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Fungi structure and reproduction

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CHAPTER 14:
Fungi structure and reproduction
Introduction

Section "A"

The fungi are a group of eukaryotic, non-vascular organism. Which are of diverse
forms, sizes, physiology and reproduces both by sexual (meiotic) and asexual
(mitotic) spores. Examples of fungi :-Mushrooms, yeasts, molds, Penicillium- the first
of the wonder drugs, penicillin, was isolated from this fungus and crop parasites.Fungi
are most often associated with the roots of some plant species, and this type of
symbiotic associations is known as mycorrhizae.

The study of fungi is known as mycology (Mykes = mushroom + logos = discourse) or


mycetology and who study about fungus is known as mycologist.

General characteristics of Fungi

Most fungi are eukaryotic,multinucleate, multicellular organisms, except yeasts


which are unicellular.
Nutrition Fungi are heterotrophic because these lack chlorophyll (green pigment)
and thus cannot create their own food through photosynthesis.Fungi acquire their
nutrients from dead organic matter by secretion of extracellular enzymes followed
by absorption. These are either :-
1. Saprophytes or Saprobes :- Most fungi are saprophytes ,obtaining nutrients
by absorbing dead organic matter.
2. Parasitic :-Some fungi are parasites, living in or on another organism (called a
host) from which they obtain their nutrients. This relationship usually harms
the host. Such parasitic fungi usually have specialized tissues called haustoria
that penetrate the host's body for food absorption.
3. Symbionts :- Some fungi live in a mutually beneficial symbiotic relationship
with another organism.
Lichen (association of fungi + algae)Some fungi are associated with
either cynobacteria or green algae and this type of symbiotic
associations is known as lichen.
Mycorrhizae (fungi + plants) :- Some fungi are most often associated
with the roots of some plant species, and this type of symbiotic
associations is known as mycorrhizae.Mutualistic association of plant
roots and fungi increase the absorptive surface area of plant roots.
(Fungus gets organic nutrients from plant; Plant gets minerals from the
soil via the fungus).
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Thallus organization (structure):-
Most of the fungi, are filamentous and the vegetative (=assimilative) stage is a
tubular branching thread-like filament called a hypha (plural = hyphae).
The hypha extends by tip growth, and multiplies by branching, creating a fine
network called a mycelium.
Two of the phyla Zygomycotina and the Chytridiomycotina (also known as lower
fungi) vegetative mycelium is without cross -walls known as non-septate (aseptate)
or coenocytes. Complete septa are only found in reproductive structures. In
aseptate mycelium numerous nuclei lie in a common mass of protoplasm.This
condition is called coenocytes.

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The other two phyla Ascomycotina and the Basidiomycotina( higher fungi)the
hyphae have cross walls that break them up into cells. A cross wall in fungi is called
a septum ( plural septa).The septum has a central pore which enable exchange of
nuclei and most organelles within the cytoplasm between adjacent compartments.
Septum formation is a simple process. Wall ingrowth towards the centre of the
compartment results in a complete or incomplete blockage of the hypha. Inward
growth may be followed by modification of the outer wall. The septum also
increases rigidity of the hypha as it can function as a structural support to the
turgor pressure within the compartment.
Ultrastructure of fungal thallus:-
Rigid cell walls are strengthened with chitin, a polymer of N-acetylglucosamine
(except oomycetes where cellulose present).
Food reserves are in the form of glycogen
Hyphae contain nuclei, mitochondria, ribosomes, golgi and membrane-bound
vesicles within a plasma-membrane bound cytoplasm. The sub-cellular structures
are supported and organized by micro-tubules and endoplasmic reticulum. The
cytoplasm and most organelles and inclusions of fungal cytoplasm are typical of
eukaryotic organisms. However, chloroplasts or plastids are absent.
Reproduction :-
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At the time of reproduction when entire cell asin case of unicellular fungus may be
converted into reproductive structure. Such a condition is known as holocarpic. But
if only a portion of thallus is converted into reproductive structure , it is called
eucarpic.
Some fungi are unicellular and are termed as yeasts. These grow by binary fission
or budding, creating new individuals from the parent cell.
Members of the Ascomycotina produce asexual conidiospores and sexual
ascospores in sac-shaped cells called asci.
Fungi from the Basidiomycotina rarely produce asexual spores, and produce their
sexual spores from club-shaped basidia in complex fruit bodies

Classification of Fungi

Over 60,000 species of fungi are known. Fungi are classified primarily by their method of
reproduction (both sexual and asexual) and fruiting bodies (asexual or sexual spores
surrounded by highly organised protective structures). In the earliest classification, there
were only two recognized kingdoms:- Plants and Animals.This two kingdom system was
used until Whitaker (1969) proposed that organisms be classified into five kingdoms :-
Monera (=Bacteria), Protista (=Mostly Algae and Protozoans), Plantae (=Plants), Mycetae
(=Fungi) and Animalia (=Animals).

