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Fruit bodies: Their production and development in relation to environment

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CHAPTER 5
Fruit Bodies: Their Production and
Development in Relation to
Environment
David Moore, Alan C. Gange, Edward G. Gange and
Lynne Boddy
Contents 1. Introduction 80
1.1 Fungal Morphogenesis 80
1.2 Morphogenetic Control Elements 84
1.3 Importance of Sexual Reproduction 85
2. Physiological Factors Favouring Fruit Body Production 86
2.1 Carbohydrates 86
2.2 Nitrogen Sources 86
2.3 Nutrient Capture 87
2.4 Non-Nutritional Environmental Variables 91
2.5 Fruiting in the Natural Environment 94
3. Fruit Body Survival 96
4. Principles of Fungal Developmental Biology 98
4.1 Underlying Principles 98
4.2 Modelling Hyphal Growth and Fruit Body Formation 98
4.3 Data Mining Fungal Genomes 99
References 99
Abstract Sexual reproduction is important because it generates genetic variation,
offers an escape from DNA parasites and provides a means to repair DNA
damage. Many fungi exhibit particular patterns of sexual fruit body morpho-
genesis but the characteristics differ between species. However, it is possible
to generalise that within developing fruit body tissues, fungal cells embark
on a particular course of differentiation in response to the interaction of
their intrinsic genetic programme with external physical signals (light, tem-
perature, gravity, humidity), and/or chemical signals from the environment
and other regions of the developing structure. Fruit body morphogenesis is
affected by carbon and mineral nutrient availability, and environmental vari-
ables including temperature, water availability, CO
2
, light and interactions
British Mycological Society Symposia Series r2008 The British Mycological Society
Published by Elsevier Ltd. All rights reserved.
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with other fungi and bacteria. Changes in the seasonal pattern of fruiting in
the UK can be detected from field records made in the last 50 years, and
while not all species behave in the same way, mean first fruiting date is now
significantly earlier and mean last fruiting date is now significantly later,
which results in an extended fruiting season. Significant numbers of species
that previously only fruited in autumn now also fruit in spring. Such analyses
show that relatively simple field observations of fungi can detect climate
change, and that fungal responses are sufficiently sensitive to react to the
climate change that has already occurred by adapting their pattern of
development. Unfortunately, though it is possible to deduce the decisive
steps in development that are open to influence, the molecular controls that
normally regulate those steps remain unknown. Extensive genomic analysis
shows that sequences crucial to multicellular development in animals or
plants do not occur in fungal genomes, so we are ignorant of the basic
control processes of fungal multicellular developmental biology.
1. INTRODUCTION
We use the term fruit bodies to encompass all the structures that develop from
fungal mycelia to produce and distribute spores or other propagules, including
basidiomata—the structures that release sexual spores (meiospores) in Basidi-
omycota, as well as a range of structures that produce asexual spores (mito-
spores) and some somatic (vegetative) structures, such as stromata and sclerotia,
that can survive adverse conditions. Obviously, the phrase encompasses a very
wide range of organs but their common feature is that they are multicellular, and
their shape and form emerge as a result of a sequence of developmental adjust-
ments. That is, they exhibit a characteristic pattern of morphogenesis.
1.1 Fungal Morphogenesis
Within the developing tissues of a fruit body, cells embark on a particular course
of differentiation in response to the interaction of their intrinsic genetic pro-
gramme with external physical signals (light, temperature, gravity, humidity),
and/or chemical signals from other regions of the developing structure. These
chemicals may be termed organisers, inducers or morphogens, and may inhibit
or stimulate entry to particular states of determination. Chemical signals may
contribute to a morphogenetic field around a structure (cell or organ), which
permits continued development of that structure but inhibits formation of
another structure of the same type within the field. All of these phenomena
contribute to the pattern formation that characterises the ‘body plan’ created by
the particular distribution of differentiated tissues in the multicellular structure.
Pattern formation depends on positional information, which prompts or allows
the cell to differentiate in a way appropriate to its position in the structure and
may be conveyed by concentration gradients of one or more morphogens emitted
from one or more spatially distinct organisers. Pattern formation thus involves an
David Moore et al.80
instructive process, which provides positional information, and a second inter-
pretive process, in which the receiving cell or tissue responds.
Fungi are ‘modular organisms’ in which growth is repetitive, and a single
individual mycelium will have localised regions at very different stages of
development (Andrews, 1995). Consideration of developmental regulatory
systems is relevant to the current discussion because any effect of the external
environment on fruit body development must operate through an influence on
the control systems that determine the distribution and growth patterns of the
multicellular structure.
The constituent cells of a fungal fruit body are generally considered to be
totipotent (able to follow any pathway of differentiation), because a mycelial
culture can be produced in vitro from a fragment of a mature, fully differentiated
structure, e.g. a fruit body stem. This feature results in a morphogenetic plasticity
which surpasses that of other organisms and provides an intellectual challenge in
terms of developmental biology, taxonomy and genetics (Watling and Moore,
1994). The only exceptions to totipotency are the meiocytes (the cells within
which meiosis occurs), which are committed to sporulation once they have pro-
gressed through meiotic prophase (Chiu and Moore, 1988a, 1988b, 1990, 1993;
Chiu, 1996). On the other hand, even meiocytes can be ‘used’ for non-sporulation
functions: the hymenium of Agaricus bisporus is packed with basidia held in an
arrested meiosis and serving a purely structural function (Allen et al., 1992).
Differentiated fungal cells require reinforcement of their differentiation
‘instructions’. This reinforcement is part of the context within which they
normally develop, but when removed from their normal environment most
differentiated hyphae revert to vegetative hyphae. Hyphal differentiation is con-
sequently an unbalanced process in comparison with vegetative hyphal growth.
In most hyphal differentiation pathways the balance must be tipped in the
direction of ‘differentiation’ by the local microenvironment, which is, presumably,
mainly defined by the local population of hyphae.
Another common feature is that morphogenesis is compartmentalised into a
collection of distinct developmental processes (called ‘subroutines’; Figure 1;
Moore, 1998a). These separate (or parallel) subroutines can be recognised at the
levels of organs (e.g. cap, stem, veil), tissues (e.g. hymenophore, context,
pileipellis), cells (e.g. basidium, paraphysis, cystidium) and cellular components
(e.g. uniform wall growth, growth in girth, growth in length, growth in wall
thickness). They are distinct genetically and physiologically and may run in
parallel or in sequence. When they are played out in their correct arrangement
the morphology that is normal to the organism results. If some of the subroutines
are disabled (genetically or through physiological stress), the rest may still pro-
ceed. This partial execution of developmental subroutines produces an abnormal
morphology. The main principles that govern fungal development as deduced
from observation, experiment and computer modelling are summarised in
Table 1 (from Moore, 2005).
