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Agaricus Morphology and Life Cycle

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650 views33 pages

Agaricus Morphology and Life Cycle

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j35476560
Copyright
© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
Available Formats
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Morphology and life cycle of Agaricus

Discipline Courses-I
Semester-II
Paper III: Mycology and Phytopathology
Unit-VI
Lesson: Morphology and life cycle of Agaricus
Lesson Developer: Geeta Sharma
College/Department: Shivaji College, University of Delhi

Institute of Lifelong Learning, University of Delhi


Morphology and life cycle of Agaricus

Table of Contents

Chapter: Agaricus

 Introduction
 Habit and Habitat
 Somatic structure (Vegetative thallus)
 Vegetative cell
 Reproduction
 Asexual
 Sexual
 Summary
 Exercise/ Practice
 Glossary
 References/ Bibliography/ Further Reading

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Morphology and life cycle of Agaricus

Introduction
The genus Agaricus comprises a group of 200 species which are heterotrophic, true
nucleated, multicellular, macroscopic and fleshy in nature. This group includes fungi whose
fruiting bodies are commonly known as mushrooms. The umbrella shaped mushrooms
(which are edible), toadstools (which are nonedible or poisonous) beautify this earth with
their presence in almost all seasons except the extreme summers and winters. The fruiting
bodies of some species appear in early spring and disappear in summers, others appear
only in rainy season and some appear sporadically whenever moisture is available. Hence
the moist season (rainy weather) is the best season to explore, observe and collect the
most beautiful, colourful (earthy flowers) fruiting bodies of mushrooms in hills as well as in
plains.

The word mykes (from which the term Mycology is derived) means mushrooms and
etymologically, mycology is the study of mushrooms. This group has been extensively
studied in detail because man collected the fruiting bodies of mushrooms from diverse
habitats to consume them as food, medicines etc.

Mature spores

Developing spores
Gills

Pileus

Veil

Stipe

Figure: The image shows different stages of development of basidiocarps and basidiospores

Source: [Link]
[Link]

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Morphology and life cycle of Agaricus

Systematic position

Mycologists have since ancient times classified mushrooms under Basidiomycetes which in
turn have been grouped in different categories of different ranks at different levels
based on specific characters like the type of mycelium ,nutrition , occurrence,
reproduction ,life cycle ,their molecular and phylogenetic relationship with other fungal
groups .Some specific systems recognized and followed are as followed:

G.C. Ainsworth (1973) :

Division : Eumycota

Subdivision : Basidiomycotina(Basidiomycetes) .

Class : Hymenomycetes .

Alexopoulos , C.J. and Mims, C. W.(1979) :

Super Kingdom : Eukaryonta .

Kingdom : Myceteae .

Division : Amastigomycota .

Subdivision : Basidiomycotina .

Class : Basidiomycetes .

Alexopoulos C. J. Mims , C. W. and Blackwell, M.(1996) :

Kingdom : Fungi

Phylum : Basidiomycota

Class : Basidiomycetes

Order : Agaricales

Webster , J . and Weber, R.W.S. (2007) :

Kingdom : Fungi(Eumycota)

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Morphology and life cycle of Agaricus

Phylum : Basidiomycota

Class : Homobasidiomycetes

Order : Agaricales

Family : Agaricaceae

Genus: Agaricus

Habit and Habitat

Agaricus is a multicellular, macroscopic, mostly saprophytic corticolous, parasitic and


mycorrhizal in nature. Agarics are cosmopolitan in nature and occur in a wide variety of
habitats ranging from the arctic to the tropics. Most of the species do show the
preference to the diverse kind of substrata for example :-

 In upland woody forests on decaying wood (lignicolous), decaying fallen leaves


(foliicolous) eg. A . lanipes, A. langei, on moist soils (terrestrial), compost and
dung (coprophyllous) eg . A. bisporus, A. bitorquis .

Figure: Agaricus lanipes a wood decayer

Source:[Link]
.JPG

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Morphology and life cycle of Agaricus

 In open areas like gardens, lawns, grassy lands and pastures, eg . A.


fuscofibrillosus, [Link], A. campestris, A. macrosporus.

Figure: A. fuscofibrillosus

Source:[Link]
[Link]

 The mycorrhizal forms occur with certain type of vegetation eg. Ectomycorrhizal
association with conifers—eg. Paxillus (gilled) ,Gomphidius (gilled) etc .