Fungi belong to Domain-Eukarya and Kingdom -Mycetae (=Fungi) The classification of


fungi, as proposed by Ainsworth (1973), is commonly followed :-

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The more recent systems of classification that is based, in part, on molecular research
are :-

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Section "B"

Division :- Myxomycota (plasmodial slime molds)

The organisms included in this division are commonly known as the plasmodial or
acellular slime molds. The exact systemic position of the Myxomycota is not clear. Olive
(1975) classified them in the Kingdom Protista. Ainsworth (1973) classification is most
accepted in which Division Myxomycota belong to Kingdom Mycetae (=Fungi).

Vegetative phase is unicellular, without a cell wall, a multinucleate mass of


protoplasm called a plasmodium.
They engulf bacteria, protozoa, and other food particles by phagocytosis.During
this mode of ingestion, the food particles, usually bacteria, beceome surrounded
by the pseudopodia of the myxamoeba. Once the food has been engulfed in this
matter, it is surrounded by a membrane or food vacuole where hydrolytic enzymes
are secreted that will digest the food.
The vegetative stage in slime molds is morphologically similar to that of an
amoeba, because of that known as myxamoeba. Plasmodium produces one or
more sporangia where meiosis takes place.

Within the Myxomycota, the class Myxomycetes, known as the acellular slime molds.
Class: Myxomycetes There are approximately 71 genera and 500 species of
Myxomycetes.

Habitats (Occurrence) :-There are commonly occur in damp places especially on


decaying wood, all of which are terrestrial (found on land).

Reproduction :-

Asexual reproduction occurs by binary fission of myxamoeba or fragmentation of


the plasmodium.
Sexual reproduction takes place by the plasmogamy (fusion of compatible
gametes) occurs between myxamoebae or swarm cells (some species are
heterothallic).Karyogamy (fusion of nucleus) occurs shortly after plasmogamy to
form zygote (2n). Zygote forms plasmodium – longest lived vegetative stage.

Life Cycle of Myxomycetes:- Bi-phasic life cycle with a haploid and a diploid phase.
The diploid stage is a naked coenocytic protoplast (that is, it consists of a
multinucleate mass of cytoplasm that is enclosed only by a plasma membrane and
does not have a cell wall).When the sporangia matured and turned grey. Normally
globose, with a definite, rather thick cell wall,unicellular, uninucelate and haploid spores
are released by rupture of the sporangial coat.The spore surface may range from almost
smooth to reticulate. Spores of myxomycetes are small (4-20 µm) and are easily picked
up by air currents, arthropods and other animals On germination a spore produces one
or four myxamoebae or flagellate cells known as swarm cells- that have one or two
anterior flagella, all of which are of the whiplash type. In most cases swarm cells possess
one long prominent flagellum and a second shorter, inconspicuous flagellum that is
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directed backward and appressed to the cell surface. Myxamoebae and swarm cells can
be interconverted. When food is abundant and environmental conditions are favorable,
myxamoebae divide repeatedly, giving rise to a large population of cells. The nuclear
divisions are centric and open (the nuclear membrane breaks down during prophase
and is reconstituted after telophase.) Under unfavorable conditions myxamoebae round
up and secrete a galactosamine wall to form microcysts (sclerotia). When favorable
conditions return, the microcysts germinate and either a myxamoeba or swarm cell
emerges from each wall.

