FUNGI

CHAPTER/

Introduction
1.1 General account of Fungi : According to E. Toofill (1984) the fungi contains
bout 5100 genera and abou 50,000 species. However according to the latest estimation
that the actual number of species may be between 100,000 and 250000 (Toofill, 1984).
According to lIngold (1967),. "the fungal kingdom is a large one with from 50000 to 100000
known species". Fungi (Singular : fungus) are found to occur everywhere i.e. they are
found in soil, water, air and even in plant and animal bodies. The literal meaning of the
term 'fungus' is mushroom, and the study of mushrooms i.e. fungi is known as mycology
-4 greek word (mykes = mushroom, and logos = discourse). The fungi constitute a large
group of living organisms devoid of chlorophyll pigment. Other characters common to
green plants are the presence of true nucleus and distinct cell wall in fungal cell, but the
vegetative body of fungi is thallus-like. Moreover the vegetative body of fungus never
possess vascular tissues. All fungi are eukaryotes because of the presence of true nuclei
and cell organelles in their cells.
A. DEFINITIOoN The fungi are chlorophyll-less thallophytes having, "heterotrophic
(saprophytic.parasitic, symbiotic or hyperparasitic) eukaryotic and spore bearing organisms
surrounded by a well defined cell wall made up of chitin, with or without fungal cellulose
along with many other complex organic molecules". They may be defined as chlorophyll-
less, non-vascular and non-autophytic i.e. heterophytic members of Thallophyta, excluding
the Bacteria and Mycetozoa, which reproduce asexually and sexually (Bessey, 1950).
According to Alexopoulos (1962), the term fungus should be restricted to the "nucleated,
spore-bearing. achlorophyllous organisms which generally reproduce sexually and
asexually, and whose usually filamentous, branched somatic structure are typically
surrounded by cell walls containing cellulose, or chitin or both."
B. HABTS OF FUNGI -On the basis of mode of nutrition, fungi are of two types mainly
eg, (i) saprophytic and (i) parasitic.
In general fungi obtain food by absorption, except a few lower groups where they take
tood by ingestion. The fungi which live on dead or decaying organic plant and animal
remains, and other rotten organic food matters are known as saprobes or saprophytes e.g.
species of Mucor, Rhizopus, Agaricus, etc. The fungi growing on dung of herbivores are
also saprophytes but they are called coprophilous fungi, e.g., Ascobolus sp.
h e vegetative body i.e., hyphae (refer page 596, C) of saprophytic fungi come in
intmate contact with the substratum and obtain food by direct diffusion through the hyphal
Walls, causing thereby disintegration of the organic matter which is utilised by those fungi.
When fungi live on other living organisms (plants and animals) and obtain their food
LE. nutrient from the living tissues of the organisms upon which they live, they are called
uases. The vegetative body i.e. hyphae of the parasitic fungi grow either intercellularly
tracellularBy in the host tissue. The hyphae if growing in between the host cells is
d intercellular. Fungi when grow intercellularly, they send out specialised and modified
fr g organs called haustoria (singular: haustorium). The victims i.e. living organisms
rom which the parasitic fungi draw food are called the hosts. These fungi obtain nutrients
00d matters from the cells of the host tissue with the help of those haustoria. In shape,
Ustoria may be knob-like, short or long, unbranched or branched forming a miniature
system (Fig. 1.1). These haustoria are developed as outgrowths from the vegetative

Vol (1)-38
 594 STUDIES IN BOTANY
body of the parasitic fungi. These haustoria are generally penetrated into the host
cells through minute pores punctured in the cell wall. The formation of haustoria is n ost'sliving
a
response to the probably
contact
HAUSTORIUM stimulus as well as to
HYPHA HOST CELL
stimulus of nutrients. the
parasitising animal tissue areFungi
not
known to
produce
haustoria.
Among fungi, different
of degrees
parasitism
and
saprophytism
can be noted, viz-
(a) Some fungi obtain their
food entirely from the
HOST CELL
HOSTCELL protoplasm of the hostsliving
and
HYPHA maintain their mode of life
throughout their life cycle only
HAUSTORIUM3
as parasites -

this type is called

obligate parasite, e.g. Puccinia
HOST- (rust fungi), Peronospora, etc.
CELL WALLX But when organisms obtain their
HOST food only from dead organic
PROTOPLAST matters and maintain their mode
of life throughout the life cycle
as saprophytes, the organisms are
Fig. 1.1-Different types of haustoria. A-Knob-like short called obligate saprophytes or
haustoria. B-Long flattened type of haustoria.
C-Branched root-like haustoria.
obligate saprobes e.g
Saprolegnia monoica. Obligate
saprophytes are incapable of infecting other living organisms.
(b) While some fungi are normally parasitic in their mode of life but later on, according
to circumstances, may pass their mode of life as saprophytes also this type is called
facultative saprophyte or facultative saprobe (e.g. Ustilago).
(c) Facultative parasites-When fungi passing their mode oflife as saprophytes in the
beginning and later on, under certain conditions become also parasites causing diseases,
they are called facultative parasites, e.g. Pestalotia.
Parasitic fungi again may be
(1) Ectoparasites - When parasitic fungi grow superficially on the surface ofthehost
plant without penetrating the host tissue, they are called ectoparasites. Ectoparasiticfung
are provided with special organs of attachment known as appressoria (singular
appressorium). According to Priston and Gullegly (1954), an appressorium i a flattened
hyphal pressing organ from which a minute infection peg grows and enters the epidermal
cells of the host; with the help of this structure ectoparasites obtain nourishment from ine

protoplasts of the host tissue.

(2) Endoparasites - When fungi penetrate the host and ramify their vegetative body
within the tissues of the host, they are called endoparasites.
The parasitic fungi which spend their entire life cycle on a single host plant are callea
autoecious and those which require more than one host to complete their life cycle are
called heteroecious.
The fungi, like all other plants are divided upto species level. If necessary species
 INTRODUCTION
units of classification
595
into further eg. varieties, biological
splitted
races etc. For example Puccinia graminis is an established strains, physiological
species which
o r c u l t u r a

ecies
om other speci of Puccinia
on certain
morphological
differsal and pathological studies clearly indicate that characters, but
it (P. graminis) is not a single
necies. Puccinia graminis
uniform s p e c i e s . is well known to
infect the members of the
eat,
amineae (e.g. whea rye, oat etc.), but it is not that family
any member of P. graminis may
infect any of the diff
grass hosts.
different For example spores of P. graminis from wheat
the
will not infect other hosts such as
olant (host) rye, oat plants etc. or vice versa
norphologically spores from.om all hosts are although
exactly similar. This obviously indicates that
of P. graminis there are several forms
species of
vithin the e.g. wheat
.