Figure: Gomphidius

Source:[Link]
us_131007.jpg

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Morphology and life cycle of Agaricus

 On the body of the basidiocarps of other mushrooms (fungicolous).

Figure: Volvariella surrecta mycoparasite on Clitocybe nebularis

Somatic structure (vegetative thallus)

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Morphology and life cycle of Agaricus

The fruiting bodies (sporophores) of mushrooms as seen on the surface of


substratum by the naked eye do not exhibit the complete fungal thallus but these
are the sexual reproductive bodies bearing meiotic basidiospores on the part of
mycelium .

The somatic structure or vegetative thallus of Agaricus consists of the mycelium


(mass of hyphae) which is septate, thin walled, hyaline typically a basidiomycetous
type. Here it arises as a primary mycelium from the monokaryotic basidiospores,
which later become dikaryotic and tertiary mycelium.

Figure: Agaricus campesteris: Diagrammatic repersentation of the anatomy of the


basidiocarp with three types of mycelia.

Source: [Link]

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Morphology and life cycle of Agaricus

The mycelium of Agaricus is of three kinds:

Primary mycelium :The monokaryotic, haploid, basidiospore of (+) and (-) strain
germinates immediately after dispersal on the substratum or on the moist soil
(at 10- 15 degree temperature, 80—90% relative humidity, and 5.8—6.o ph)
by absorbing mineral nutrients directly in soluble form to give rise a germling
or primary hyphal initial . This hyphal initial cell undergoes mitotic division to
form monokaryotic, uninucleated, haploid, hyaline, thinwalled primary mycelium
which has a very short life in nature.

Secondary mycelium: The primary hyphae or monokaryotic hyphae with


(+) or (-) strain haploid nuclei come in contact with each other to form
dikaryotic or hetrokaryotic mycelium by two methods : -

i. Somatogamy: The somatic cells ( vegetative cells) of primary or


monokaryotic hyphae of opposite srains (+ and - ) when they come
in contact with each other directly on the substratum , the cell wall
between the two cells ( + and- ) dissolves , fusion of two protoplasms
( plasmogamy) takes place, two nuclei (+and-) lie near each other to
form a dikaryotic, haploid cell (somatogamy) . This dikaryotic (n) cell
undergoes multiple mitotic divisions to form thin walled,
multicellular, branched secondary mycelium. Each cell of this mycelium
carry two genetically distinct (compatible) nuclei (+ and-) and is strictly
heterokaryotic, dikaryon. fig.

ii. Clamp connection: This method is very common to develop


heterokaryotic, secondary mycelium from primary mycelium of opposite
mating types. In this process the two compatible cells of opposite strain
(+&-) of monokaryotic hyphae come in contact with each other, the
cell wall between them dissolves, nuclear migration takes place .
The nucleus of this dikaryotic cell divides mitotically and the daughter
nucleus migrates in the adjuscent cell with the help of a characteristic
structure or a connection i.e. a short appressed hyphal branch which
bypasses the transverse septum and connects the two adjuscent
daughter cells laterally called Clamp ( or buckle) connection. It sis a
method which ensures that when a dikaryotic mycelium divides, each
cell of these hyphae will contain two genetically distinct nuclei.

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Morphology and life cycle of Agaricus

Figure: Formation of secondary mycelium through somatogamy and clamp connections

Source: Author

Secondary, dikaryotic or heterokaryotic mycelium is multicellular, thinwalled, branched,


cottony in appearance, the hyphal cells of which are binucleated or
multinucleated, haploid dikaryons.

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Morphology and life cycle of Agaricus

Tertiary mycelium : The dikaryotic, secondary mycelium carrying opposite


strain( +and-) haploid nuclei in the hyphal cells branch profusely by mitosis and
the hyphal branches anastomose with each other to form a complex tissue
of fruiting body (basidiocarp).

Vegetative or hyphal cell

The Agaricus hyphal cell consists of outer cell wall (chitin microfibrils and
glucan),inner plasma membrane (selectively permeable), true nucleus, haploid of +
and - strain or both, cytoplasm, mitochondria, ribosomes, endoplasmic reticulum,
golgi apparatus, small vacuoles and glycogen granules as reserve food material
etc.

 Structure of septum: The septum ( the intersecting wall between each


hyphal cell of septate hypha ) in each cell of monokaryotic and dikaryotic
hyphae of Agaricus is pierced by a narrow 0.1- 0. 2 ,um width which is
surrounded by a barrel called septal swelling. This kind of septal pore is
known as Dolipore septum. This pore is overarched with a perforated cap
i.e an extension of endoplasmic reticulum. This perforated cap is called
parenthesome or pore cap which has several pores in it from which
organelles like mitochondria can pass.

