy” paragraph. All blue lines indicate outcrosses and red lines backcrosses. 230

6.1 Examples of Agaricomycetes species with diverse lifestyles and fruiting body morphologies. Top row from left to right three white rot species, Flammulina velutipes, Nemecomyces mongolicus and Cyclocybe aegerita. The middle row shows litter decomposers, Coprinopsis pseudonivea and Mycena renati, the latter being a representative of a genus with the most expanded gene coding repertoires known to date. Also shown are two ectomycorrhiza species, the puffball-shaped Pisolithus tinctorius and the death cap Amanita phalloides, the latter being a focus of intense research on toxin systems. Bottom row: Calocera cornea, an early-diverging mushroom-forming species in the Dacrymycetes that causes brown rot; Hygrophoropsis aurantiaca (Boletales), another brown rot species, and Amanita vittadinii an edible Amanita that is considered a saprotrophic litter decomposer. 241

6.2 The accumulation of genomes of mushroom-forming fungi over the last 2 decades (A) and the frequency of keywords in abstracts of corresponding papers illustrated as a word cloud (B). Word size corresponds to frequency of occurrence. 243

6.3 The distribution of assembly sizes (A) and the number of protein coding genes (B) in published genomes available in MycoCosm (as of January 2024). Data based on statistics provided by the Joint Genome Institute. Data for Mycena olivaceomarginata (97000 genes) have been omitted from (B) for visualization purposes. 255

8.1 Mechanism of the CRISPR/Cas9 gene editing system. 308

9.1 Mutation accumulation studies are designed to estimation mutation rates by growing cells in the absence of selection. A) The fluctuation assay was used originally to show that mutations are random and not caused by the selective environment (Luria-Delbrück experiment). In this example, mutation at the ura3 locus is used to estimate rates of mutation to null alleles. A single genotype is grown in permissive conditions that counter selects for mutations, in this case minimal medium lacking uracil. After growing numerous small populations in numerous replicates, some populations are used to estimate total population per well (e.g., via a Coulter counter), and other populations are plated in their entirety onto selective media (e.g., 5FOA) to estimate the total number of ura- cells per population. B) Mutation accumulation with yeast involves streaking single colonies out from plate to plate. Using the estimated number of cells per colony allows the number of transfers to be translated to the number of generations. Transfers can be streaks or using spread plating of diluted colonies. C) With a mold species such as Aspergillus, an individual clone from a spore can be isolated and used to establish lines that are independent. After a certain number of days, the margin of the mycelium can be sampled for conidia, and a small number of conidia used to start the next passage after dilution in water. D) For species that do not sporulate, mycelium can be transferred in between passages. In order to reduce the effective population size Bezemova et al. (2020) used thin tubes of either 4 mm or 0.8 mm thickness to reduce competition among nuclei at the leading margin of the growing mycelium. After the mycelium reaches the end of the tube, the hyphae are seeded into clean tubes of media for another passage. 330

9.2 Genetic diversity in cultivated A. bisporus strains is limited. Phylogeny of major clades of Agaricus bisporus and related species based on mitochondrial DNA SNPs. Cultivars and their escapes are only known from Clades Europe II & III, which show low diversity within each clade. Shown in pie charts are the proportion of strains assigned to cultivar, wild, or escape. Branch lengths are scaled to millions of years using a molecular clock. 333

9.3 Evidence of strain degeneration in Cordyceps militaris. A-B. Healthy fruiting body production observed on typical strain grown on wheat (A) and vigorous fruiting on an inoculated pupa (B). C-D. Degenerated strains after repeated subculturing and produce fewer and poorly pigmented fruiting bodies in jar cultures (C) and fail to produce fruiting bodies when inoculated on a pupa (D). Photos generously provided by Mengqian Liu and Caihong Dong, Chinese Academy of Sciences. 335

10.1 Mass of tissue symptom of Dry Bubble Disease 353

10.2 Split stem symptom of Dry Bubble Disease 354

10.3 Trichoderma aggressivum f. aggressivum (Ta2) growing on the surface of the casing. It is dark green with a white fringe. 357

10.4 a) White fluffy growth symptom of cobweb disease and b) Fleshy color spotting symptom 360