ecting forms, rye infecting forms and so on. So morphological infecting

charactersforms,
(suchoat
as
LAi
ability
to infect and develop upon a particular type of host etc.) by means of which a
acies is divided into a number of sub-species or forma speciales e.g.
speci
Ch.cnecies P. graminis tritici on the host plant wheat
(Triticum aestivum)
P. graminis avenae 99 99
oat (Avena sativa)
P. graminis secalis » 99
rye (Secale cereale)
Again it has that sub-species is also not an uniform assemblage; further
been seen

cnlitting of a sub-species on the same criterion is also possible. A host species e.g. Triticum
aestivum (wheat) is also composed of several varieties such as N.P. 710, N.P. 798, Ridley
etc, The wheat infecting sub-species (P. graminis tritici) is not equally pathogenic on all
the varieties of wheat. So there are forms within this sub-species each of which is adapted
differently on different varieties of wheat. This is an extreme instance of the phenomenon
of host range reaction amongst obligate parasites and which is referred to as physiologic
specialization. " And the forms within the species of an obligate parasite which are

morphologically indistinguishable but are separable from one another on the basis of the
reacion againsthost varieties are known as physiologic race."Physiologic specialization
is also poted in Peronospora, Cystopus and many other rust fungi.
esides as parasites or as saprophytes, fungi sometimes live in close association
with algae (eg. lichens) - such mode of life is known as symbiosis. In this type of

PSEUDOPARENCHYMATOUS
MANTLE

INTERCELLULAR
MYCORRHIZA- HYPHA

NORMAL ROOT
A B

Fig
g2-Diagram showing, A- Mycorrhiza on a portion of root and B-Fungus mantle on the
outer surface of the root and hyphae in intercellular spaces of cortex.
Clation both the organisms are mutually benefitted. Sometimes fungi also live in
a5Ociation with the roots of gymnosperms and angiosperms - this association
 STUDIES IN BOTANY
596
between fungal hyphae and roots of higher plants is known as mycore
ia (literally,
and the associated fungi are called mycorrhi fungi.
fungus-root)
are not harmful but rather beneficial yet
Althou; these
these
types of associations two
looked upon as mild type of parasitism, because some endotrophic m associatio
form naustoria by means of which they
absorb utrients from
çells of
nizal fungiare
Mycorrhiza may be ectotrophic and endotrophic.
hos
mer type occurs in almost tissue.
trees (both gymnosperms and angiosperms), in this type many intercellular f forest
lular fungal hyphae
occur largely in the cortex and remain connected with well developed externalr
-
this forms a mantle on the outer surface of the short root and replaces mycelium
the pilifer
layer with its root hairs. Most members of Agaricaceae are ecto
ctotrophic mycorrhiza. In
case of endotrophic mycorrhiza, the infected roots contain well develope
in the intercellular spaces and penetrating many of the living cortical hyphae runnin
cells. Exter
hyphae may be only poorly developed. All orchids have endophytic mycorrhizas.Sne
of Rhizoctonia are good example of endophytic mycorrhizas.
Some fungi may grow epiphytically on the surface of higher plant parts withoe
causing any major harm - these are called epiphytic fungi. hout
C. VEGETATIVE i.e. SoMATIC STRUCTURES-Vegetative body of fungi may be as simnle
unicellular (e.g. Chytrids and Yeasts), or in majority the body is made up oflongand
slender filaments called hyphae (singular: hypha). Filaments constituting the hyphaeof
fungi elongate by apical growth. Hyphae usually branch freely and intertwine toform a
mass ofmore or less loosely interwoven hyphae constituting the mycelium (plural:mycelia
Hyphae may be hyaline or coloured, simple or branched, septate i.e. cellular or aseptate
i.e. without septa (Fig. 1.3) forming long, much-branched coenocytic cell.
In septate hypha, the protoplasm is interrupted at regular intervals by the formation

HYPHAL CELLS SEPTUM

PROTOPLAST
'.

Fig. 1.3-Somatic hyphae of fungi; A-Portion of aseptate (coenocytic) hypha.

B-Portion of septate hypha.
of partitions i.e. cross walls
(septa) during mitosis meiosis-as a result the nypd,
or
becomes divided into cellular structure. In
septate hyphae, the septa in most cases
possess a central pore - this type of septa is termed porous or
which cytoplasmic strands including nucleus
perforated septa, throug
migrate from one cell to other. Agan
may be without pores (termed non-porous septa).
In aseptate hypha, the protoplasm is continuous without being interrupted by coSS
 INTRODUCTION 597
nce the cylindrical walls of coenocytic cells are lined with cytoplasm containing
many small nuclei. However, in aseptate i.e. cocnocytic hyphae secondary septa (septa not
associated with nuclear divisions) are formed to delimit reproductive structures or older
narts of hyphae. Mycelium may be ectophytic (growing on the surface of the substratum)
and endophytic (growing within the substratum). Development of the mycelium of a
fungus generally begins as a short germ tube emerging from a germinating spore (Fig.
14). In thickness hypha varies according to types from 0.5 to 1004. In size the entire
mycelium may be only a few microns in length or it may be very long as far as covering
metres.
The composition of the hyphal wall ie. cell wall varies in different classes of fungi. In
most fungi, the cell wall is composed of microfibrils of chitin having the formula
(C.HNO,,) n. According to Bracker et al (1976) the chitin microfibrils are formed by
theenzyme chitin synthetase. Chitin is a polymer composed of N-acetyl -2-glucosamine
units, it is often impregnated with some salts or similar other substances. In some fungi the
wall is made up of cellulose or other polysaccharides, e.g. glucans, mannan etc. In few
cases cellulose and chitin have been reported to occur together. In the cell wall of many
fungi, callose (a complex carbohydrate), lignin-like substances and other organic material
have also been detected.
In septate i.e. cellular hypha, hyphal cells may contain one nucleus (uninucleate), two
nuclei (binucleate) or many nuclei
(multinucleate) in each cell. In higher fungi,
two phases are noted in the life cycle-firstly,
A in which the hyphal cells are uninucleate, is
called monokaryon or monokaryotic phase
and the corresponding mycelium is called
monokaryotic mycelium; and secondly, in
which the cells are binucleate, is called
dikaryon or dikaryotic phase and the
corresponding mycelium is called dikaryotic
mycelium. Monokaryotic and dikaryotic
mycelium are also known as primary and
secondary mycelium respectively. In other
C
members, hyphal cells are multinucleate.
The vegetative fungal cells has almost the
same ultrastructure like other eukaryotes. The
Fig. 14 Different stages (A-D) in cell remains surrounded by distinct cell wall.
ascospore germination. Composition of cell wall is described
previously. The cell wall is followed by plasmalemma: it regulates the movement of soluble
materials into and out of a hypha. In between the plasmalemma and cell wall or at the
Surface of plasmalemma some membranous structures known as lomasomes are present.
The cell contains either many small vacuoles or a single large vacuole, each of which is
Surrounded by a
tonoplast.
Distinct double membrane bound nucleus with well developed nucleolus consisting
mostly of RNA and distinct chromatin strand which becomes organised into chromosomes
during nuclear division are also present, the nuclei are either globose or ellipsoid, the
nuclear membrane are porous; other double membrane bound organelles (about 1
ng) such as mitochondria, tubular endoplasmic reticulum, Golgi apparatus, ribosomes
a r e also present. However, like prokaryotes, the ribosomes lie free in the cytoplasm
datached to endoplasmic reticulum. The cytoplasm is surrounded by a
 598 STUDIES IN BOTANY

plasmamembrane i.e. plasmalemma. Mitochondria are small and oval. Pres

lomasomes in the form of particles, vesicles or tubules at some places between of
wall and the plasmamembrane is an unsusal feature of fungal cells. All f e
chlorophyl1, but some other pigments has been reported e.g. reddish violet ack
known as 'neocercosporin' (C,H,O from Cercosporina kikuchi has Temeat
Matsueda et al (1978). Plastids and starch grains are absent. eported by
CELL WALL
RIBOSOMES LOMASOMES