image1405

When free water is available myxamoeba can differentiate into flagellated swarm cells
Swarm cells and myxamoeba may function as gametes (both in homothallic or
heterothallic strains).The compatible gametes fuse in pairs (two swarm cells, two
myxamoebae) to form a diploid zygote.As the zygote grows, its nucleus undergoes
successive synchronous mitotic divisions without cytokinesis. And the cell becomes
transformed into a multinucleate, amoeboid structure, the plasmodium.
Plasmodium:-The plasmodium is a diploid structure.Plasmodium is a naked,
multinucleate, motile mass of protoplasm; no cell wall around it. However, in most
species, the plasmodium is enveloped by a gelatinous slime sheath that contains
microfibrils. Just inside the slime sheath is the plasma membrane that surrounds and
confines the cytoplasm Plasmodia are of various colors. Examples- Physarum
polycephalum it is a bright yellow, slimy structure and Didymium iridis the plasmodium
is colorless In nature plasmodia probably feed on bacteria, spores of fungi and plants,
and possibly on protozoa and even on bits of nonliving organic matter Growth is
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accompanied by successive mitotic divisions of the nuclei embedded in the cytoplasm. In
growing plasmodia of P. polycephalum, nuclear division occurs almost simultaneously
every 8-10 hours throughout the plasmodium and requires 20 to 40 minutes for
completion. There are three types of plasmodia :- protoplasmodium - microscopic
throughout its existence; gives rise to only a single sporangium when it fruits.
aphanoplasmodium - resembles a protoplasmodium in its initial stages, but soon
elongates, branches, and becomes a network of very fine, transparent strands;
Stemonitales. phaneroplasmodium - characteristic of Physarales, also resembles a
protoplasmodium at first; it grows larger and becomes more massive. Its protoplasm is
very granular, and the plasmodium is visible even at an early stage of development. The
gelified and fluid portions of the veins are easily distinguishable and the rhythmic,
reversible streaming is very conspicuous

Sporulation and Sporophores :-

Under favorable conditions, the plasmodium will migrate and feed for a period of
time before being converted to one or more sporophores.Sporophores posses
brightly coloured sporangia.
Entire plasmodium of a myxomycete usually is converted into one or more
sporophores so that the somatic and reproductive phases seldom coexist in the
same individual.
Sporangium consists of the following parts: -
1. Peridium (persistent or evanescent):- The fragile, outer layer of the
sporangium is the peridium (pl.=peridia), which may be persistent or
degenerate by the time the sporangium is ready to disperse its spores.
2. Hypothallus - The hypothallus is a plasmodial remnant forming the base for
one or more fruiting bodies. The hypothallus connects the stalk or stipe to
the substrate. It may be dull or brightly colored, thin and delicate or
coarse.The hypothallus may be composed of calcium carbonate
3. Stalk :- Basal portion of sporangium, may or may not be present, may be
hollow or filled with material Stalks formed from secretions of plasmodium
and are acellular (in contrast to cellular slime molds
4. Columella and Pseudocolumella :-The columella appears as an extension of
the stalk into the spore mass, although it may not resemble the stalk. In a
sessile fruiting body, the columella may be an area on the inside of the
peridium where it contacts the substrate or appears as a dome-shaped
structure. A pseudocolumella (pseudo=false) is a columella that does not
attach to the stalk. The pseudocolumella is found only in the order
Physarales, existing as a lime mass within the spore mass. Capillitial elements
may be attached to the columella or pseudocolumella.
5. Capillitium and Pseudocapillitium :-The capillitium consists of threadlike
elements inside the sporophores, intermingled with the spores.Some
elements of capillitium may be elastic, allowing for expansion when the
peridium opens, while other types are hygroscopic and capable of dispersing
spores by a twisting motion. A pseudocapillitium is present in some aethalia
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and pseudoaethalia producing species. Pseudocapillitial elements are highly
variable in size and shape, and may appear as bristles, threads or perforated
plates.
6. Spores.

Section "C"

General characteristics of Mastigomycotina :-

One large group of the Mastigomycotina is aquatic. While another group of the
Mastigomycotina are primarily terrestrial, although the organisms still form motile
zoospores when open water is available
The members of Mastigomycotina produce flagellated zoospores in their life cycle.
Most of them are filamentous and have coenocytic mycelium. However, unicellular
form are present, and some genera show the pseudosepta (false cross wall)
formation.
Rhizoids are present in some of unicellular forms.
They show centric nuclear division. Their centrioles remain functional during
nuclear division.
Live either as saprophytes or parasites.Due to presence of haustoria in a majority
of Mastigomycotina , the mode of nutrition is typically absorptive.
The sexual reproduction takes place by different methods , oospores formation
are common in almost all Mastigomycotina

Three classes are included in this sub-division, on the basis of zoospore and oospore and
comprise 204 genera and 1160 species:-

Chytridiomycetes produces posteriorly uniflagellate zoospores Chytridiomycetous


fungi occur as saprobes on plants and animal remains in water while other
members occur as parasites on algae and aquatic animals.
Hyphochytridiomycetes :- Zoospores are anteriorly uniflagellate.The
Hyphochytridiomycetes are those aquatic fungi whose thallus is holocarpic or
eucarpic, monocentric or polycentric and their vegetative system is rhizoidal or
hypha-like with intercalary swellings.
Oomycetes :- The Oomycetes contain 74 genera and 580 species, which are mostly
aquatic, though some are terrestrial and live as parasites or saprophytes.Includes
classic “water molds” in the Order Saprolegniales and the “downy mildews” in the
Order Peronosporales.