Fungal morphogenesis must be totally different from animals, because fungal
cells have walls, and from plants (whose cells also have walls) because hyphae
grow only at their tips and hyphal cross-walls form only at right angles to the
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Fruit Bodies 81
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Figure 1 Flowchart showing a Simplified View of the Processes involved in Development of
Fruit Bodies and other Multicellular Structures in Fungi (from Moore, 1998a) AU :5
.
David Moore et al.82
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Table 1 The Eleven Principles that Govern Fungal Development
Principle 1 The fundamental cell biology of fungi on which development
depends is that hyphae extend only at their apex, and cross-
walls form only at right angles to the long axis of the hypha
Principle 2 Fungal morphogenesis depends on the placement of hyphal
branches
Principle 3 The molecular biology of the management of cell-to-cell
interactions in fungi is completely different from that found
in animals and plants
Principle 4 Fungal morphogenetic programmes are organised into
developmental subroutines, which are integrated collections
of genetic information that contribute to individual isolated
features of the whole programme. Execution of all the
developmental subroutines at the right time and in the right
place results in a normal structure
Principle 5 Because hyphae grow only at their apex, global change to
tropic reactions of all the hyphal tips in a structure is
sufficient to generate basic fruit body shapes
Principle 6 Over localised spatial scales coordination is achieved by an
inducer hypha regulating the behaviour of a surrounding
knot of hyphae and/or branches (these are called Reijnders’
hyphal knots)
Principle 7 The response of tissues to tropic signals and the response of
Reijnders’ hyphal knots to their inducer hyphae, coupled
with the absence of lateral contacts between fungal hyphae
analogous to the plasmodesmata, gap junctions and cell
processes that interconnect neighbouring cells in plant and
animal tissues suggest that development in fungi is
regulated by morphogens communicated mainly through
the extracellular environment
Principle 8 Fungi can show extremes of cell differentiation in adjacent
hyphal compartments even when pores in the cross-wall
appear to be open (as judged by transmission electron
microscopy)
Principle 9 Meiocytes appear to be the only hyphal cells that become
committed to their developmental fate. Other highly
differentiated cells retain totipotency — the ability to
generate vegetative hyphal tips that grow out of the
differentiated cell to re-establish a vegetative mycelium
Principle 10 In arriving at a morphogenetic structure and/or a state of
differentiation, fungi are tolerant of considerable
imprecision (¼expression of fuzzy logic), which results in
even the most abnormal fruit bodies (caused by errors in
execution of the developmental subroutines) being still able
Fruit Bodies 83
long axis of the hypha. Consequently, fungal morphogenesis depends on the
placement of hyphal branches. A hypha must branch to proliferate. To form a
multicellular structure, the position at which the branch emerges and its direction
of growth must be controlled. A major aspect of that directional control is an
autotropism—a tropism to self—in which growth direction of each hyphal
branch is influenced by the position of the rest of the mycelium. Exploratory
mycelia experience a negative autotropism, which causes them to grow away
from the main mycelium and this maintains the outward exploration of the
substratum. On the other hand, to create a multicellular structure like a fruit
body, positive autotropism is essential to cause hyphae to grow together for
hyphal branches to cooperate and coordinate their activities. Tropic reactions
imply a signalling system, a signal sensing system and a reaction system. Math-
ematical models of these systems can be created very successfully (Stoc
ˇkus and
Moore, 1996; Mes
ˇkauskas et al., 1998, 1999a, 1999b, 2004a, 2004b; Moore et al.,
2006), but we know nothing yet about their biochemistry, cell biology or molec-
ular nature. However, it is clear that what mechanisms exist must be different to
animals and plants because gene sequences known to regulate development in
animals and plants do not occur in fungal genomes (Moore et al., 2005; Moore and
Mes
ˇkauskas, 2006).
1.2 Morphogenetic Control Elements
The only major morphogenetic control elements known in fungi are the mating
type factors, which regulate pheromone production and pheromone receptors
involved in mating, ranging from recognition between sexually competent cells
in yeast to governing growth of clamp connections, internuclear recognition and
regulation of the distance between the two nuclei in Basidiomycota (Casselton,
2002). However, not all fungi possess mating type factors, and, indeed, even in
species that have a well-developed mating type system apparently normal fruit
bodies can be formed by haploid cultures, and fruit body formation can usually
be separated from other parts of the sexual pathway by mutation (see Chapter 5
in Moore, 1998a).
Generally, vegetative compatibility genes define the individuals of fungal
populations, while mating type factors are usually interpreted as favouring the
outbreeding of a fungal population (Chiu and Moore, 1999). Consequently,
mating type genes contribute to management of the genetics of the population as
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Table 1. (Continued )
to distribute viable spores, and poorly (or wrongly)
differentiated cells still serving a useful function
Principle 11 Mechanical interactions influence the form and shape of the
whole fruit body as it inflates and matures, and often
generate the shape with which we are most familiar
Source: From Moore (2005).
David Moore et al.84
well as to the sexual development of the individual. Sexual reproduction
generates genetic variation, offers an escape from DNA parasites and provides a
means to repair DNA damage (Bernstein et al., 1985).
1.3 Importance of Sexual Reproduction
The crucial step in sexual reproduction, which provides the contrast with asexual
reproduction, is the fusion of nuclei derived from different individuals. If the
individuals involved in a mating have different genotypes, the fusion nucleus
will be heterozygous and the products of the meiotic division can be recombinant
genotypes. Thus, in one sexual cycle, new combinations of characters can be
created in the next generation for selection. Consequently, the most common
‘explanation’ for sex is that it promotes genetic variability through out-crossing
and that variability is needed for the species to evolve to deal with competitors
and environmental changes. There is plenty of evidence to show that asexual
lineages change little in time and that out-crossing certainly does promote var-
iability in a population, which enables the organism to survive environmental
challenges (Hurst and Peck, 1996; Burnett, 2003).
This, though, is a ‘group selectionist’ interpretation. It argues that variation
generated in an individual meiosis benefits the group or population to which the
individual belongs. Yet current theory prefers to emphasise that selection acts
on individuals (Carlile, 1987; Dawkins, 1989). A feature that is advantageous in
selection must be so because of benefit to the individual itself or its immediate
progeny. As noted above, an alternative interpretation of the selective value of a
sexual cycle suggests that repair of damaged DNA is the crucial advantage of
meiosis (Bernstein et al., 1985). It is argued that bringing together genomes from
two different individuals enables DNA damage in one parental chromosome,
caused by mutation or faulty replication, to be repaired by comparison and
recombination with the normal chromosome provided by the other parent.
Genetic fitness would be increased but only when out-crossing ensures
heterozygosis. Even an incomplete sexual cycle might be of advantage in this
case.