Figure: Dolipore septum (arrow)

Source: [Link]

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Morphology and life cycle of Agaricus

Reproduction
Asexual reproduction

The species of Agaricus undergo asexual reproduction very rarely. It produces two
types of asexual spores i.e. oidium (thin walled, fragmented spore), and
chlamydospores (thick walled, resting spore).

 Oideum or fragmented spore : Oidia or arthrospores are thin walled,


hyaline, fragmented, spores which are produced by the dissolution of middle
lamellar wall between the adjuscent cells on the aerial branches of monokaryotic
and dikaryotic hyphae. The monokaryotic oideum fuses with the hyphal cell
of the opposite strain to form the secondary heterokaryotic mycelium .The
oidia of dikaryotic type germinate directly to give rise heterokaryotic
mycelium eg. Coprinus lagopus. These type of spores are produced during
favourable conditions i.e temperature 10 -15Oc , relative humidity 80-90% and
ph 6-6.5 .

Figure: Asexual reproduction by oidia formation (rare)

Source: Author

 Chlamydospores or resting spores: It is a thick walled resting spore


or perennating spore (seed like) produced during unfavourable conditions
(drying of substratum, rise in temperature, and change in ph). The hyphal
cells loose extra moisture content, glycogen particles (reserve food) get
converted into fat droplets, cell wall becomes thickened by chitin deposition.
These spores produced in chains are initially light coloured but later
become dark brown, get separated from the rest of the parental mycelium

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Morphology and life cycle of Agaricus

and parennate in unfavourable conditions in the soil or on the substratum,


eg. Coprinus lagopus (Lewis 1961) and Volvariella volvacea (Chang and Yan
1971) . Chlamydospores during rainy season germinate by absorbing
moisture and minerals from the substratum at 10-20Oc , ph 5.8-6.0 and
give rise the young mycelium on the substratum.

Figure: Chlamydospores and their germination

Source:Author

Sexual reproduction

Agaricus does not have well defined sex organs or sexual gametes. The somatic
or vegetative cells of primary hyphae of opposite mating types (+&- strains )
function as sexual gametes. The two somatic cells of haploid primary hyphae
of opposite strains (+&-) come in contact with each other by a process of
somatogamy, oidization or by clamp connection and fusion of protoplasms (
plasmogamy) bring two compatible nuclei (+&-) near each other. Homothallism is
rare in agaricales eg. (Coprinus sterculinus). Most of the species exhibit either
unifactorial or bifactorial homothallism . The two significant steps that is
karyogamy (nuclear fusion of opposite mating +&- strains) and meiosis ( reduction
devision) take place in the basidial cells of fertile layer (hymenium) of fruiting
body i.e. basidiocarp . The basidiospores (meiotic ) are produced on the probasidial
cells exogenously at the abaxial(ventral) surface of the basidiocarp .

Basidiocarp

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Morphology and life cycle of Agaricus

Figure: Mature basidiocarp showing basal remnant of mycelium, stipe annulus and pileus.

Source: [Link]
II+revised+[Link]

The fruiting body or the basidiocarp formation is regulated or influenced by the


interaction of multiple environmental factors:

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Morphology and life cycle of Agaricus

Light: Diffused light .

Relative humidity : 80-90 % .

Temperature : 12-18’c .

pH of the substratum : 5.8-6.0 .

Mineral nutrients (ammonium salts etc.) and aeration.

These conditions favour the formation of fruiting bodies or the


basidiocarp on the substratum. The initial steps in fruiting body formation are :

1. An intricate hyphal lattice is formed by the interaction of hyphal


branches .

2. From this hyphal lattice the bunch of aerial hyphae appear


which eventually produce a round aggregation of tightly interwoven
hyphae to form a button or a bud like structure, creamish white in
colour called bud primordium ( first basidiocarp stage) on the
substratum.

3. The bud primordium grows in size and as it reaches 1.0 mm in


diameter , a presumptive stipe, hymenium (fertile layer) and terminal
pileus (cap) get differentiated simultaneously .

4. The elongation of stipe, expansion of terminal pileus in to upper


(adaxial)smooth and lower(abaxial)infolded gilled lamellar structure and
production of basidiospores occur quite rapidly and dramatically in
the life cycle of Agaricus .

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Morphology and life cycle of Agaricus

Figure: Diagrammatic representation of development of basidiocarp from pin head to


mature fruiting body.