10.5 S. megalocarpus sporangiospores on a branched sporangium 362

10.6 Zygospores of S. megalocarpus that show the dark colored, thick-walled spores 363

10.7 S. megalocarpus growing on an A. bisporus with the web or bread hair-like growth 363

10.8 Internal decay of the mushroom fruiting body caused by S. megalocarpus 364

10.9 Wet Bubble Disease showing the amber colored drops symptom 365

10.10 Bacterial Blotch discoloration symptom 367

10.11 Misshapen mushrooms symptom of mummy disease 370

10.12 a) Premature cap opening symptom of LaFrance Disease b) Drumstick-shaped mushroom symptom 373

10.13 Symptoms of MVX-PD: bare patches in crop 375

10.14 Symptoms of MVX: coffee-colored mushrooms 376

10.15 Mushroom phorid fly (Megaselia halterata) life cycle 378

10.16 S. carpocapsae nematodes at different life stages. A) S. carpocapsae infective juveniles. B) Mature adult S. carpocapsae bursting from a dead M. halterata larva. C) S. carpocapsae within a larval cadaver. 380

10.17 Stratiolaelaps scimitus predatory mites. A) A S. scimitus mite (PM) next to a Tyro-phagus spp. mold mite with a penny for scale. B) Adult S. scimitus mites feeding on a M. halterata larva. 381

10.18 Attract-and-kill station on a farm. This attract-and-kill station used natural light as an attractant at a vent window, with an electrostatically charged net blocking fly movement. Powdered insecticides were applied to the insecticidal nets to kill adults that were behaviorally trapped at the station. 382

10.19 Fungus gnat (Lycoriella ingenua) life cycle 383

11.1 Examples of white and cremini button mushrooms in growth rooms (A and B), and all three types from markets (C: top, portobello; middle, cremini; bottom, white). 397

11.2 Full colonization of the substrate by A. bisporus mycelia 402

11.3 Mushroom picking is a labour-intensive process that requires significant hand skills to be effective 406

11.4 Packaged whole and sliced button mushrooms as well as portobello mushrooms 407

12.1 Substrate preparation for growing oyster mushrooms 423

12.2 Bagging and sterilization of substrates for growing oyster mushrooms 424

12.3 Incubation of fruiting substrates inoculated with oyster strains 425

12.4 Fruiting of oyster mushrooms 426

13.1 Fruiting body of L. edodes 433

13.2 Mixing sawdust and other materials for growing Lentinula edodes 439

13.3 Bagging of mixed substrate for growing Lentinula edodes 440

13.4 Traditional method for sterilizing shiitake-growing substrates 441

13.5 Inoculation of Shiitake spawns into sterilized substrate 442

13.6 Mycelium growth and colonization of substrates by Shiitake strains 443

13.7 Formation of dark brown mycelium 443

13.8 Harvesting shiitake grown on inclined artificial logs 445

13.9 Injecting water into the bag 446

14.1 Life cycle of the straw mushroom 453

14.2 Cultivation equipment and facility for the straw mushroom. A) straw mushroom cultivation room; B) the corridor equipment of straw mushroom cultivation room; C) Straw mushroom cultivation shelf; D) Cultivated straw mushrooms 458

15.1 Life cycle of Flammulina filiformis 467

15.2 Flow chart of Flammulina filiformis cultivation technology 472

15.3 Wrapping with plastic sheet 478

16.1 Seedlings of Pinus armandii before addition of truffle spore inoculum 488

16.2 Differences in the growth of seedlings of Pinus armandii inoculated with truffle spores (right) or not (left). 489

16.3 Ectomycorrhizas formed by Pinus armandii and T. indicum. a: naked eye view of mycorrhizas under one root system; b: one mycorrhiza under dissecting microscope 489