PLASMA
MEMBRANE

NUCLEUS

ENDOPLASMIC
RETICULUM

MITOCHONDRION GLYCOGEN VACUOLE

Fig. 1.5-Ultrastructure of a portion of fungal hypha.

The nuclei of fungi are very small - therefore their study is very difficult. The division
of the nucleus is both mitotic and meiotic. It has been observed with the help of electron
microscope thatnuclear membrane, during mitotic division, does not disappear but constricts
like a dumbbell (Fig. 1.6)-this type of nuclear membrane-behaviour during mitotic division
is called karyochoresis. In meiosis however the nuclear membrane
disappears.
The chief reserve food material occurs as granules of glycogen which is an
aglucopyranose polymer-like starch; this glycogen gives brown colouration with iodine
solution. In some fungi manitol (alcohol) is also found as food reserve. Sometimes,
lipids and proteins are also found as food reserve.

NUCLECLUS

NUCLEAR MEMBRANE

CHROMOsOME

CHROMOSOME
SPINDLE A

CENTRIOLE

Fig.1.6-Different stages of the mitotic division in fungi. A-Metaphase.

B-Telophase. C-Interphase.
D. NUTRmiON- Fungi are unable to manufacture their own food material due absence

of the green pigment. Hence

majority of fungi obtain food either from living orgai
 INTRODUCTION 599
i t e s or from dead organic substances as saprophytes. Alexopoulos and Mims (1979)

mentioned that usually fungi require C,

all

K, Mg. S, B, Mn, Cu, Zn, Fe,

0, H, N, P,
in the form of essential element
Mo and Ca
nutritional requirement. Fungi also
for their
some compounds which are
synthesise
functioning as vitamins. Glucose,

ammonium and nitrogenous compounds

as

best food of fungi. Excess

nitrates form the
in the form of glycogen
is stored by fungi
and lipids.
Í.e. ORGANISATION OF
E. MoDIFICATION
THE MYCELIUM Tangled mass of hyphae B

ie. themycelium of most fungi may be

modified as such:
(a) Plectenchyma
-When hyphae
grow together, inter-twine, and become
organised into loosely or compactly
woven
Fig. 1.7Fungal tissues (plectenchyma).
massive tissues, it is called plectenchyma. A- Prosenchyma. B-Pseudo-parenchymna.
Practically all organised fungal tissues are
called plectenchyma. Plectenchyma may be oftwo types- in
tissue is loosely woven
1. Prosenchyma or Prosoplectenchyma-In this case, the
REPRODUCTIVE BODY

STROMATIC TISSUE

Fig. 1.8- Stroma and sclerotium.

A-Section through a stroma. B- Structural Fig.1.9-Rhizomorph.
detail of stromatic tissue. C- Sclerotium.
D-T.s. of sclerotium.
to one another and their elongated
the component hyphae lie more or less parallel
wnich
cells are easily distinguishable as such.
 600 STUDIES IN BOTANY
2. Pseudoparenchyma or Paraplectenchyma- In this type of tissue hyphal indivl .
is lost, as a result hyphal cells are not distinguishable as such; hyphae lie sideaty
closely so that in a cross-section cells look isodiametric or oval and resemble nare SIde
cells of higher plants. ma
(b) Sclerotium (plural : Sclerotia)- It is also usually composed of pseudoparenchy
in which interior cells are hyaline and stored with food and outer cells are thickwall.ma
dark brown or black and crust-like. Structurally it is rounded or cushion-shaped. Sclert
are tough and resting bodies, e.g. Claviceps sp.
(c) Stroma (plural; Stromata) - It is a somewhat indefinite solid body formed by manv

fungi from either prosenchyma or pseudoparenchyma. Reproductive structures ar

fructifications commonly develop in or from this structure, e.g. Daldinia sp. (Fig. 1.8).
(d) Rhizomorph - Here the hyphae run parallely side by side and adhere to each
other forming dark brown, hard and thick strand of somatic hyphae in which the hyphae
have lost their individuality here the whole mass behaves as an organised unit. The
structure of the growing tip of the rhizomorph somewhat resembles that of a root tip.
These structures are resistant to unfavourable conditions and remain dormant until
favourable conditions return. In a mature rhizomorph there is present a central core
consisting of large, elongated, thin-walled cells. The central core remains enveloped by
a rind or outer covering composed of thick-walled small cells. Rhizomorphs may creep
under bark of trees or underground and serve to spread fungus from one favourable
place to another place of suitable food. (Fig.1.9)
E GRowTH OF THE MYCELIUM:-The filaments constituting the body of a fungus grow
only at the tips. Growth involves cell enlargement followed by cell division and cytoplasm
synthesis. The growing hypha forms branches in acropetal order behind the growing tip.
Most fungi grow between 0° and 35°C. but the optimum temperatures lie in the range of
20- 30C. In case of septate hyphae, following methods are also important
for growth.
I. By cell division-In most of the cases, growth
of the
hypha takes place by nuclear division followed by
transverse wall formation between the daughter nuclei.
. By conjugate divison- In fact this method is a
specialised type of cell division which occurs in case of
hypha of binucleate cells, i.e. dikaryotic mycelium. In this
A C case, the nuclei are oriented side by side and both of them
divide simultaneously. Transverse wall is next formed in

A between the two pairs of nuclei.