Source: Author

In agaricales the three basic types of basidiocarp development have been


recognized(Singer 1975) :

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Morphology and life cycle of Agaricus

Hemiangiocarpous development:The hemiangiocarpous basidiocarp development is


characterized by the following facts:

 The hymenium or fertile layer is enclosed by tissues (tertiary mycelium) of


the basidiocarp even during early stages of its development .

 The margin of the pileus is connected to the basel stipe by a membrane


called inner veil .

 The hymenium remains enclosed till the terminal cap (pileus) expands and
matures .

 The veil tears from the margin of the pileus shortly before the spores
mature and remains attached to the stipe as a ring or annulus .

 ( In some species the veil tears in such a manner that it hangs down
from the cap(pileus) like a thin cobwebby curtain called Cortina and in
others the entire primordium is covered by a universal veil. This universal
veil breaks and leaves a cup shaped body around and bulbous base of
the stipe. The remains of the parts of universal veil are seen as scales on
the cap. These features help in classification and identification of many
species as this feature is evanescent in other species ).

Gymnocarpous and Pseudoangiocarpous development : In these two methods


the hymenial (fertile) layer is formed initially on the outside of the fruiting
(basidiocarp) body. In gymnocarpous development the hymenial tissue is not
covered and remains naked till maturity. In pseudoangiocarpous method the
hymenial tissue later becomes enclosed by the development of the outgrowth of
the incurving margin of pileus and the stipe. The hymenium remains enclosed till
the basidiocarp matures and the pileus expands to reveal the hymenial tissue
.In these methods the vestigial structures are not seen at the maturity of the
fruiting bodies (basidiocarp).

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Morphology and life cycle of Agaricus

Figure: Figure representing different types of basidiocarp development

Source: Author

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Morphology and life cycle of Agaricus

Structure of Basidiocarp:The basidiocarp of Agaricus which bears basidiospores


consists of the following tissues:

a) Hymenium: The fertile layer of anastomosed secondary hyphae is found


beneath the pileus (terminal cap). It lines in the gills or lamellae
exogenously which hang below in the ventral (abaxial) surface of the pileus.

b) Gills: The gills are the thin strips of tissues which radiate from the margin
of the pileus in towards the stipe. The structure and position of the inner
edge of gills towards stipe is a valuable and interesting feature in many
agaricales for taxonomists. The gills in some species are free from stalk
while in others they are attached directly to the stipe called adnate. The
decurrent gills are attached and run down the stipe for some distance.

c) Trama : The trama ( L. Trama-Woof) is the inner most tissue which lie in
the center of the gills. It is an important feature of taxonomic interest Singer
(1975). The central tissue (trama) is of two types:

 The trama consists of plectenchymatous tissue which is made up of


elongated hyphae.

 The trama tissue contains large, globose or oval cells called sphaerocysts
scattered along the elongated hyphal cells (eg. Russula and Lactarius ).
The trama is surrounded on both sides by the fertile layer (hymenium) in
which closely packed layer of basidia are interspersed with basidioles or
cystidia or both in some species . The cystidia are long and extend from
one gill to the next to keep the gills apart from each other (Coprinus) .

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Morphology and life cycle of Agaricus

A B

C D

Figure: Diagrammatic representation of a A. V.L.S. fruiting body; B. V.L.S. gills; C-D. Part
of gill magnified

Source: Author

Basidium : The basidium is a cell or a structure found at the terminal end of


dikaryotic hypha in the hymenial layer of the gill exogenously. In the basidial cell
the dikaryotic nuclei fuse with each other ( karyogamy) to form diploid nucleus (2n)
which immediately undergoes meiosis (reduction division) to form a definite number
of basidiospores (n). Each basidial cell bears a definite number of haploid
basidiospores (2 or 4) which at maturity are forcibly shot off to disperse and fall
below the pileus under the influence of gravity on the substratum. In still air these

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Morphology and life cycle of Agaricus

haploid basidiospores (+ and-) get deposited below the pileus in a mass to form
the spore print.

Figure: Development of basidium and formation of spores

Source: Author

Basidiospore: The haploid, monokaryotic basidiospore varies in size, shape, colour


and is thick walled and multilayered (3-5), eg. three layered in Psilocybe sp. and
Agaricus brunnescens (bisporus). It consists of protoplasm, haploid nucleus (+or -
), ribosomes, mitochondria, lipid bodies, vacuoles, glycogen granules as reserve food
material . The basidiospores exhibit distinct colours eg. in Chlorophyllum
molybdites spores are green, in Russula sp. the spores are yellow and in other

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Morphology and life cycle of Agaricus

species spores are white, pink, brown, black and smoky-grey. The spore colour is
not always the same as that of gills and is determined by making a spore print
on white and black paper .