16.4 Measuring the percentage of truffle mycorrhizal colonization 489

17.1 Auricularia heimuer sexual life cycle 503

17.2 Representative cultivars of Auricularia cornea 507

17.3 Small-scale (left) and large-scale (right) autoclave 509

17.4 Manual (left) and automated (right) spawn inoculations. 510

17.5 A: Incubation room components and set up. B: Full mycelia colonization. 511

17.6 Bag openings for inducing fruiting body formation after full colonization 512

17.7 Hole-puncher for fungal fruiting bags 513

18.1 Production facility of P. portentosus in Jinghong City of Yunnan Province, China 521

18.2 P. portentosus produced in large-scale factory facility. 526

18.3 Appearance of Cultivar YL1701-2 530

19.1 Fruiting body of Hericium erinaceus 541

19.2 Fruiting room for Hericium erinaceus 548

20.1 Tree cutting and wood sectioning for growing Ganoderma mushrooms 561

20.2 Bagging of wood sections for growing Ganoderma mushrooms 562

20.3 Stacking and sterilizing bagged wood segments for growing Ganoderma mushrooms 563

20.4 Arrangement, bag removal, and soil covering of wood sections colonized with Lingzhi mycelia 564

20.5 Growth management of Ganoderma fruiting bodies 567

20.6 Collection of Ganoderma spore powder using paper bag sleeves 568

20.7 Collection of Ganoderma spore powder in a nonwoven shed 536

20.8 Management of Lingzhi production in shelf-based cultivation of substitute substrates 571

20.9 Root art and auspicious cloud Lingzhi 572

20.10 Cultivation of large Lingzhi fruiting bodies 573

20.11 Cultivation of antler-shaped ganoderma 573

20.12 Cultivation of tower-shaped bonsai Lingzhi 574

21.1 Cyclocybe chaxingu (HFJAU1332). A, B: Basidiomata; C: Basidia; D: Pleurocystidia; E: Basidiospores; F: Pileocystidia; G: Cheilocystidia; H: Colonies. Bars=10 μm. 579

21.2 Differences in pileus color exhibited by different cultivars. a, Gancha AS-1; b, Gucha 1; c, Gucha 2; d, A cultivar named Baicha with white pileus 584

21.3 The mycelium of Cyclocybe chaxingu growing inside mushroom bags 589

21.4 Wall bag cultivation of Cyclocybe chaxingu 589

22.1 Four cultivated cordycipitoid fungal species from natural habitat. A, Ophiocordyceps sinensis; B, Cordyceps militaris; C, Tolypocladium guangdongense; D, Cordyceps chanhua. 597

22.2 Fruiting body cultivation of Ophiocordyceps sinensis on host insects 601

22.3 Fruiting body cultivation of Cordyceps militaris on artificial media (left) and insects (right) 602

22.4 Fruiting body cultivation of Tolypocladium guangdongense on artificial media 603

22.5 Fruiting body cultivation of Cordyceps chanhua on artificial media 604

23.1 A: Maximum likelihood (ML) phylogenetic analysis of species in the genus Morchella based on a four-gene data set (ITS plus EF1-a plus RPB1 plus RPB2), with Verpa and Disciotis as outgroups. B: The relationships among the proposed four groups (blushing morels, yellow morels, black morels, and semi-free-capped morels), three clades (Rufobrunnea Clade, Esculenta Clade, and Elata Clade), and three sections (Rufobrunnea, Morchella, and Distantes). 611

23.2 The revised life cycle of Morchella based on Du and Yang (2021) 616

23.3 Main cultivars. A: Morchella importuna; B: Morchella sextelata; C: Morchella eximia 618

23.4 Cultivation stepwise protocols and cultivation modes of true morels 620

23.5 Harvesting and processing of cultivated true morels 625

24.1 Examples of mushroom stamps 641

24.2 Local merchandise based on large underground mushroom mycelia reported in scientific and popular press 642

24.3 Underground mycelial network of Termitomyces 643

24.4 Harvested fruiting bodies of Tricholoma matsutake 643

24.5 Commercially cultivated and harvested Flammulina filiformis and naturally found fruiting bodies 644

24.6 Representative pictures of Ophiocordyceps sinensis 646

24.7 Ganoderma in building wall and porcelain 652

24.8 Tianquan Mushroom Museum in Changshan, Zhejiang, China 653

24.9 Representative mushroom dishes 654

25.1 Biosynthesis pathway of terpenoid in Basidiomycetes and Ascomycetes. This schematic pathway was drawn based on the information reviewed in reference 662