II. By clamp connections In all major types or
Basidiomycetes, an interesting mechanism is notedo
ensure that daughter nuclei arising from conjugate division

D
of the dikaryon become separatèd in the two daughter
E F
cells. This mechanism takes place by the help of speca
Fig.1.10-Clamp connections Curved outgrowths called clamp connections which
arc

showingdifferent stages (A-F) formed during nuclear division of binucleate cells.

binucleate cell is ready to divide, a short lateral outgro
a

called the clamp, arises between the two nuclei (Fig.1.10, A); this clamp
Deg
form
ahook-like structure and
represents clamp connection. the
Next one 0) nucleus migrates into the outgrowth and the other (x) remains win
cell. Now both the nuclei (r and y) divide simultaneously. Oblique one

division result in the formation orientauo

of one daughter nucleus (y) in the clamp
connecu
 INTRODUCTION 601
daughter nucleus (y) in the dividing cell (Fig.1.10). Simultaneously the orientation
the uecond division result in the formation of one daughter nucleus (x) near the lower
o ft h e s e c o n d

cell and the daughter nucleus (x) near the nucleus (0).
sal end ofthe
t h e clamp bends over and its free end fuses with the cell. The wall at the point
of fusion dissolves and the daughter
nucleus (y) approaches another
daughter nucleus (x). Now the
clamp becomes closed by the
formation of a septum at the point
of its origin. A second vertical
septum develops just below the
level of the out-growth of the
clamp-this second vertical septum
divides the parent cell into two

B
daughter cells of which the terminal
cell contains the daughter nuclei (r*
and y') whereas the lower daughter
cell contains the daughter nuclei (x)
and ().
G. Reproduction-Reproduction
A-Transverse
Fig. 1.11-Vegetative reproduction. is the phenomenon of the formation
cell division (fission). B-Budding8 of new individuals having identical
Three general types of reproductions are recognised
characters typical of the species.
asexual and (c) sexual. Vegetative and asexual
in fungi, such a s : (a) vegetative, (6) but sexual
union of nuclei, sex cells (gametes) or sex organs,
types do not involve
union of two nuclei or sex cells. At the time of the
reproduction is characterised by the when the entire vegetative
formation of either asexual or sexual reproductive structures,
structures then such fungi are
thallus gets converted into one or more reproductive
differentiated into vegetative (somatic) and
called holocarpic. When the thallus is
reproductive parts, the fungi, are called eucarpic. the following
(a) Vegetative reproduction'-This type reproduction takes place by
of
methods -
some portion i.e. hyphal filaments
1. Fragmentation- Breaking up ofthe entire thallus or
a new individual.
into one or more pieces, where each piece grows into
where the vegetative
2. Fission-This (Fig. 1.11-A) is noted in unicellular types (Yeasts)
transverse wall.
cell simply splits into two daughter cells by constriction or
3. Budding- In case ofbudding, vegetative, ie. parent
cell gives rise to small outgrowths
cell and grows into a new individual e.8.
called buds, each bud separates from the parent sometimes gives
Yeasts (Fig. 1.11,B). In some fungi ascospores and basidiospores
higher
nse to lateral outgrowths which cut off buds by budding-
suitable conditions are available
4.Resting bodies like sclerotia also germinate as soon as
and those give rise to new vegetative bodies.
method of the formation of new
(6) Asexual reproduction It is often defined
-
as a
means of one or
by non-sexual means. Asexual reproduction takes place by
VIdual units called spores. Morphologically
ANOTe kind of specialised cells i.e. asexual reproductive
non-motile, endogenously or exogenously
POres may be one or many-celled, motile or
ormed. Biologically spores may be asexual or sexual.
some mycologists.
Vgetative reproduction is also considered as a type of sexual reproduction by
 602 STUDIES IN BOTANY
The different kinds of spores in fungi may be classified as follows
SPORES
I Aserual (These spores are IL. Asexual but related to II. Sexual (These
vegetative in nature, they are meant sexual reproductive cycle. the spores are
for methods of
products of the union of
accessory sex cells i.e.
multiplication and do not possess gametes-hence
these are called
any relation to a sexual process, sexual
spores. Sexual spores are
hence they do not take part in diploid and take part in
alternation of generations). alternation of generations).

1.Oidia Ascospore Basidiospore

2. Conidia
3. Zoospores
4. Sporangiospores or Aplanospores
5. Chlamydospores
Oospore Zygospore
6. Spores of Rust fungi
) Uredospores
i) Aeciospores
1. (a) Oidia' (Singular : Oidium)-These are vegetatively formed cells arising directly
from the breaking up of hyphae; the hyphae break up into numerous, somewhat rectangular
or rounded fragments called oidia (fig. 1.12,A). Oidia behave like spores. These are also
known as arthrospores. Examples Trichosporon beigeli (Deuteromycetes), Coprinus
lagopus (Basidiomycetes) etc.
(b) Conidia (Singular: Conidium) -These are non-motile spores cut off externally
either singly (Phytophthora) or in chains (Penicillium) from the tip of a special aerial
hyphae called conidiophores or phialides (Fig. 1.13, A-B). Conidiophore may be
unbranched (Aspergillus) or branched (Penicillium). The type and arrangement of conidia
and conidiophore in few other fungi are as follows
of
In Trichothecium conidia are two-celled. In Fusarium and Microsporum two types
conidia are formed e.g. some are small unicellular called microconidia and others
are

form globular clusters

large, multicellular called macroconidia. In Trichoderma conidia
on the conidiophores.
According to Bessey (1950), conidia may arise by the fragmentation of the whole
cel- this type is regarded
hyphae or of special hyphae into cylindrical, ovoid or spherical
by him as oidial mode of conidium formation.
(c) Zoospores or Swarmspores-These are endogenously produced unicellular, naked
are developed
and motile spores having one or two flagella (Fig. 1.13, D_H. Zoospores
within the sac-like structure called zoosporangium formed at the tip of zoosporangiophore,

Rust fungi are characterised in having peculiar form of life cycle specially when growing o
s
in them. Uredospore
different kinds of hosts. Consequently different types of spores are noticed
for basidium proaueuo
essentially meant for asexual reproduction. Teliospore is apreparatory stage
Pycniospore is uninucleate and meant for producing binucleate condition ultimately. Acciospo
the starting of binucleate condition.
Formation of oidia and their germination into individual have been regarded as a method
Alexopoulos an
vegetative reproduction by some authors (Alexopoulos, 1962; Srivastava, 1960; res

Mims, 1979; and others). According to Stevenson (1967) oidia are asexual reproductive surue
i.e. spores.
 INTRODUCTION
603
Saprolegnia (Fig. 2.7, B), Pyhium (Fig. 2.10,B), Phytophthora (Fig. 2.15, C-D) etc.
C a of zo0spores may be of tinsel type (i.e. hairy along its entire
type length)rigid and
non-hairy). Whiplash type is provided with two parts, viz. lower
or of
whiplash
longer
part, and upper short and flexible portion.
authors suggest that zoospores of Many
but motile form of fungi
are nothing
sporangiospores.
(d) Sporangiospores or
are unicellular,
Aplanospores-These
and
endogenously produced, non-motile
a-OIDIUM
non-flagellate spores formed within the sac-
like structure called
sporangium (Fig.1.13, C)
developed at the tip of special erect hypha called
sporangiophore, e.g. Mucor, Rhizopus, etc.
(e) Chlamydospores- These are unicellular,
non-motile, thick-walled, dark-coloured, non-
deciduous resting spores (Fig. 1.12, B), formed
endogenously when terminal or intercalary hyphal
cells enlarge, round up, accumulate much food
reserves and form a thick wall, e.g. Fusarium,
Phytophthora, Mucor, etc.
CHLAMYDOSPORE () Thallospores -In this type, the vegetative
part of the thallus develops into spore-like structure
called thallospores e.g. in some yeasts and some
other fungi. Blastospores, arthrospores etc. are also
Fig. 1.12-Formation of oidia (A) and called thallospores.