A B

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Morphology and life cycle of Agaricus

Figure: Basidiospores and formation primary hyphae

Figure: Diagrammatic life cycle of Agaricus sp.

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Morphology and life cycle of Agaricus

Source:Author

Figure: Life cycle of a typical agaric (Amanita sp.)

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Morphology and life cycle of Agaricus

Fairy Rings
The fruiting bodies of mushrooms appear in rings on the moist grassy green lands, gardens
and on the decaying cellulosic substrata in the forests during spring, early summers and
winters exhibiting the creamish white rhizomatous subterranean peripheral mycelial growth
in abundance. The term “fairy ring” for these beautiful bodies appearing on the ground was
initially used by early philosophical taxonomists and naturalists for the fairies visiting these
areas, dancing, roaming around and enjoying in circles the previous nights which resulted in
the appearance of these beautiful (young unopened floral bud like structures) fruiting bodies
on the footsteps of these fairies in the early hours of the successive mornings. Similarly
multiple stories of our grannies express the miraculously emergence of white or colourful
fruiting bodies in rings on the grassy lawns in the early hours of the morning of spring,
summer and winters.

Figure: Fairy rings


Source: [Link]
,[Link]

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Morphology and life cycle of Agaricus

Scientifically the seed like mushroom (+&- strain, haploid, uninucleate ) basidiospores
absorb moisture and minerals from the substratum and germinate to form (monokariotic)
primary mycelia, which later by somatogamy form dikaryotic, binucleated, haploid
secondary mycelia. These dikaryotic mycelia anastomose with each other longitudinally to
form long thick strands of rhizomatous mycelium which spread around to form initially the
discoid mycelial creamish white colony on the substratum.

Figure: Photograph of rhizomatous mycelia on substrata

Source:[Link]

[Link]

As the colony matures the primary mycelia (the life span of which is very short) in the
center of the colony die and the rhizomatic strands towards the periphery of colony
anastomose to form small tissues or primordial buds ( button stage) in the arcs or complete
circle which is named as fairy ring. These primordial buds (basidiocarps) mature with in
forty eight hours, get differentiated into basel stipe and terminal pileus, underneath of which
the abaxial layer infolds to form gills or lamellae. The gills exhibit the fertile
zone(hymenium)where probasidial cells undergo karyogamy and meiosis to form two or four
basidiospores, initially hyaline and yellow to brown at maturity with in three to four days.
The mature basidiocarps (dark brown) shed their basidiospores on the substratum and die
but the rhizomatous secondary mycelia survive and perennate in the soil or on the
substratum for years (400 years in Marasmius sp. Butler and Jones,1949).

The parennating rhyzomatic (commonly called shoe string) mycelia in the soil are
responsible for the reccurrence of fairy rings in the same areas every year. The most
common examples are Marasmius oreades,hlorophyllum molybdites, Agaricus arvensis, A.
campestris,A. tabularis, Coprinus sp., Clitocybe sp. etc. The area inside the fairy ring
exhibits a distict zone of grass, more greener than the grass of the rest of the lawn. This

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Morphology and life cycle of Agaricus

effect is due to the degeneration of older hyphae of the fungal mycelium which released the
nitrogenous compounds in the soil and became available to the grass as fertilizer along with
the moisture available in the center.

Bioluminescent fungi
The rhyzomatic mycelium of some basidiomycetes exhibit an interesting feature of emitting
visible light at night in the decaying woods, old exposed roots of the trees and the decaying
organic matter in the forests. These fungi are bioluminescent in nature.

Figure: Bioluminescent fungi A. Marasmius elegans, B. Omphalotus C. Mycena


haematopus; D. Mycena chlorophos; E. M. interrupta ; F. O. olearus

Source: [Link]
[Link]
[Link],
[Link]
[Link]
.jpg/220px-Mycena_interrupta_01_Pengo.jpg,
[Link]

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Morphology and life cycle of Agaricus

The mycelium of these fungi produce the compact mass of hyphal strands of great length in
which the hyphae loose their individuality and the entire strands act like thick units of
creamish white or black, brittle rhizomorphs (commonly called shoe strings) which penetrate
and spread extensively under the bark (outer ligno-cellulosic protective zone) and in the
cortical layers of old forest trees to nourish the fungus parasitically and glow in the dark.
This parasitic nature of bioluminescent basidiomycetes kill the infected trees and continue to
live saprophytically on dead trees which are left glowing as `ghost` trees in the forests eg.
Armillariella mellea(Honey agaric) , Clitocybe illudens, Mycena galericulata ,
Omphalotus olearius etc. This phenomenon of glowing of rhizomatous mycelium on
woods or trees is called fox–fire.