Gene mutations can be recessive and damaging, and different mutations are
likely to occur in different mitotically generated cell lines. Just the formation of
the diploid (or heterokaryon in most Basidiomycota) by out-crossing will benefit
the mated individual if recessive adverse mutations are masked by non-mutant
(‘wild-type’) alleles in the nuclei of the other parent. Out-crossing might also give
rise to heterozygous advantage, where the heterozygous phenotype is better than
either of its homozygous parents. This has been demonstrated frequently in
plants and animals, and also in Saccharomyces cerevisiae (James, 1960).
Clearly, the genotype of the parental mycelium makes a crucial contribution
to the genetics of the progeny population, but to produce a progeny population
the parental mycelium must first produce a crop of fruit bodies and to do that it
must grow into and through the substratum to capture, translocate and accu-
mulate sufficient nutrients to support the formation of what can be massive
multicellular structures.
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Fruit Bodies 85
2. PHYSIOLOGICAL FACTORS FAVOURING FRUIT BODY
PRODUCTION
Fungi enjoy an adaptable and flexible metabolism. It is unlikely that there is a
compound, organic or inorganic, on the planet that some fungus cannot utilise,
transform, modify or otherwise metabolise (see Chapter 3 in Moore, 1998a).
These versatile biochemical capabilities are used in a variety of ways during
morphogenesis in fungi and over the past century there have been numerous in
vitro studies of the nutritional physiology of fruit body production. Nutrients that
are inferred to be ‘favourable’ for fruiting are those that allow the organism to
exert its own intrinsic controls over the progress of its metabolism (Hawker,
1950).
2.1 Carbohydrates
An enormous volume of research has been done on this topic (for reviews see
Moore-Landecker, 1993; Jennings, 1995; Moore, 1998a), though it is important to
remember that conditions in the laboratory are far removed from the natural
environment. The crucial insights came from Hawker’s (1939, 1947) experiments:
simple sugars tend to favour asexual spore production while oligo- and poly-
saccharides are especially good carbon sources for production of fruit bodies.
Glucose often represses fruit body production, even in very low concentrations.
The rate with which a fungus can hydrolyse a carbohydrate determines the abil-
ity of the carbohydrate to promote fruit body formation (Hawker and Chaudhuri,
1946), so what seems to matter most is the rate of supply and ease of use of
substrates as determinants of their value in promoting fruit body formation. It
comes as no surprise, therefore, that saprotrophic Basidiomycota on dung fruit
more readily than those utilising leaf litter, and in turn than wood decomposers,
though, of course, these resources also differ in mineral nutrient content. Like-
wise, fungi that participate early in community development within a resource
fruit more readily than most later colonizers (Cooke and Rayner, 1984; Rayner
and Boddy, 1988; Chapter 11), whose carbon sources are more recalcitrant.
2.2 Nitrogen Sources
Similar conclusions are reached when attention turns to the ‘best’ nitrogen
source, which usually proves to be one amino acid or a mixture of amino acids. In
most cases inorganic nitrogen and ammonium salts fail to support fruit body
development although they may support production of primordia, but amino
acids are required to produce the mature fruit bodies (reviewed in Moore, 1998a).
This suggests that the formation of fruit body initials may be an activity of the
vegetative mycelium and it is their further development which constitutes the
fundamental ‘mode switch’ into the fruit body morphogenetic pathway. At least
some of the deleterious effects of ammonium salts may be due to their influence
on the pH of the medium, though metabolite repression caused by ammonium
ions in many Ascomycota may be another cause. Fruit body formation in some
fungi is favoured by provision of protein as source of nitrogen. Several
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David Moore et al.86
basidiomycetes (A. bisporus,Coprinus cinereus (¼Coprinopsis cinerea) and Volvari-
ella volvacea) are able to use protein as a carbon source as efficiently as they use
glucose (Kalisz et al., 1986), so an advantage of protein is that it serves as a source
of carbon, nitrogen and sulphur. In more natural conditions, A. bisporus and a
wide range of other filamentous fungi can utilise dead bacteria as sole source of
carbon, nitrogen, sulphur and phosphorus (Fermor and Wood, 1981; Grant et al.,
1986).
Higher carbon than nitrogen concentrations are usually required for fruit
body production but the optimum C:N ratio varies from 30:1 to 5:1 (references
in Moore-Landecker, 1993). High concentrations of amino acids tend to delay
and/or depress maturation of fruit bodies even in organisms in which fruit body
formation is optimal on media containing lower concentrations of amino acids,
an effect that may result from the production of large quantities of ammonium as
a nitrogen-excretion product on such substrates. When grown on protein as sole
carbon source, nitrogen needs to be excreted from the mycelium; when this
happens in vitro the ammonium concentration of the medium increases drastically
during mycelial growth. One-third to one-half of the supplied protein-nitrogen
was metabolised to ammonia by batch cultures of three saprotrophic bas-
idiomycetes when protein was the sole source of carbon (Kalisz et al., 1986).
2.3 Nutrient Capture
Hyphae absorb sufficient nutrients to support their active vegetative growth and
to allow accumulation of reserve materials, which may subsequently be trans-
located to sites of need, including developing fruit bodies. Fruit body primordia
may be fairly uniformly dispersed, but locations of enlarging and maturing fruit
bodies may be much less evenly spread. For example, in Coprinus lagopus, certain
favourably placed young fruit bodies may initiate a flow of nutrients in their
direction, others that are deprived then fail to mature (Madelin, 1956a, 1956b,
1960). When C. lagopus colonies were physically divided in half early in growth,
the two halves yielded similar fruit body biomass, whereas the two sides of an
intact colony could differ by as much as 10:1, implying that in the latter case the
‘minority’ half is exporting its nutrients to the ‘majority’ half (Madelin, 1956b).
Mycelia must have access to sufficient substrates before fruiting is possible.
Buller (1931, p. 165) discussed the requirement for a minimum amount of
mycelium to support a minimum fruit body in Coprinus sterquilinus, arguing that
one of the functions of hyphal fusions between (clonal) germlings is to ensure the
rapid formation of that minimal size mycelium encompassing a corresponding
minimum quantity of substrate. Obviously, the minimum quantity of substrate
required varies between species depending on size of the fruit bodies produced.
Fungi producing small fruit bodies are able to do so with only a small amount of
resource, e.g. minute Marasmius and Mycena species restricted to leaf petioles,
small portions of leaf lamina, beech cupules, etc. (Figure 2). A very large mycelial
domain is required to produce the large, perennial brackets of heart-rot fungi
(Rayner and Boddy, 1988). It was estimated that all of the nitrogen in 13.6 g of
wood would be required to supply 1 g of Ganoderma applanatum basidiome, and
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Fruit Bodies 87
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David Moore et al.88
36.1 g wood to supply 1 g of spores, based on mean nitrogen content of fruit
bodies (1.13%), spores (3.05%) and Betula sapwood (0.83%; Merrill and Cowling,
1966). Since fruit bodies are commonly 1 kg or more, and several grams of spores
are produced each year (Fomes fomentarius produced 1.115 g spores in 20 days
(Meyer, 1936)), a mycelium would need to draw upon the entire nitrogen content
of more than 14 kg wood.