chlamydospores (B). II. (a) Ascospores-Characteristic spores of

the sub-division
Ascomycotina. These are
unicellular or multicellular and non-motile spores formed endogenously in a definite number
(4 or 8) after meiosis within a special type of sporangium called ascus, e.g.,
Saccharomyces
(Fig. 3.12, A), Peziza (Fig, 3.29, B), Ascobolus, etc.
(6) Basidiospores- Characteristic spores of the sub-division
are non-motile, unicellular Basidiomycotina -these
spores formed exogenously on a basidium as a result of meiosis,
Agaricus (Fig.4.16, D), Coprinus, etc.
The formation of two above mentioned
spores is associated with the phenomenon of
sexual reproduction; their formation is preceeded by the union of two nuclei (may be
regarded as male and female) followed by meiosis.
. (a) zygospores-These are sexually produced diploid spores, characteristic of

Lygomycetes (Mucor, Rhizopus, etc.) under the sub-division Phycomycotina. Zygospores

are formed as a result of the union of two morphologically undifferentiated gametes.
6) Oospores-These spores are formed as a result of the fusion of two morphologically
Oiterentiated gametes i.e. male and female. These are characteristics of the class Oomycetes
under the sub-division
Phycomycotina and of some Ascomycetes.
Sexual reproduction - It is the method of the production of new individual by
exual means. In fungi as in other living organisms sexual reproduction involves the fusion
OT two compatible (i.e. mutually tolerent) nuclei to form a zygote-nucleus. There are
Pcaly three distinct phases in the process of sexual reproduction and these are
DrPlasmogamy -It is first of the three phases. This phase involves the union of wo
PrOOplasts bringing thereby the two haploid nuclei close together in one cell.
Karyogamy - It is the second phase. This phase involves the fusion of two haploid
604 STUDIES IN BOTANY
nuclei (which are brought together by plasmogamy) into one diploid i.e.
zygote nucleus
There may be gap in time between plasmogamy and karyogamy or
karyogamy follow
plasmogamy immediately as seen in many members of lower fungi. In higher funo
karyogamy is delayed more or less, and the plasmogamy results only in the establishment
of a binucleate cell containing one nucleus from each parent -such a
pair of nuclei is

CONIDIUM
SPORANGIOSPORES

SPORANGIUM
A

**i°1

B
D

Fig. 1.13-Different types of spores in fungi A-Conidia in chain borne on a

B-Single conidium on conidiophore. C-Sporangiospores borne in a conidiophoresporangium
D-Uniflagellate zoospores with whiplash flagellum. E-Biflagellate primary zoospores w
tinsel and whiplash flagella. F-Biflagellate secondary zoospore with tinsel and whiplash flageus

called a dikaryon. The fusion of these two nuclei i.e. karyogamy takes place consideraouy
in the later
part of the life history of the fungus. In the mean time, during growth and
c
division of the binucleate cell, the dikaryotic condition may be perpetuated trom ccell to
cell by the simultaneous (i.e. conjugate) division of the two such closely associated nucic
and by the separation of the
resulting daughter nuclei into cells.
daughter
The process by which the dikaryotic condition is established is called dikaryolizaio
and the karyogamy by which the diploid condition of a nucleus is established is cail
diploidization.
(iii) Meiosis -It constitutes the third phase. Karyogamy, sooner or later, 15 wed
mheby
meiosis which again reduces the diploid number of chromosomes to the haploid nuin
 INTRODUCTION 605
Ormiin ologies used in sexual reproduction in Fungi - Fungi may be
Terms a n d t e r
both distinguishable male and female sex organs occur
homothallic where bo
onoeciousi.e.
.. dioecious i.
bedioecious i.e. heterothallic wherethe male and female
thalus,
or they may other words, fungi may be bisexual and unisexual.
differe thalli. In
same
an the occurs
on
s e x o r g a n s

i.c. or bisexual fungi are represented by t sign while

monoecious
homothallic

Theothallic i . e . dioecious
orunisexual fungi are represented by the +or sign depending
naleness respectively. The sex organs are called gametangia (singular;
and male
femaleness
upon
ametangium), gametangia
gia may produce differentiated sexual units called gametes or may
one or more units called gamete-nuclei. Gametangia may be
c e undifferentiated
ilar
rphologically simila
p r o d
i.e. isogametangia or may be dissimilar i.e. heterogametangia.
ogametangia producei
morphologically similar gametes i.e. isogametes. Heterogametangia
aredifferentiated into male gametangia called antheridia (singular antheridium) and
The gametes produced by antheridium and oogonium
gametangia called oogonium.
female
phologically different
and are called male and female gametes respectively. In
may be 1flagellate (planogametes) or non-flagellate (aplanogametes).
are

fungi gametes may

fungi gametes and
On the basis of the development, the sex organs (antheridia oogonia) may be
diclinous and androgynous. When antheridia and oogonia are developed on different hyphal
hranches, they are called diclinou5. In androgynous type, both the sex organs are developed
onthe same hyphal branch and the antheridium arises immediately below the oogonium-
stalk.
Again on the basis of the method offertilization, sex organs (antheridia and oogonia)
may be amphigynous and paragynous. In amphigynous condition of sex organs, the
oogonium punctures the antheridium and passes through it and takes the form of a globose
structure above the antheridium; the antheridium is thus present in the form of a funnel
shaped collar around the base of the mature oogonium (e.g. Phytophthora infestans). When

SPERM

EGG

Fg 1.14-Sexual reproduction (gametic copulation) in fungi. A-Isogamy by isogametes.

B-Anisogamy by anisogametes. C-Anisogamy by anisogametes (oogamy)
in Monoblepharella sp.
heanheridium arises from the branch of the same hypha (bearing oogonium) or from a
O u n g branch and the antheridium itself is applied to the wall of the oogor
 STUDIES IN BOTANY
606
the time of fertilization, it is called paragynous. Paragynous condition of sex organs atthe
time of fertilization is noted in androgynous type while amphigynous condition at the
time of fertilization is noted in diclinous type.
The various methods of sexual reproduetion in fungi may be grouped into following

types
1. Gametic copulation - This process involves the fusion of two gametes. When fusion
takes place between two isogametes, whether both of which are planogamous (motile) or
aplanogamous (non-motile), the process is called isogamy When fusion takes place between
two heterogametes or anisogametes, the process is called anisogamy or heterogamy. In
heterogamy both the copulating gametes may be planogamous (1.14, B) or one of the
heterogametes, i.e. smaller one may be motile (planogamous) and the other i.e. larger one
being non-motile (aplanogamous): theformer enters the oogonium and fertilises the latter
(Fig. 1.14, C), eg. Monoblepharis, Monoblepharella, etc.