The luminescent effect of the fungus can be exhibited in the mycology laboratory by
bringing in the piecies of wood completely colonized by mycelium of Armillariella mellea and
also by culturing the mycelium of this fungus or some other agarics on PDA agar medium
which show this effect.

The basidiocarps, the gills and basidiospores of bioluminescent fungi can be used in the labs
to demonstrate the glowing effects e.g. Omphalotus olearius (jack-o`lantern),
Lampteromyces japonicus(poisonous moon light),Mycena lux –coeli and Mycena rorida
respectively.

The bioluminescent fungi exhibit the best luminous effect at 10-25 ± 1 temperature and 80-
90% relative humidity on the substratum and the light given off is the byproduct of the
biochemical (chemiluminescent) reaction which involved the direct conversion of chemical
energy in to light energy which is actually responsible to remove oxygen from the fungal
cells.

The formula of bioluminescent reaction involved in emission of light energy(cold) and the
expulsion of oxygen from fungal cell:

ATP + Luciferin + Luciferase + O2 = CO2 + Light

Molecular energy+ substrate + enzyme + anaerobic condition=Removal of O2 + light


energy (cold)

The fungal luminescence is a function of living tissue in which a continuous process of


chemical reaction involves a substrate (D-luciferin) combining with molecular energy (ATP)
and oxygen, that is regulated by the fungal enzyme (luciferase) releasing a specific molecule
(luminescent) , energy level of which increases and results decay of the substratum, which
in turn it releases the photons hence producing the light. This process does not require any

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Morphology and life cycle of Agaricus

light stimulus and continues during day and night in the hyphal cells to produce light energy
called cold light e.g. the Mycena lampadis, a singal large specimen that exhibits the
emission of the light through its hyphal cells in an excellent amount that it is sufficient to
read a book even at night.

The bioluminescence is an important function in the bioluminescent (glowing fungi) fungi,


also called wood decayers, to release the photonic light energy responsible to remove the
oxygen from fungal hyphal cells which do not require oxygen and direct sunlight for their
growth andnutrition.

Summary
Basidiomycetes are highly advanced fungi .The thallus or mycelium is well developed
septate , hyaline , uni, bi or multinucleated, highly branched ,creamish white in colour.
They are either saprophytes or parasites and exhibit mycorrhizal association with
roots of forest trees . They reproduce asexually by fragmented spores. The sexual
reproduction takes place by simple fusion of two somatic cells of opposite strains of
primary hyphae by somatogamy .The basidiospores or meiospores are produced on the
secondary probasidial cells in the hymenial layer of the gills exogenously on the
basidiocarp . There is no differentiation of sex organs in basidiomycetes which reflects
that there is degeneration of sex organs and it is same in ascomycetes where the
ascogenous hyphae take part in sexual reproduction and produce ascospores or
meiospores in ascus mother cell endogenously in ascocarp (fruiting body) . There is no
motile phase in both of these classes .

Exercise/ Practice
1. Define the following with example :
 Basidium
 Hymenium
 Clamp connection
 Somatogamy
 Dikaryon
 Basidiocarp
2. Draw well labelled diagram of the following :

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Morphology and life cycle of Agaricus

 Dolipore septum

 V.S. Gill

 L .S. Basidiocarp

 Clamp connection

3. Fill in the blanks :

 The terminal cap of basidiocarp is called ---------.

 The fruiting body of mushroom is called --------.

 The fertile layer of basidiocarp is ------------.

 The mass of hyphae is called -----------.

 Fusion of two somatic cells is called ------.

 Fairy rings are formed by -----------.