Culture studies indicate that once the minimum substrate size is reached fruit
body distribution is governed by a flow of nutrients towards particular devel-
oping fruit bodies, rather than localised nutrient depletion or inhibition of de-
velopment. The generality of this interpretation is based on two consistent
observations. First, that many fruit body primordia are generally formed, but
only a comparatively small number of them develop into mature fruit bodies; but
if fruit body size is related to local nutrient supply, one would expect that all of
the primordia on a colony would develop into mature but small fruit bodies, each
using those quantities of materials which are available locally and adjusting its
size accordingly. Second, a crop consisting of several fruit bodies will often
develop as a group, so that any general inhibitory action is unlikely. The concept
that nutrients flow towards a favoured centre would permit several neighbouring
primordia to mature in a clump, while still withholding nutrients from unfa-
vourably situated primordia. Clearly, different species emphasise different
aspects of this physiology in their fruiting behaviour and some are character-
istically solitary, e.g. Phallus impudicus, while others are caespitose, e.g. Hypholoma
fasciculare and Psathyrella multipedata (Figure 2). Some Basidiomycota, notably
Corticiaceae, form fruit bodies over the entire resource surface that they have
access to, e.g. Vuilleminia comedens on branches in the canopy. Large, skin-like
fruit bodies of some Corticiaceae may form at individual sites, subsequently
coalescing on contact. Detail is, however, lacking as much less research has been
done on these species than on Agarics.
In vitro experiments consistently indicate a general correlation between nu-
trient exhaustion of the medium and the onset of multicellular morphogenesis;
however, reproduction is not an alternative to vegetative hyphal growth but an
aspect of the differentiation of vegetative hyphae. Continued growth of the
vegetative mycelium is necessary to provide sustenance to its developing fruit
bodies. Correlation of fruiting with nutrient exhaustion of the medium does not
mean that development is prompted by a mycelium that is starving, because the
mycelium has accumulated nutrient reserves. Further, the timing of fruiting and
the amount of biomass that a fungus commits to fruiting varies with life history
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Figure 2 Some Fruit Bodies of Saprotrophic Basidiomycota, illustrating a Range of Sizes and
Resources: (a) the Solitary Macrolepiota rhacodes with a Coin Size Marker (20 mm Diameter);
(b) a Fruit Body of Marasmius setosus with the Same Coin Size Marker; (c) even Smaller
Marasmius Specimen on the Petiole of a Beech Cupule; (d) Collybia peronata on a Pine Cone;
(e) the Decidedly Caespitose Psathyrella multipedata; (f) Terence Ingold Posing with Fomes
fomentarius on a Beech Tree in Knole Park, Sevenoaks, Kent, 1969 (see Ingold, 2002).
Photographs (a)–(e) by David Moore of Specimens Collected by Members of the Mid-
Yorkshire Fungus Group at Harlow Carr Gardens.
Fruit Bodies 89
strategy (Cooke and Rayner, 1984; Rayner and Boddy, 1988; Chapter 11). Rapid
and extensive commitment of mycelial biomass is an R-selected (ruderal) char-
acteristic, typical of fungi that rapidly dominate following disturbance. Such
fungi are usually not combative and are often rapidly replaced by later arriving,
more combative species. They, therefore, must commit to reproduction before
they are killed and replaced. By contrast, slower and intermittent commitment to
reproduction is characteristic of fungi in stressful environments and/or that are
combative, dominating middle stages of community development. Laboratory
studies have largely employed species, e.g. Coprinopsis spp., Pleurotus spp.
and Schizophyllum commune, that fruit readily in culture, which is a ruderal
characteristic; thus, we must be cautious in extrapolating to fungi with other life
history strategies.
As we have discussed above, only preconditioned mycelium is capable of
undergoing morphogenesis. The preconditioned mycelium must be beyond a
particular minimum size, perhaps be of a particular minimum age, and the
underlying nature of both these preconditions is that the mycelium has been able
to accumulate sufficient supplies of reserve materials to support development of
the minimum reproductive structure. For some fungi, exhaustion of a particular
metabolite from the medium or substrate may be a signal that prompts morpho-
genesis in a mycelium that is not starving, but is healthy and well provisioned.
Exhaustion of one or more constituents of the medium changes the balance of
nutrient flow. If the medium is no longer fully supportive, the requirements of
active hyphal growth can no longer be met by import from outside the hyphae
and the balance must shift from ‘reserve material accumulation’ to ‘reserve
material mobilisation’. That change from balanced growth to growth under lim-
itation in external nutrient supply is what signals the onset of morphogenesis.
Cellular differentiation leading to fruit body morphogenesis is an expression of
unbalanced growth which is precipitated by one or more changes in the balance
of metabolism, and itself causes further cycles in which cellular components are
re-allocated. Even though nutritional dependence on the external substrate may
still be demonstrated, the emphasis shifts towards intramycelial regulation.
While this metabolic change is proceeding there is a change in the behaviour
of hyphal branches. For some branches, negative autotropism becomes positive
autotropism, so that neighbouring hyphae, often those of the surface or more
aerial parts of the mycelium, can interact. They form centres of rapid but self-
restricting growth and branching which become the hyphal aggregates or
mycelial tufts, perhaps 100–200 mm in diameter, that are the ‘initials’ of the
reproductive structure the organism can produce. Frequently, and especially in
culture, these aggregates are formed in great number over the whole surface of
the colony. As supplies of nutrients in the medium approach exhaustion repres-
sion of the morphogenesis of these hyphal aggregates is lifted and they proceed
to develop further. As mentioned above, only a small number of the first-formed
hyphal aggregates usually undergo further development and these become the
focus for translocation of nutrients, mobilised from the stores in other parts of the
colony and transported through the hyphal network to the developing repro-
ductive structures.
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David Moore et al.90
Illumination may be required, either to promote further morphogenesis or to
direct development into one of a small number of morphogenetic pathways (see
below). Particular temperatures may also be required for particular pathways of
development. Development usually proceeds in a series of steps that may be
coordinated by environmental cues (illumination, temperature, atmosphere) and
often involve sweeping re-allocation of cellular components. Within the young
fruit body, therefore, new accumulations of ‘stored’ nutrients arise, and there
may be a number of these accumulation–mobilisation–translocation–accumula-
tion cycles during the development of the reproductive structure.
2.4 Non-Nutritional Environmental Variables
As well as carbon and nitrogen nutrition, discussed above, many more environ-
mental variables affect fruit body initiation and development (reviewed by
Jennings, 1995; Moore, 1998a; Scrase and Elliott, 1998; Ku
¨es and Liu, 2000). Such
is the bulk of the literature that we can do little more here than list the major
observations.