25.2 Shikimate biosynthetic pathway in mushroom. The full name and the respective abbreviation of compounds are as follows, 5-enolpyruvylshikimate-3-phosphate (EPSP); phosphoenolpyruvate (PEP); 3-deoxy-D-arabino-heptulosonate-7-phosphate (DAHP); dehydroquinate (DHQ); 3-dehydroshikimate (DHS); shikimate-3-phosphate (S-3-P); The bullets/numbering (1)–(7) indicate respective enzymes catalyzing the seven steps of shikimate pathway: (1) DAHP synthase; (2) dehydroquinate synthase (DHQS); (3) dehydroquinate dehydratase (DHQase); (4) Shikimate dehydrogenase (SD); (5) Shikimate kinase (SK); (6) EPSP synthase; (7) chorismate synthase. 665

27.1 Structures of toxic cyclopeptides 740

28.1 Structures of C8-oxylipins, their perceived odour and biosynthetic pathway of oct-1-en-3-ol synthesis via the oxygenation of linoleic acid to 10-(S)-hydroperoxy-9,11-octadecadienoic acid (10-HPOD) by a dioxygenase (DOX) and the subsequent cleavage by a hydro-peroxide lyase (HPL). 757

28.2 Schematic biosynthesis of isopentenyl diphosphate (IPP) and dimethylallyl diphosphate (DMAPP) in fungi, which are linked to generate the geranyl diphosphate (GPP), farnesyl diphosphate (FPP) and geranylgeranyl diphosphate (GGPP), the precursors for volatile and semi-volatile terpenes. The enzymes in the boxes are acetyl-CoA C-acetyltransferase (ACAT), 3-hydroxy-3-methylglutaryl-CoA synthase (HMGS), 3-hydroxy-3-methylglutaryl-CoA reductase (HMGR), mevalonate kinase (MVK), phosphomevalonate kinase (PMVK), mevalonate diphosphate decarboxylase (MVD), isopentenyl diphosphate isomerase (IDI), GPP synthase (GPPS), FPP synthase (FPPS), GGPP synthase (GGPPS). 762

28.3 Structures of several volatile terpenes found in mushrooms. Beneath the trivial names are listed the perceived odours, if described. 766

28.4 Structures of volatile sulphur compounds (VSC) found in mushrooms 770

29.1 Lion’s mane (Hericium erinaceus) growing from a kit indoors 792

29.2 Blue Oyster Mushrooms (Pleurotus ostreatus) ready for harvest, growing under 6500K LED lights. 792

29.3 Fruiting mushroom blocks on a shelf at Nature Lion’s mushroom farm 793

29.4 Harvesting lion’s mane mushrooms. (Hericium erinaceus) 793

29.5 Adding chopped straw to a sack and pasteurize in boiling water. 798

30.1 The real mycelium-based leather products of Reishi™ from MycoWorks and Mylea™ from Mycotech Lab. 821

30.2 The real mycelium-based packing products from Grown.bio 822

30.3 The real mycelium-based building products from Mogu 823

30.4 The real mycelium-based furniture interior design products from Grown.bio and Ecovative 823

30.5 The mycelium-based elect electronic components. (a) mycelium skin with Cu-Au traces; (b) demonstration photograph of a mycelium-based sensor board with different elements; (c) demonstration of the mycelium-based battery; (d) section view of the mycelium zinc-carbon battery. 824

30.6 Commercial-scale production of mycelium-based materials. (a) Bioreactor of mycelium leather production at VTT, (b) vertical farming of aerial mycelium foam ©Bolt Threads, (c) and (d) commercial-scale production farm of ©Ecovative 828

Tables

1.1 Jaccard similarity among six continents 9

A1 The classification of edible and medicinal mushrooms above family level, primarily sourced from the Mycobank database along with additional references. 12