2. Gametangial contact -In majority of fungi, gametès are not released from the
gametangia to the outside, instead gamete or gametes are directly transferred from one
gametangium into the other. In this process, the two gametangia of opposite sex (i.e. male
and female gametangia, otherwise known as antheridium and oogonium) come in contact
with each other; next one or more gamete-nuclei migrate from the male gametangium
(antheridium) to the female gametangium (oogonium) either through a pore formed by
NUCLE!
(MALE GAMETES)
ANTHERIDIUM

FERTILIZATION
TUBE
SOMATIC HYPHA

NUCLEUJS
.

EGG
OOGONIUM (FEMALE GAMETE)

Fig. 1.15-Sexual reproduction through gametangial contact in Pythium aphanidematum,

the
formation of one or more fertilization tubes (serving as a passage) developed from the
male gametangium. Examples-Pythium aphanidermatum, P. debaryanum, etc. After the
migration of gamete or gametes from the male gametangium or the antheridium both the
gametangial.e. oogonium and antheridium may eventually disintegrate or the antheridiu
only disintegrates while the oogonium continues its development in various ways. Win
the exception of few fungi, the gametes of the male gametangia are reduced
undifferentiated protoplast, each consisting of a nucleus.
3. Gametangial copulation- In this method the fusion of the entire contents of thne
 INTRODUCTION
607
l a t i n g gametangia takes place. Ihis process may take place in any one of the two
two copula

ways
Transfer ofthe
th entire contents of one
gametangium into the other
() through a pore
FEMALE HOST CELL
MALE GAMETANGIUM GAMETANGIUM- WALL

.'

HOST CELL
WALL'
A
B
Fig. 1.16-Sexual reproduction through gametangial copulation in Rhizophidiun couchii.
developed in the gametangial walls at the point of contact of the two gametangia. The
male gametangium attaching itself to and emptying its entire content into the female
gametangium. Example- Rhizophidium couchii. This method is noted in holocarpic form
where the entire thallus acts as a gametangium.

A
GAMETANGIA-
MATURE ZYGOTE
(ZYGOSPORE)

Fig1.17-Sexual reproduction through gametangial copulation is Sporodinia grandis.

) By direct fusion of the two gametangia into one cell- this takes place by the
Olution of the common contacting walls of the two gametangia resulting in the fornation
One common cell in which the protoplast of the two gametangia mix with each other.
Examples -Sporodinia grandis, Mucor sp.,
t
permatization-This is the process by which small, uninucleate, unicellular spore
DOIes called spermatia (singular : spermatium) arecarried through the various agencies
 STUDIES IN BOTANY
608
like insect, water, wind etc. to the female gametangia, or to special recentive
sometimes to somatic hyphae to which they remain applied. The con hyphae or
pass into those receptive structures (which functions as female organ) through n spermatia
formed
SPERMATIOPHORE
TRICHOGYNE
SPERMATIUM
(RECEPTIVE
FEMALE ORGAN)
TRICHOGYNE
sPERMATIUM

ASCOGONIUM

SOMATIC HYPHA-

B D
A

Fig. 1.18-Sexual reproduction by means of spermatization in Podospora anserina

A-Spermatophore withspermatium. B-Ascogonium with trichogyne. C-Spermatization.
D-Plasmogamy.
by the dissolution of walls at the point of contact. The spermatia serve as male organs.
Examples-Podospora anserina, Puccinia sp. etc.
5. Somatogamy -In most of the higher fungi, sex organs are absent. In such fungi
somatic cells take over the sexual reproduction i.e. anastomosis between the cells of the

SOMATOGAMY
SPORE SPORE

i i

Fig. 1.19-Sexual reproduction through somatogamy is Peniophora S

somatic hyphae takes place; in this process the compatible nucleus of one cell passes
the corresponding somatic cell by dissolving the walls between them and thus a dikaryo

condition is established. Example -Peniophora sp. (Fig. 1.19) union

H. Sexual Compatibility - In fungi, sexual reproduction actually means tne
where

between two nuclei which are contributed by the male and female gametes n cment of
ent of
there is visible sexuality, i.e. where sexual reproduction takes place by the developn
morphologically distinguishable male and female gametangia and gametes. It has bee du
is due
finitykn
noted that the two nuclei having some affinity unite with each other and thisafin known

nuclei are
to the presence ofsome factors which help them to unite-such uniting
 INTRODUCTION 609
.Tethere
there are no such factors, the nuclei fail to unite, and the nuclei are
If
atible
compatible,

therefore
as
mDatible.
called incompatible. Similar condition is noted in
majority of fungi which produce
and female sex organs on the same
male and
hable male
distinguishable thallus, but they (thalli)
clearly s erile because their male organs are incompatible with their female
sexually

be
of som factors controlling
may

organs
dueto
absence
compatibility.
o On the basis of sex, Alexopoulos (1962)has classified most fungi into three categories
a sfollows
ermaphrodite ((also designated as monoecious)- Here each thallus bears both
1. Hermaphrodite
female sex organs (gametangia).
male and
2. Dioecious (also called dimorphic)- In this, some thalli bear only male sex organs
ome other thalli bear only female sex organs.
whiles

iated- In
3. Sexually undiferentiat this case, sex organs are produced but they are not
marphologically distinguishable into male and female.

On the basis of compatibility, fungi in the above sex categories may belong to any one
three groups
ofthe following
(a) Homothallic fungi - Those fungi in which every thallus is sexually selffertile

fungi under this group can reproduce sexually by itself without the help of another thallus.
(b) Heterothallic fungi - Those fungi in which every thallus is sexually self-sterile and

therefore requires the help of another compatible thallus of a different mating type for
sexual reproduction.
Heterothallic fungi again belong to any one of the following two categories
) Bipolar heterothallic -Fungi in this group consist of two mating types of individuals
which differ in their genetic make-up for the compatibility factor. One mating type carries
the gene A in each nucleus, and the other mating type
thalli
carries the gene a in each nucleus.
Only whose nuclei carry opposite genes of this Mendelian Aa are pair compatible.
(i) Tetrapolar heterothallic -Fungi in this group consist of four mating types of
individuals. Here compatibility is governed by two pairs of factors such as Aa and
Bb
located on different chromosomes. Only thalli whose nuclei
carry opposite genes of
both Mendelian pairs Aa and Bb are
compatible, the resulting zygote has the genotype
AaBb.
(c) Secondary Homothallic fungi -