As the above discussions of metabolism imply, fruit body development
requires oxidative metabolism (glycolysis and TCA cycle activity are often am-
plified) and good aeration is, not surprisingly, associated with successful fruiting.
This means not only oxygen but also various volatile metabolites including car-
bon dioxide. Elevated carbon dioxide concentrations can suppress basidiome
initiation in S. commune (Raudaskoski and Salonen, 1984). In Agaricus, increased
elongation of the stem occurs with elevated CO
2
, accumulated naturally from
respiration, whereas cap and gills expand and spores mature more rapidly when
CO
2
is removed (Turner, 1977). It has been argued that the morphogenetic effect
on maturation of the fruit body may have ecological advantage: CO
2
-enhanced
elongation of the stem would raise the gills away from the surface of the sub-
stratum where the concentration of CO
2
might be expected to be higher than in
the wider atmosphere because of the respiratory activity of microorganisms in
the casing soil (Turner, 1977).
High CO
2
levels promote formation of long hyphal compartments in S. com-
mune. It has been argued (Raudaskoski and Salonen, 1984) that a wood decom-
poser like S. commune is likely to experience elevated CO
2
within the wood as
respiratory CO
2
accumulates. Mycelium that reaches the surface of the wood,
however, will be exposed to CO
2
reduced to the atmospheric normal. Such
mycelium will be able to form the shortened cells and more compact branching
habit, and be predisposed to fruit body formation.
Light has diverse effects on formation of reproductive structures in different
basidiomycetes, increasing or decreasing their number, affecting their develop-
ment or determining whether or not they are produced (Carlile, 1970; Elliott,
1994). In general, the most effective parts of the spectrum are the near-ultraviolet
and blue wavelengths, typical of the shaded and litter-covered forest floor. There
are indications that the photoreceptor involved in fruit body morphogenesis may
be membrane bound. In some fungi levels of intermediary metabolites and
coenzymes, and activities of several enzymes respond very rapidly to changes in
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Fruit Bodies 91
illumination. The vegetative mycelia of many Ascomycota require exposure to
light before they will produce fruit bodies and/or asexual spores, and show
specificity not only for particular wavelengths but also for a particular dosage of
light radiation. In some Basidiomycota, sequential light exposures are respon-
sible for initiating and programming fruit body morphogenesis, and periods of
darkness between illumination events are important. Again, blue (400–520 nm) to
near-ultraviolet (320–400 nm) light is the most effective and the work suggests
that at least two photosensitive systems operate in fungi, one stimulated by near-
ultraviolet and the other by blue light. Because their absorption spectra parallel
the action spectra of the blue light photoresponses, carotenes and flavins appear
to be the best candidates for photoreceptors.
Production of fruit bodies in vitro typically occurs over a more restricted range
of temperature than that which will support mycelial growth. Optimum temper-
atures for fruit body production are generally lower than those most favourable
for mycelial growth. In Basidiomycota most information relates to species
adopted as laboratory models or for commercial cultivation. Cultivated species
frequently need a temperature downshift (by 5–101C) and lower CO
2
concentra-
tions for fruiting, e.g. A. bisporus,C. cinereus,Flammulina velutipes,Kuehneromyces
mutabilis,Lentinula edodes,Pholiota nameko,Pleurotus ostreatus,Stropharia rugosa-
annulata and V. volvacea (Chang and Hayes, 1978; Stamets, 1993). This list includes
compost-grown fungi as well as some wood-chip/straw and log-grown wood
decomposers, and is not unrepresentative of the wider community of
saprotrophic fungi, so it may be that most Basidiomycota require a temperature
downshift. A prolonged downshift is not always required; thus, fruit body
initiation in F. velutipes, which fruits in nature during late autumn to spring,
occurs at a continuous regime of 201C or following 12 h at 151C (Kinugawa
and Furukawa, 1965). Interestingly, the optimum temperature for both mycelial
growth and production of fruit body initials by A. bisporus is 241C (Flegg,
1972, 1978a, 1978b). However, temperature downshift is required for further
development of initials beyond a cap diameter of 2 mm.The fruit bodies develop
normally when the temperature is lowered to 161C. So, as with the reaction to
nitrogen sources mentioned above, the implication is that formation of fruit body
initials/primordia is an aspect of mycelial growth, but their proper development
requires a further morphogenetic switch. It is tempting to conclude that these
in vitro responses reflect the organism’s natural response to seasonal changes.
Relative humidity (RH) affects fruit body initiation. Relatively high humidity
is usually conducive to initiation of fruiting (Stamets, 1993), though it prevents
initiation in Polyporus ciliatus (Plunkett, 1956). The water content of the resource
may be even more critical. There is a balance between too high a water content
that reduces aeration and too low a water potential that provides insufficient
water for development (Scrase and Elliott, 1998; Ohga, 1999a; Kashangura et al.,
2006). There is variability between strains; Pleurotus sajor-caju was able to produce
primordia at 2.5 MPa but none at 3.5 MPa even though they were able to grow
under these xeric conditions (Kashangura et al., 2006). pH can affect fruit body
development, being optimal for several species at 6–7 (Ku
¨es and Liu, 2000), but
pH 4 for L. edodes (Ohga, 1999b).
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David Moore et al.92
Physical constraints influence fruit body formation in vitro. Sexual reproduc-
tion is often initiated when the growing mycelium reaches an obstacle such as the
edge of the dish or barriers placed onto the surface of the medium (the ‘edge
effect’ or ‘check to growth’). Reproductive structures often arise when mycelial
growth had been arrested, by either physical or chemical means (Moore, 1998a).
A physical barrier is not absolutely necessary for the ‘edge effect’, rather the
important determining factor is the disturbance in metabolism which results
from either encountering the edge of the dish or a major change in nutritional
value of the substrate. Thus, different sorts of barrier and different sorts of
medium transition are able to disturb the progress of metabolism sufficiently to
initiate fruit body formation. The same applies to physical injury to the
mycelium, which can stimulate fruit body formation (Leslie and Leonard,
1979a). Fruiting response to mechanical injury in S. commune is determined by at
least four genes (Leslie and Leonard, 1979a, 1979b), showing that a number of
different parallel routes lead to fruit body formation.
Inter- and intraspecific interactions can stimulate reproductive development.