2.1 Molecular structure of mat loci of cultivated mushroom species from the Basidiomycota, Agaricomycetes 101

3.1 Common cultivated mushrooms and their germplasm preservation centers in the world 158

3.2 Representative examples of population genetic studies of edible mushrooms 177

5.1 An overview from all genetic linkage maps generated for edible mushrooms since 1993 218

5.2 Quantitative trait loci (QTL) studies in edible mushrooms done from 1999 up to 2023 220

7.1 Advances in research on edible fungi transformation in the past two decades 284

7.2 Examples of successful gene editing and silencing in edible fungi 287

10.1 Prevalence of two viral diseases on Agaricus bisporus mushrooms 372

11.1 Compositions of two common types of composts used for growing A. bisporus 401

16.1 Advantages and disadvantages of three different types of inoculums in producing mycorrhizal seedlings 487

20.1 Commonly used formulas for Lingzhi substitute substrate cultivation 570

22.1 Taxonomy and host range of four cultivated cordycipitoid fungi 597

25.1 Representative mushroom pigments sorted by their biosynthesis pathways and the mushroom producers 669

26.1 Effects of adding edible mushrooms on the rheological, cooking, and microstructural properties of staple composite foods 714

26.2 Selected dietary supplements based on mushrooms 717

26.3 Cosmetic products containing mushrooms and their ingredients 722

27.1 The new species of poisonous mushrooms reported from 2012 to 2022 737

28.1 Key aroma compounds of some of the most valuable mushrooms 774

29.1 A list of select mushroom grain spawn and grow kit suppliers 795

29.2 Hardwood sawdust & wheat bran growing block recipe 797

29.2 Straw growing block recipe 797

30.1 The spent mushroom substrates (SMS) used for organic fertilizer productions 811

30.2 The spent mushroom substrates (SMS) used for new mushroom cultivations 813

30.3. The spent mushroom substrates (SMS) used for biofuel productions 815

30.4 Examples of spent mushroom substrates (SMS) used for crop disease and pest management 816

30.5 Bioremediation of environmental contaminants by spent mushroom substrates (SMS) 818

30.6 Mushroom species used for mycelium-based materials 826

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Mushroomology

Cover Mushroomology
E-Book ISBN:
9789004751699
Publisher:
Wageningen Academic
Print Publication Date:
25 Feb 2026
Front Matter
Preliminary Material
Copyright Page
Figures and Tables
Chapter 1 Diversity and taxonomy of edible and medicinal mushrooms
Chapter 2 Diversity of life cycles, breeding systems, and mating types in mushrooms
Chapter 3 Mushroom population genetics and germplasm
Chapter 4 Mushroom physiology and nutritional requirements
Chapter 5 Quantitative trait loci mapping and mushroom breeding
Chapter 6 Mushroom genomics – the history, driving forces and current state of the genomics of mushroom-forming fungi
Chapter 7 Molecular biology and biotechnology of mushrooms and genetically modified mushrooms
Chapter 8 Gene editing in edible and medicinal mushrooms and applications
Chapter 9 Mutation and strain degeneration in mushrooms
Chapter 10 Agaricus mushroom pests and pathogens
Chapter 11 Agaricus bisporus, the button mushroom
Chapter 12 Biology and cultivation of oyster mushrooms
Chapter 13 Lentinula edodes, the Shiitake Mushroom
Chapter 14 Volvariella volvacea, the straw mushroom
Chapter 15 Flammulina filiformis, the golden needle mushroom
Chapter 16 Tuber spp., the true truffles
Chapter 17 Biology and cultivation of Auricularia heimuer and Auricularia cornea
Chapter 18 Phlebopus portentosus, the cultivated bolete
Chapter 19 Hericium erinaceus, the lion’s mane mushroom
Chapter 20 Ganoderma sichuanense, the cultivated lingzhi
Chapter 21 Cyclocybe chaxingu, the chashugu
Chapter 22 Cordycipitoid fungi (Cordyceps sensu lato)
Chapter 23 Biology and cultivation of true morels (Morchella spp.): current progress and future trends
Chapter 24 Mushroom culture
Chapter 25 Mushroom pigments: biosynthesis, diversity, ecological roles, and applications
Chapter 26 Mushroom processing and processed products
Chapter 27 Species diversity and poisoning classification of poisonous mushrooms
Chapter 28 Mushroom volatiles – diversity, synthesis, function, and applications
Chapter 29 Introduction to home growing mushrooms
Chapter 30 Mushrooms in a changing world
Back Matter
Index

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