3. Illustrate with the help of labeled diagrams only the life cycle of Agaricus
bisporous.

4. Write the salient features of basidiomycetes .

Glossary
Annulus : (ring) A remnant of the inner veil (a ring) around the stipe in some
mushrooms.
Basidiocarp: (Gr. Basidion- small base;karpos – fruit) A fruiting body
that bears basidia .
Basidiospore : (Gr. Basidion – small base + spora – spore , seed) A spore borne on
the outside of a basidium , following karyogamy and meiosis .
Basidiole or basidiolum : (L. dimin. Of basidium) A type of sterile element in
the hymenium of certain basidiomycetes ; resembles a basidium without
basidiospores .
Basidium : (pl. basidial ; Gr. Basidion – a small base) A structure bearing on its
surface a definite number of basidiospores (typically four) that are usually formed
following karyogamy and meiosis .
Binding hyphae : Thick walled , typically aseptate , highly branched hphae
present in the basidiocarps of some fungi .

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Morphology and life cycle of Agaricus

Clamp connection : A bridge like hyphal connection characteristic of the


secondary mycelium of many basidiomycetes .
Cortina : (L. Cortina – curtin) A curtin like , cobwebby veil hanging from the margin
of the cap of certain mushrooms .
Cystidium : (kystis – bladder + idion, dimin. Suffix) A sterile element Occurring in
the hymenium of certain basidiomycetes; Cystidia are generally larger than other
hymenial elements and protrude beyond them .
Dikaryon : (NL. Di – two + Gr. Karyon – nut, nucleus) A pair of closely associated
nuclei , each usually derived from a different parent cell .
Dikaryotic : It is pertaining to cell that contains a dikarion .
Dolipore septum : (L. dolium - large jar + pore) A septum with a central pore
surround by a barrel-shaped swelling of the septal wall and covered
on both sides by a perforated membrane termed either the septal pore cap ;
common in many basidiomycetes .
Fairy ring : A ring of fruiting body of mushrooms on the substratum or the
ground representing the periphery of the mycelial growth of a basidiomycete .
Inner veil : The hyphal membrane covering the gills of some young mushrooms .
Karyogamy : (Gr. karyon- nut , nucleus + gamos –marriage , union) The Fusion of two
nuclei.
Meiosis : (Gr. meiosis – reduction) A saries of two nuclear divisions usually in a
quick succession in which the chromosomenumber is redused by one- half .
Monokaryotic : (Gr. monos – single+ karion – nucleus , nut) A cell containing a single
nucleus .
Mushroom : A fleshy , sometimes tough umbrella-like basidiocarp or Fruiting body
of certain basidiomycetes .
Mycelium : (pl. mycelia; Gr. mykes – mushroom , fungus) The mass of hyphae
constituting the body (thallus) of a fungus .
Mycology : (Gr. mykes – mushroom , fungus + logos – discourse) The study of fungi
.
Mycophagy : (Gr. mykes – mushroom + phagein – to eat) The eating of
Mushrooms .
Oidiophore : (Gr. oidion – small egg + phoreus – bearer) A specialized hypha that
bears oidia .
Oideum : (pl. oidia; Gr. oidion – small egg) A thin walled , free , hyphal cell
derived from the fragmentation of a somatic hypha in to its component cells or

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Morphology and life cycle of Agaricus

from an oideophore; it behaves as a sporeor as a spermatium ; normally used in


reference to such
cells produced by some basidiomycetes .
Pileus : (L. pileus – cap) A terminal portion or cap of certain types of ascocarps
and basidiocarps .
Plasmogamy : (Gr. plasma- a molded substance , gamos – union , marriage )The
fusion of two protoplasts .
Plectenchyma : (Gr. pleko – 1weave;enchyma – infusion , i. e. a woven tissue) The
general term employed to designate all types of fungal tissues; the two most
common types of tissues are prosenchyma and pseudoparenchyma .
Primordium : (L. primordium –beginning) The first stage of any structure .
Probasidium : The portion of the basidium in which karyogamy takes place .
Somatogamy : The fusion of somatic cells during plasmogamy .
Sphaerocyst : The spherical cells present in the trama of the russullaceae.
Spore : A minute propagative unit functioning as a seed , but from it as the
spore does not contain an embryo .
Stipe : The stalk of the stipitate basidiocarp .
Trama : The fungal tissue composing the pileus or bearing the hymenium of the
Holobasidiomycetidae .
Zygote : Adiploid cell resulting from the union of two haploid cells .