In interactions with other fungi this is at least partly a result of damage to
vegetative hyphae (Rayner and Boddy, 1988). Many A. bisporus strains fruit only
when associated with bacteria, e.g. pseudomonads, apparently not due to pro-
duction of stimulatory compounds but to removal of inhibitory compounds (De
Groot et al., 1998). When competing with C. cinereus in agar culture, C. congregatus
fruited from a much smaller resource volume than when growing alone (Schmit,
1999). In contrast, interactions can result in a fungus being confined to territory,
e.g. a decay column in wood, that is too small to support fruit body production
by that species. Fruit bodies are assembled from contributions of a number of
cooperating hyphal systems, usually of the same individual. Hyphal interactions
are controlled by the somatic and mating incompatibility systems (Chiu and
Moore, 1999) that maintain mycelial individuality. Fruit bodies of somatically
compatible Basidiomycota can fuse when the fruit bodies develop in extremely
close proximity, as is commonly seen when resupinate fruit bodies meet on
wood, and also with stipitate basidiomata, e.g. a fused cap with three stems of
Boletus (Xerocomus) chrysenteron in Kibby (2006). However, hyphal cooperation is
so fundamental that it can even lead to the formation of chimeric fruit bodies.
Mixed cultures of two genetically different heterokaryons can produce basidio-
mata comprising both dikaryons, as seen with P. nameko (Babasaki et al., 2003).
Even more extreme is the case of fruit bodies of Coprinus consisting of two
different species, C. miser and C. pellucidus (Kemp, 1977). The hymenium com-
prised a mixed population of basidia bearing the distinctive spores of the two
species but the chimera extended throughout the fruit body as both species could
be recovered by outgrowth from stem segments. All of these features can be
interpreted as aspects of the tolerance of imprecision in fungal morphogenesis
which has been discussed elsewhere (Moore, 1998a, 1998b, 2005; Moore et al.,
1998).
Once fruit bodies have been produced environment, particularly temperature
and RH, can affect spore production. For example, in the field spore production by
Hericium erinaceus is highest at about midday reflecting diurnal temperature and
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Fruit Bodies 93
RH (McCracken, 1970). In the laboratory, at 85–95% RH, spore production in-
creased from a minimum at 01C to a maximum at 24–271C, and ceased at 31–331C.
At 201C, sporulation was greater at 30% RH than at 90% RH (McCracken, 1970).
2.5 Fruiting in the Natural Environment
It is well known that the majority of Basidiomycota fruit in autumn, following
mycelial growth and decomposer activity in spring and summer. Temperature
and rainfall are considered to be the two main factors affecting productivity
(Salerni et al., 2002). In a 21-year fruit body survey of a forest plot in Switzerland,
there was considerable variation between years in species richness and produc-
tivity, only litter decomposing saprotrophs, Collybia butyracea var. asema and
C. dryophila, appearing in all years (Straatsma et al., 2001). Appearance of fruit
bodies was correlated with July and August temperatures, an increase of 11C
resulting in a delay of fruiting by saprotrophs of 7 days. In contrast, fruit body
productivity was correlated with precipitation from June to October (Straatsma
et al., 2001), and similar relationships have also been found in Britain and Sweden
(Wilkins and Harris, 1946; Wasterlund and Ingelog, 1981).
In a 3-year study of Mediterranean oak forests, there was no evidence for
influence of temperature on fruit body species diversity or productivity by most
saprotrophs, though there was strong positive correlation between species
diversity of wood decay fungi and maximum temperature, and with spring and
summer rainfall (Salerni et al., 2002). Temperature and rainfall in the 5 days prior
to surveying seemed to have little effect on fruiting, but did so between 10 and 30
days prior to survey.
Climate change has resulted in phenological changes in plants, insects and
birds (Parmesan and Yohe, 2003), and this has recently been shown to be the case
for fungi (Gange et al., 2007). Analysis of a data set of fruiting records of 200
species of decomposer Basidiomycota in Wiltshire, UK, each of which had been
recorded over more than 20 years during 1950–2005, revealed that mean first
fruiting date averaged across all species is now significantly earlier, while mean
last fruiting date is now significantly later (Figure 3; A.C. Gange, E.G. Gange, T.H.
Sparks and L. Boddy, unpublished data). Thus, the fruiting season has been
extended since the 1970s. Not all species fruit earlier (47% show an advance-
ment), or produce fruit bodies later into the year (55% continue fruiting later) but
of those saprotrophic Basidiomycota that showed significantly earlier fruiting
dates (n¼94), the average advancement was 7.9 days per decade, while for those
with significantly later last fruiting dates (n¼110) the delay was 7.2 days per
decade. The response differs depending on habitat type: 13% of grassland species
fruiting earlier, 48% having later last fruiting; 53% of wood decay fungi fruited
earlier, with 20% having later last fruiting. There was a significant relationship
between mean fruiting date of those species that normally fruit early in the
season (September) and late summer temperature and rainfall (Figure 4). Local
July and August mean temperatures have significantly increased (July, Po0.05;
August, Po0.01), while rainfall has decreased, though less markedly, over the
56 years of the survey.
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David Moore et al.94
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Figure 4 Relationship between Mean Fruiting Date of Saprotrophic Basidiomycota Species
that Normally Fruit Early in the Season (September) and (a) August Temperature (R
2
¼0.299,
F(1,54) ¼23.056, P¼0.007) and (b) August Rainfall (R
2
¼0.126, F(1,54) ¼7.790, P¼0.000) (A.C.
Gange, E.G. Gange, T.H. Sparks and L. Boddy, Unpublished Data) AU :6
.
Figure 3 Mean First Fruiting Date (Lower Line) and Mean Last Fruiting Date (Upper Line) for
200 Saprotrophic Basidiomycota over 56 Years. Splitting the Data into Two Equal (28 Year)
Periods Reveals no Trend in the First Half (P¼0.97) but a Highly Significant Trend (Po0.001)
in the Second Half (A.C. Gange, E.G. Gange, T.H. Sparks and L. Boddy, Unpublished Data).
Fruit Bodies 95
As well as changes to autumn fruiting patterns, significant numbers of species
that previously only fruited in autumn now also fruit in spring (Figure 5). Since
mycelia must be active in uptake of water, nutrients and energy sources before
fruit bodies can be produced this suggests that these fungi may now be more
active in winter and spring than they were in the past.
Other aspects of the environment can also influence fruiting by affecting
microclimate (e.g. ground vegetation and logging waste), providing additional
resources or inhibitory compounds. For example, in managed forests: there was
lower fruit body biomass where Pteridium aquilinum was abundant; in dry years
Mycena species were more abundant in areas with logging waste, but in wet years
they were equally or more abundant in areas without logging waste; fruit body
biomass was negatively correlated with grass cover in dry autumns, but posi-
tively correlated in wet autumns (Wasterlund and Ingelog, 1981).
3. FRUIT BODY SURVIVAL
As well as the physical size of a fruit body, a significant feature in the ecology of
the organism is the length of time that the fruit body remains sufficiently intact to
distribute spores. This varies from a few days or weeks for fleshy fungi to several
years for perennial brackets, longevity of the latter being associated with struc-
tural physical characteristics and production of chemicals that inhibit inverte-
brate feeding or are toxic to them (Kahlos et al., 1994 AU :1; Stadler and Sterner, 1998).