In some bipolar heterothallic fungi during spore

formation, two nuclei of opposite mating type are regularly incorporated in each spore.
Each spore on
germination therefore produces a thallus which contains both A and a
nuclei and
naturally behaves as though it was homothallic. This condition is called
secondary homothallism.
Life cycle - Like other living organisms, fungi also possess a cycle of haploid
and diploid structures corresponding to gametophyte and sporophyte of higher plants.
Tungi are haploid organisms, they have haploid nuclei in their cells. Asexual cycle
maintained through asexual reproduction with the help of haploid spores. The diploid
Pnase begins with the fusion of two nuclei i.e. karyogamy and ends with the meiosis.
&rue i.e normal sexual cycle, three important processes viz, plasmogamy, karyogamy
cear fusion i.e. diploidization) and meiosis (haploidization) occur in a regular sequence
Vol (1)-39
 STUDIES IN BOTANY
610
and generally at specified points in the life cycle of an organism as follows
Spore (n)
Spore-bearing on germination
structure
Asexual
Vegetative body (n)- Reproductive structures (n)
(haplophase)
Plasmogamy
Sexual
on
Dikaryon (n+n)
germination (Dikaryophase)
Karyogamy
Spores (n) Meiosis Zygote (2n)
(haplophase) (Diplophase)
In majority of fungi, distinct alternation of generation is absent. In the life cycle of
most fungi, "the diploid phase occupies a very much smaller portion of the life cycle than
does the haploid phase." The hyphae of fungi are also called haplonts or diplonts according
to the haploid (n) or diploid (2n) number of chromosomes respectively.
The diagram of the life cycle mentioned above shows the general scheme of the asexual
and sexual cycle in fungi. However, the sexual cycle may again be elaborated into following
four types :
1. Haploid cycle In this type meiosis occurs immediately after nuclear fusion
karyogamy) and the meiotic products are then dispersed. Therefore the diploid phase
is of short duration. Members of many Phycomycotina and some Ascomycotina exhibit
this type of life cycle.
karyogamy
Haplophase (n) Diplophase (2n)
meiosis
2. Haploid-dikaryotic cycle This type is similar to former type except that paired
compatible nuclei persist in close association "in the same hyphal segment (hence dikaryon
and divide synchronously, for a greater or lesser period". This type is seen in many
Ascomycotina and some Basidiomycotina (Ustilaginales).
Plasmogamy Dikaryophase (n+n)
Haplophase (n)
meiosisS
Diplophase
(2n) Karyogamy
3. Haploid-diploid cycle - In this type haplophase and diplophase alternate in regia
succession. It is found in a few species of aquatic Oomycetes of the sub-a
Phycomycotina.
plasmogamy & karyogamy
Haplophase (n) Diplophase (2n)
meiosis t
se is
4. Diploid cycle -In this type the diploid phase is elaborate and the haplopna
nediately by
restricted to sexual reproductive structures; the haplopha is followed
diplophase. This type is usually noted among large number of the class Oomycc
 INTRODUCTION
611
Phycomycotina.

Diplophase (2n)
meiosis
Reproductive structures (n)
plasmmogamy & karyogamy
PARASEXUAIITY - It has been also observed that some fungi do not
go through a
ie true sexual cycle, in them plasmogamy, karyogamy and meiosis take place
norma

but not
at specifie points or at a specified time in the life cycle. Such a cycle is known
cycle d the process is parasexuality. Parasexuality was first discovered
and
as parasex
2 by Pontecorvo and Roper of the University of Glasgow in Aspergillus nidulans
nerfect
and stage
perfect sta; of Emericella nidulans. (For details see in connection with
Deuteromycotina. Chapter 5)
committee of the international code of Botanical Nomenclature recommended the
se of the following "suffixes" for the divisions, sub-divisions and other units of Fungi
Division should end in - mycota; Sub-division should end in - mycotina

- mycetes; Sub-classes "" - mycetidae

Classes
99
- ales; Families
Order 99
- aceae

FUNGI

Mycelium usually Mycelium septate

aseptate

Class I Phycomycetes Characteristic Characteristic Ascospores,

spores are spores are basidioopores
ascospores basidiospores are absent
Mycelium Myceliumn
absent or well
rudimentary developed Class II Class Ill Class IV
Ascomycetes Basidiomyctes Fungi Imperfecti
Sexual Sexual
1.Archimycetes reproduction by reproduction by Number of Number of
e.g. Synchytrium, dissimilar similar gametes,
Olpidium, etc. basidiospores basidiospores
gametes, the the result is indefinitee definite
result is oospore
zygospore
2. Oomycetes e.g. 3. Zygomycetes 1. Hemibasidio-
Saprolegnia, e.g. Mucor, mycetes Basidium Basidium
Pythium, Rhizopus, etc. e.g. Ustilago, septate not septate,
Phytophthora, etc. Tilletia, etc.
continuous
AScOcarp rounded, Ascocarp saucer Asci in parallel 2. Protobasidi- 3. Autobasi
Asci in ascocarp
shaped. Asci in series. omycetes diomycetes
irregularly ascocarp lie in Ascocarp flask e.g. Puccinia, e.g. Polyporus,
aranged parallel series. shaped. etc. Agaricus,
1. Plectomycetes Phallus, etc.
2. Discomycetes 3. Pyrenomycetes
e.g. Saccharomyces,
e.g. Peziza. e.g. Claviceps,
Penicillium, Ascobolus, Xylaria,
Eurotium, Helvella, etc. Laboulbenia, etc.
Erysiphe, etc.
 STUDIES IN BOTANY
612
K. Classification Fungi are divided by most of the authors into four classes main
Of all the classifications, the most standard and widely accepted classification is that adon
to the septation of the mycelitum
by Gwynne-Vaughan and Barnes (1927). According
the characters of the principal spores, they have divided the fungi into three main elac
such as Phycomycetes, Ascomycetes and Basidiomycetes. Although Fungi Imperfecti haas
been given the rank of a class (fourth) still the group does not form the main class as the
members of Fungi Imperfecti are reduced i.e. incompletely known. The outline of the
classification of H. C. I. Gwynne-Vaughan and B. Barnes (1937) is given in the page 611
E. A. Bessey (1950) classified true fungi into four classes, viz. Phycomycetes
Ascomycetes, Basidiomycetes and Fungi Imperfecti. He included Mycetozoa under Funoi
separating Mycetozoa as a sub-class from the fungi; therefore Bessey designated all funoi
excluding Mycetozoa as the True Fungi.
Martin (1931, 1941) classified fungi into three classes, viz, Phycomycetes, Ascomycetes
and Basidiomycetes, and two form-classes viz. Fungi Imperfecti and Lichens. In 1961 he
treated Fungi as sub-division Eumycotina which includes three classes viz.Phycomycetes,
Ascomycetes and Basidiomycetes, and two form-classes viz. Lichens and Deuteromycetes
(Fungi Imperfecti).
Alexopoulos (1962) designated the term Mycota for all fungi. Mycota is given t
rank of a division. The division Mycota is divided into two sub-divisions, viz. the
Myxomycotina (true slime moulds) and the Eumycotina (true fungi). The sub-division
Eumycotina ie. true fungi are sub-divided into eight classes and one form-class as follows:-
1. Class Chytridiomycetes-Fungi with variety of thalli structures, whose motile cells
are provided with a single posterior whiplash flagellum.
2. Class Hyphochytridiomycetes-A small group of aquatic fungi whose motile cells
are provided with a single anterior tinsel flagellum.
3. Class OomycetesFungi with variety ofthalli structures, whose motile cells possess
two oppositely directed flagella, one whiplash type and other tinsel type. Sexual reproduction
resulting in the formation of resting spore (oospore) formed from the fertilised egg.
4. Class Plasmodiophoromycetes -Parasitic fungi with non-cellular, multinucleate
thalli living in the cells of their hosts. Resting spores produced in masses, but not in distinct
fruiting bodies. Motile cells possess two anterior whiplash flagella.
5. Class Zygomycetes -Saprophytic or parasitic fungi with a well developed
coenocytic or septate mycelium. Sexual reproduction resulting in the formation of a
resting spore (zygospore) formed by the fusion of two usually equal gametangia. Motile
cells are not formed.
6. Class 7richomycetes- Fungi with simple or branched filamentous
coenocytic thallus,
attached to digestive tract or the external cuticle of
living arthropods. Asexual reproductuon
by a variety of spores. Sexual reproduction where known is like that of Zygomycetes.
7. Class
Ascomycetes-Fungi which form spores, resulting from and
karyogamy meiosi
within special sac-like structures called asci
(singular : ascus).
8. Form-class Deuteromycetes (Fungi Imperfecti) - In this form-class placed fung
which, in their general structure and asexual
reproduction, resemble the
Basidiomycetes, but in which sexual stages have not been discovered.
Ascomycetes a
9. Class Basidiomycetes -Fungi which form spores, resulting from karyogamy and d