References
Greuter ,B . and D , Rast . 1975 . Ultrastructure of the dormant Agaricus bisporus
spore . Can . J . Bot . 53 : 2096-2101 .
Litten. W .1975 . The most poisonous mushrooms . Scientific American. 232: 91-101.
Malloch, D .1976 . Agaricus brunnescens , the cultivated mushroom .Mycologia 68:
910-919 .
Singer,R .1975 . The Agaricales in Modern Taxonomy . 3 rd Edition. J . Cramer ,
Weinheim .
Stocks,D . L . and W . M . Hess .1970 .Ultra structure of dormant and germinated
basidiospores of a species of Psilocybe . Mycologia 62:176-191 .
Alexopoulos C . J . and Charles W . Mims . 1979 . Introductory Mycology .3rd edn .
Wiley Eastern Ltd .
Alexpoulos, C . J ., Mims ,C . W . and Blackwell , M .1996 . Introductory Mycology . 4th
edn . John Wiley and Sons, New York .

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Morphology and life cycle of Agaricus

Talbot , P . H . B . 1971 . Principles of fungal taxonomy . Macmillon , London .


Webster , J . and Weber , R . 2007 . Introduction to Fungi . 3rd edn .Cambridge
University .
Sharma , P . D . 2005 . Fungi and Allied Organisms .Narosa Publ . N . Delhi .
Inderjeet , K . Sethi , and Walia , S . K . 2011 . Textbook of Fungi and their Allies .
Macmillan Publ . India , Ltd .

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Common questions

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Changes in pH and temperature significantly impact Agaricus' lifecycle. Optimal conditions include a pH of 5.8-6.0 and temperatures between 12-18°C. Deviations can hinder mycelial growth and basidiocarp formation, affecting overall reproductive success and spore dispersal. Such environmental sensitivity underscores the necessity for fungi to adapt to specific niche conditions to thrive .

Fairy rings form when rhizomatous mycelium spreads outwards from a central point, depleting nutrients centrally and forcing growth to the periphery, resulting in a circular arrangement. This leads to greener grass due to the nitrogen release from decaying hyphae, which acts as a fertilizer. This phenomenon reflects the recycling of nutrients and the significant influence of fungi on soil health and plant ecology .

In hemiangiocarpous development, the hymenium is initially enclosed by a veil, which breaks as the pileus matures. Gymnocarpous development exposes the hymenium since its inception, while in pseudoangiocarpous development, the hymenium is exposed initially but later covered by outgrowths of the pileus, which only reveal it upon maturity. These distinctions affect how spores are protected during their development .

Primary mycelium in Agaricus originates from monokaryotic, haploid basidiospores and has a short life span. It consists of thin, hyaline, monokaryotic hyphae. Secondary mycelium forms when haploid primary hyphae of opposite strains come in contact through somatogamy to produce dikaryotic, binucleated mycelium which persists longer. Tertiary mycelium creates the structural tissue of the basidiocarp, enclosing the hymenium during initial development .

The formation of the basidiocarp in Agaricus species is influenced by several environmental factors, including diffused light, a relative humidity of 80-90%, temperature ranging from 12-18°C, a pH of the substratum between 5.8-6.0, along with the availability of mineral nutrients and aeration. These conditions are critical for the development and maturation of the fruiting bodies .

Somatogamy in Agaricus involves the fusion of somatic cells from primary mycelial hyphae of opposite mating types (+ and -), resulting in the formation of dikaryotic secondary mycelium. This dikaryotic state is crucial for the development of the fruiting body and subsequent sexual reproduction, ensuring genetic diversity through meiosis in basidia .

Agaricus fruiting bodies exhibit several adaptations for reproductive success, including gills beneath the pileus to maximize surface area for basidiospore production, an annulus to protect developing spores, and a stem (stipe) to elevate the body into the air for spore dispersal. These morphological features optimize spore release into the wind, enhancing propagation .

Agaricus participates in mycorrhizal associations, particularly with coniferous trees, enhancing nutrient exchange between fungi and host plants. This symbiosis aids in nutrient absorption, improves soil structure, and increases plant tolerance to stress, contributing significantly to ecosystem productivity and resilience .

Bioluminescence in basidiomycetes is facilitated by a biochemical process where the substrate luciferin combines with ATP and oxygen, catalyzed by the enzyme luciferase, to produce light without heat ('cold light') and release CO2. This phenomenon occurs in the living tissue and does not require external light or oxygen, likely serving a role in deterring predators and attracting spore dispersal agents .

In Agaricales, the hymenium is crucial as it houses the reproductive cells, which undergo meiosis to produce basidiospores. It is located beneath the pileus, lining the gills or lamellae, providing a structured environment for spore maturation and release. The structural characteristics of the hymenium are often used for taxonomic classification, highlighting its importance beyond mere reproduction .

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