There appears only to be one detailed study of the lifespan of an agaric, an
analysis of the fruit bodies of A. bisporus grown in an experimental mushroom
farm over 36 days (Umar and Van Griensven, 1997). The fruit bodies remained
healthy for 18 days before localised cytological indications of senescence became
evident (nuclear and cytoplasmic lysis, permeable cytoplasmic membranes and
structural changes to the cell wall). Cells of the fruit body collapsed irregularly
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Figure 5 The Proportion of Saprotrophic Basidiomycota in Different Habitat Groups that,
Before 1975, were not Recorded as Fruiting in Spring, but after This Time did so in at least 1
Year (A.C. Gange, E.G. Gange, T.H. Sparks and L. Boddy, Unpublished Data).
David Moore et al.96
and the remnants of the lysed cells aggregated around and between the remain-
ing living hyphal cells. Most of the stem hyphae became empty cylinders. After
36 days, electron microscopy showed that most of the cells throughout the fruit
body were severely degenerated and malformed, yet a number of basidia and
subhymenial cells remained intact and alive even at 36 days. Interestingly, when
mushrooms were cultivated using conventional commercial farming procedures,
50% of the fruit bodies were infected by Trichoderma harzianum and/or
Pseudomonas tolaasi by 18 days. All such fruit bodies died at 24 days due to
generalised severe bacterial and fungal infections leading to tissue necrosis and
decay of the caps and stems.
Observations of a wild troop of Clitocybe nebularis in a garden in Stockport,
Cheshire, began on 21 October 2006, at which time the fruit bodies were young,
but close to maturity (5 cm diameter), and continued for 29 days (Figure 6). By 19
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Figure 6 Life and Death of Clitocybe nebularis Fruit Bodies in a Suburban Garden in
Stockport, Autumn 2006. Observations began on 21 October and Continued for 29 days to 19
November. Troops of Fruit Bodies of Coprinus micaceus Emerged, Matured and Decayed 26
October and November 1 (the Latter are Illustrated). Some Disturbance and Grazing
(Squirrels?) was Evident on 10 November, and Collapsed Fruit Bodies by 18 November.
Fruit Bodies 97
November most of the fruit bodies were beginning to collapse. These basidio-
mata of C. nebularis were still actively releasing spores on 7/8 and 12/13
November, clearly indicating that agarics with large fruit bodies can distribute
spores for 3–4 weeks, though viability was not tested. During the observation of
C. nebularis, two troops of fruit bodies of Coprinus micaceus emerged, matured and
decayed (26 October and 1 November), illustrating the alternative (R-selected)
strategy of rapid production of short-lived fruit bodies.
The longevity of fruit bodies is obviously important for dispersal, but so also
is the period over which spores are actively produced and released, and the
viability/germinability of spores produced at different times. While some species
retain high germinability of spores produced over several weeks, e.g. Poria tenuis
and Trametes hispida, with others there is a decline, e.g. germinability of Poria
placenta and Gloeophyllum trabeum declined from 494 to 19 and 44%, respectively,
5 weeks after fruiting was initiated in culture (Schmidt and French, 1983).
4. PRINCIPLES OF FUNGAL DEVELOPMENTAL BIOLOGY
Numerous observations show that all aspects of the environment can influence
the production and development of fungal fruit bodies. To understand how
this occurs we need to formalise fruit body development sufficiently to allow
recognition of the decisive steps that are open to influence, and we must also
identify the molecular controls that normally regulate those steps.
4.1 Underlying Principles
Three generalisations can be extracted from the past century of observations on
fruiting physiology. First, the organism internalises nutrients rapidly to gain reg-
ulatory control over nutrient access and distribution. By so doing the vegetative
mycelium becomes competent to produce multicellular structures like fruit bodies.
Second, factors that promote fruiting, whether physical or chemical, seem to work
by disturbing the normal progress of cellular metabolism. It is the disturbance itself
that is the effective factor, overcoming some block to progress and inducing the next
stage to proceed. Consequently, parallel pathways cover some stages of fruit body
development and for these stages different factors seem to be interchangeable (e.g.
a particular nutritional state may replace a particular illumination requirement).
Third, even relatively simple developmental pathways can be subdivided into
stages (at least, initiation, development and maturation) and there seems to be a
need for successive signals (successive metabolic disturbances) to maintain
progress of the developmental process. Each stage involves change in hyphal
behaviour and physiology, taking the tissue to a higher order of differentiation.
4.2 Modelling Hyphal Growth and Fruit Body Formation
Hyphal growth is well suited to mathematical modelling, and the recent neigh-
bour-sensing model brings together the basic essentials of hyphal growth kinetics
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David Moore et al.98
into a vector-based mathematical model that ‘grows’ a life-like virtual mycelium
(or ‘cyberfungus’) on the user’s computer monitor (Mes
ˇkauskas et al., 2004a,
2004b; Moore et al., 2006). The program has been used in a series of experiments
(Mes
ˇkauskas et al., 2004a, 2004b) to show that complex fungal fruit body shapes
can be simulated by applying the same regulatory functions to all of the growth
points active in a structure at any specific time. No global control of fruit body
geometry is necessary; rather, the shape of the fruit body emerges as the entire
population of hyphal tips respond together, in the same way, to the same signals.
These computer simulations thus demonstrate that because of the kinetics of
hyphal tip growth, very little regulation of cell-to-cell interaction is required to
generate fungal fruit body structures. The program includes parameters that can
be used to mimic the effects of cell-to-cell signalling and environmental variables.
These give the experimenter the opportunity to study the effects of such variables
on fungal growth in silico.
4.3 Data Mining Fungal Genomes
The notion that control mechanisms of fungal multicellular developmental
biology are probably very different from those known in animals and plants that
emerges from the work described so far is supported by sequence searches of
genomic databases. The unique cell biology of filamentous fungi has clearly
caused control of their multicellular development to evolve in a radically differ-
ent fashion from that in animals and plants. There are no Wnt,Hedgehog,Notch,
TGF,p53,SINA or NAM sequences in fungi (Moore et al., 2005; Moore and
Mes
ˇkauskas, 2006), but there are presumably analogous or homologous processes
in fungal multicellular structures that need to be regulated.
Unfortunately, the demonstration that developmental control sequences of
animals and plants lack fungal homologues leaves us knowing nothing about the
molecules that do govern multicellular development in fungi. Yet these are the
molecules and mechanisms that generate fungal fruit bodies. The molecular
control elements of development are the things with which the environment
interacts to cause its effects. While we remain ignorant of the basic control proc-
esses of fungal developmental biology we will also remain ignorant of the way
environment impacts on fungal biology.