meiosis, on the surface of special structures called basidia (singular : basidium)

ohn Webster (1970,77) classified fungi following the scheme proposed by Ains
(1966), which is outlined below
FUNGI (Mycota)
Division I. Myxomycota
Class (1) Acrasiomycetes
 INTRODUCTION 613
Order Acrasiales
Class (2) Hydromyxomycetes
Order Hydromyxales
Order Labyrinthulales
Class (3) Myxomycetes (Slime moulds)
Class (4) Plasmodiophoromycetes
Order Plasmodiophorales
Division II. Eumycota (Mycobionta : eumycetes)
Sub-divisionl, Mastigomycotina
Class (1) Chytridiomycetes
Order Blastocladiales
Order Chytridiales (chytrids)
Order Monoblepharidales
Class (2) Hyphochytridiomycetes
Order Hyphochytriales
Class (3) Oomycetes
Order Lagenidiales
Order Leptomitales
Order Peronosporales (downy mildews etc.)
Order Saprolegniales
Sub-division 2. Zygomycotina
Class (1) Zygomycetes
Order Entomophthorales
Order Mucorales
Class (2) Trichomycetes
Sub-divison 3. Ascomycotina (Ascomycetes)
Class (1) Hemiascomycetes
Order Endomycetales
Order Taphrinales
Class (2) Plectomycetes
Order Erysiphalcs (powdery mildews etc.)
Order Eurotiales
Class (3) Pyrenomycetes
Order Hypocreales
Order Sphaeriales
Class (4)
Discomycetes (inoperculate discomycetes)
Order Helotiales
Order Phacidiales (operculate discomycetes)
Order Pezizales (hypogaeous discomycetes)
Order Tuberales (truffles)
Class (5) Laboulbeniomycetes
Order Laboulbeniales
Class (6)
Order
Loculoascomycetes (bitunicate ascomycetes)
Capnodiales
Order Dothideales
Order Hysteriales
Order Microthyriales (Syn-Hemisphaeriales)
Order Myriangiales
OrderPleosporales
Sub-divison 4. Basidiomycotina
Class (1) (basidiomycetes)
Order
Hemibasidiomycetes
Uredinales (rusts)
 614 STUDIES IN BOTANY

Order Ustilaginales (smuts)

Class (2) Hymenomycetes
Order Agaricales (agaricus, boleti)
Order Aphyllophorales (polypores etc.)
Order Tulasnellales
Class (3) Gasteromycetes
Order Hymenogastrales
Order Lycoperdales
Order Nidulariales
Order Phallales
Order Sclerodermatales
Sub-divison 5. Deuteromycotina (Fungi imperfecti)
Class (1) Coelomycetes
Order Melanconiales
Order Sphaeropsidales
Class (2) Ayphomycetes
Order Hyphales (Syn. Moniliales)
Class (3) Agonomycetes (Mycelia sterilia)
Order Agonomycetales (Syn. Myceliales)
G. C. Ainsworth (1973) first proposed the well known classification of fungi in 1966
and followed by himself in the "Dictionary of Fungi" (1971) and also in his book Fungi
: An advanced Treatise "in 1973. This classification is phylogenetic that reflects natural
relationship. In this classifcation, fungi with plasmodium or pseudoplasmodium are placed
in a separate division 'Myxomycota, whereas majority of filamentous fungi without having
plasmodium or pseudoplasmodium are placed in true fungi "Eumycota."
1 . Classification of Fungi as proposed by G. C. Ainsworth (1973) followed by
John Webster (1980)

Divisions Fungi
MYXOMYcOTA EUMYCOTA
Plasmodium or Pseudoplasmodium present) (Plasmodium or
pseudoplasmodium absent)

Classes
Acrasiomycetes (free Labyrinthulales (net Myxomycetes Plasmodiophoromycetes
living assimilatory plasmodium is (presence of free living (presence of parasitic
phase of amoebae present) plasmodium within host
saprobic plasmodium)
unite as a
cells)
pseudoplasmodiumn
before reproduction) Sub-divisions

Mastigomycotina Zygomycotina Basidiomycotina Deuteromycotina

Ascomycoina (Zoospores and
(Zoospores (Zoospores (Zoospores and (Zoospores and
perfect-state spores like
present; perfect absent; perfect ZYgospores are Zygospores are
Zygospores, ascospores
state spores are state spores are absent; perfect- absent; perfect and basidiospores are
oospores) zygospores) state spores are state spores are
absent)
ascospores) basidiospores)
Classes

Classes Classes Classes Classes

1. Chytridiomycetes 1.Zygomycetes 1. Hemiascomycetes 1. Teliomyceles
1. Blastomycetes

2. Hyphochytridio 2. Trichomycetes 2. Loculoascomycetes 2. Hymenomycetes

2. Hyphomyceles

3. Coelomycees
mycetes 3. Plectomycetes 3. Gasteromycetes
3. Oomycetes 4. Laboulbeniomycetes
5. Pyrenomycetes
6. Discomycetes