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... Como se ha mencionado en varios estudios, (Boddy & Heilmann-Clausen, 2008) altas concentraciones de humedad en el suelo y en la madera no favorecen la producción de cuerpos fructíferos. Esto se comprueba al observar las pocas especies que formaron cuerpos fructíferos durante el 2008, año que además de presentarse la precipitación más alta (4 000 mm), también se dieron las temperaturas más bajas, dos factores que pueden influir en el comportamiento de los hongos (Moore, Gange, Gange, & Boddy, 2008). En el 2009 las precipitaciones disminuyeron (2 000 mm), y la aparición de cuerpos fructíferos de diferentes especies fue baja, pero superior a la del 2008. ...
... No se puede descartar; sin embargo, la posibilidad de que algunas de las especies estuvieran presentes en el bosque primario, pero que no fructificaran durante las visitas efectuadas a este bosque, como ha sido mencionado en otros trabajos (Moore et al., 2008;Mueller et al., 2006;Nordén & Paltto, 2001). ...
... La humedad y la temperatura son dos de los factores que se han relacionado con la aparición de fructificaciones de los hongos (Boddy & Heilmann-Clausen, 2008;Moore et al., 2008;Mueller et al., 2006;Pyle & Brown, 1999). La zona de estudio se caracteriza por precipitaciones y temperaturas anuales de alrededor de 2 700 mm y de 10-14°C, respectivamente y una corta estación seca. ...
... Por outro lado, vários estudos propõem que a fenologia é um indicador sensível de mudanças climáticas recentes, no entanto, a resposta fenológica a essas mudanças não é compartilhada pelos membros de uma comunidade. Para os fungos, os eventos fenológicos incluem a formação de basidiomas e o crescimento vegetativo das hifas (Moore et al., 2008). Este último evento dificulta a realização de estudos fenológicos com fungos, uma vez que são geralmente encontrados sob o solo ou outros substratos, sendo os basidiomas geralmente utilizados para esse fim (Bünten et al., 2012). ...
Chapter
Os Fungos Micorrízicos Arbusculares (FMA), famosos pela formação de simbiose com raízes da maioria das famílias de plantas, são comuns em quase todos os ecossistemas terrestres. Entretanto, os ambientes aquáticos têm sido pouco investigados principalmente quanto a diversidade desses microrganismos benéficos. Apesar da notória negligência nas pesquisas sobre diversidade de FMA em áreas alagadas, há informações que requerem atenção. Dessa forma, o objetivo do trabalho foi inventariar espécies de FMA em 7 ambientes aquáticos lênticos, oligotróficos do estado do Rio Grande do Norte, Brasil. Para isso, sedimento rizosféricos de 10 famílias de ma- crófitas aquáticas foram coletados, os glomerosporos extraídos por peneiramento úmido e centrifugação em água e sacarose 50%, montados em lâminas para microscopia, identificados e quantificados. Surpreendentemente, 105 espécies foram observadas distribuídas em 5 ordens, 11 famílias e 21 gêneros. A maior frequência foi das famílias Acaulosporaceae e Glomeraceae, com os gêneros Acaulospora e Glomus como mais representativos. Ambispora appendicula foi a única espécie comum às 7 lagoas. Os índices de diversidade foram elevados, exceto para a Lagoa do Boqueirão e Lagoa Azul, que apresentaram maiores índices de dominância. Os hospedeiros hidrófitos com maior riqueza foram Cyperaceae e Lentibulariaceae. O número de táxons encontrado é elevado, assim como a esporulação, revelando o potencial dos ecossistemas aquáticos em abrigar ampla riqueza de FMA.
... According to Martínez de Aragón et al. (2007), fungi require in the first stage of their development a minimum of precipitation and then appropriate temperatures for the appearance of sporophores. Similar data have been reported by many authors (El-Assfouri et al. 2005, Moore et al. 2008, Gévry and Villeneuve 2009and Bâ et al. 2011. ...
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... However, our sampling was performed when the ring effects in the vegetation were most visible, similar to other studies on fairy rings (Marí et al., 2020;Yang, Li, et al., 2018;Zotti et al., 2020). Finally, nutrient requirements of fairy ring fungi in La Bertolina could be lower than those in other locations, due to the absence of fruiting bodies, which normally require a high nutrient investment (Gençcelep et al., 2009;Moore et al., 2008). ...
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Shiitake mushroom (Lentinula edodes (Berk.) Sing.) is widely cultivated in China, Japan, Korea and many other Asian countries. It is one of the most extensively grown and consumed edible fungi in the world, with an exceptional high agricultural yield. Taxonomically, L. edodes belongs to the phylum Basideomycotina and family Agaricaceae. It is broadly distributed in the wild, mainly in the subtropical to temperate regions of the northern hemisphere. Due to its excellent taste qualities, high nutritional value, and medicinal properties, L. edodes is of great importance for the food industry and has medicinal applications. Various active medical ingredients such as lentinan, lentin, lectin and eritadenin, have been isolated from L. edodes culture media, fruiting bodies or mycelium. Enzymes, such as laccase, produced by L. edodes, have potential for industrial applications related to paper production (biopulping), residue treatment and improvement in the digestibility of animal rations. Like for other edible fungi cultivation, the raw materials for shiitake mushroom cultivation mostly constitute agricultural waste such as sawdust, straw and cottonseed husk. Moreover, the waste from the shiitake mushroom cultivation itself can be further used as a bio-organic fertilizer, greatly contributing to the process of crop rotation. Due to the changing natural environment along with the continuous improvement of living standards, mushroom cultivation is facing constant challenges. In order to obtain shiitake mushroom strains adapted to different climatic conditions, different cultivation methods and processing practices, breeders continuously screen for new shiitake varieties implementing diverse methods. In this chapter, we present an overview of the origin, distribution, taxonomic position, genetic characteristics, cultivation patterns and history of shiitake mushroom breeding by traditional and modern breeding methods in China.
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Mushroom cultivation presents an economically important biotechnological industry that has markedly expanded all over the world in the past few decades. Mushrooms serve as delicacies for human consumption and as nutriceuticals, as "food that also cures". Mushrooms, the fruiting bodies of basidiomycetous fungi, contain substances of various kinds that are highly valued as medicines, flavourings and perfumes. Nevertheless, the biological potential of mushrooms is probably far from exploited. A major problem up to now is that only a few species can be induced to fruit in culture. Our current knowledge on the biological processes of fruiting body initiation and development is limited and arises mostly from studies of selected model organisms that are accessible to molecular genetics. A better understanding of the developmental processes underlying fruiting in. these model organisms is expected to help mushroom cultivation of other basidiomycetes in the future.
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Some effects of light and temperature on fruiting of Coprinus lagopus in pure culture are described. Fruiting, which in darkness did not commence until about the 15th day, was accelerated by continuous light or by brief exposures to light between the 7th and 13th days of incubation. Very small exposures sufficed, provided they were of a wavelength no longer than that of green light, the response being restricted to the area of mycelium actually exposed. In both light and dark a temperature near 25° C. was optimal for growth and fruiting.