General characteristics of Class :- Oomycetes :-

Vegetative body is filamentous and coenocytic except the unicellular Lagenidiales.


Holocarpic or Eucarpic
1. Holocarpic - Entire thallus converted into reproductive structures
2. Eucarpic - Reproductive organs arise from only a portion of the thallus and
remainder continues as somatic Majority of species are eucarpic.
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Cell wall contains cellulose and glucans.Chitin is absent.
Asexual reproduction is by biflagellate heterokont (different) and anisokont
(unequal) zoospores that are produced in zoosporangia.
Zoosporangia- Modified hyphae that are usually terminal and delimited by a
septum
Zoospores are diploid formed by mitosis
Anteriorly directed flagellum is tinsel type and posteriorly directed is whiplash type
.Depending on genera single type-monomorphic or two types of zoospores are
formed-dimorphic . Two types of zoospores are formed in the life cycle are :-
1. Primary zoospores :- First formed, pip-shaped, and the flagella are located
anteriorly. Primary zoospore is released from the zoosporangium, encyst and
germinates to form the secondary zoospore.
2. Secondary zoospores :- The secondary zoospore which is reniform or bean-
shaped and laterally flagellated.
Zooporangium and zoospores are the major dispersal agents for most species.
Sexual reproduction:- Sexual reproduction is heterogamous (oogamous) by
oogonia (female) and antheridia (male). Female gamete (oosphere) produced by an
oogonium. Depending on taxon, there may be one to many oospheres per
oogonium Male gamete is produced by antheridium and transferred to the
oogonium by gametangial contact and migration of male nuclei into oogonia and
fertilize oospheres Homothallic– self-fertile or Heterothallic– opposite mating types
required for sexual reproduction. A swimming sperm is absent in the Oomycetes.
This type of sexual reproduction is referred to as gametangial copulation. In
antheridia and oogonia meiosis take place. The eggs and sperms are products of
meiosis and the only parts of the life cycle that are haploid. Diploid zygote develops
into thick-walled resistant oospore that germinates and give rise to vegatative
diploid hyphae that reproduce asexually by production of zoospores.

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image1406

The vegetative body is diploid and the life cycle is diplontic.


Class Oomycetes is divided into four orders.

Lagenidiales (Salilagenidiales)
Leptomitales
Saproleginales
Peronosporales :- This order has some of the most well known pathogens (fungi
cause diseases) cause diseases to many a crop plants. Peronosporales:- divided
into three families :-
1. Pythiaceae - Pithium,Phytophthora
2. Peronosporaceae-Plasmopara
3. Albuginaceae -Albugo.
Peronosporales differs from the Saprolegniales in producing only secondary
zoospores in a zoosporangium
That is differentiated from hyphae (eucarpic) and one oosphere (egg) per
oogonium.
Zoosporangia often deciduous

image1410

Zoospores often formed in vesicle


They are aquatic, amphibious, terrestrial and some of the most destructive plant
pathogens .

The most economically important group of Oomycetes is the Peronosporales that


contain the late blight of potato fungus Phytophthora infestans and relatives such as
Peronospora, Bremia, Plasmopara and others that cause “downy mildews”, the “damping
off” fungi, Pythium spp., and the white rust fungi, Albugo spp.

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However, although members of Mastigomycotina are morphologically similar, they share
no close phylogenetic relationships to fungi. The combination of cellulose cell wall,
biflatellated zoospores, one flagellum of the tinsel type and the other of the whiplash
type, and gametangial copulation are characteristics that are shared with some
members of the algal divisions Phaeophyta and Chrysophyta. This has recently led to
recognition of yet another kingdom, Stramenopila, which includes the divisions
Bacillariophyta, Chrysophyta, Phaeophyta, Hyphochytridiomycota and Oomycota. These
divisions are now thought to be derived from a common ancestor .

Section "D"

Sub-division : -Zygomycotina

Most of the Zygomycotina are present in soil and dung, occurring mostly as
saprophytes; few are parasitic on plants and animals. About 1000 fungal species
belong to Zygomycotina.
Vegetative ( somatic) body is Haploid .
Thallus is usually mycelial, hyphae coenocytic.
Cell wall is made up of chitin and chitosan.
Asexual reproduction occurs most commonly by the formation of nonmotile,
unicelled sporangiospores in uni- or multispored sporangia or merosporangia. In
addition, arthrospores, chlamydospores, and yeast cells can be formed by some
species.
These characteristics are shared with the divisions of flagellated fungi
(Mastigomycotina).
Spores are dispersed either violently or passively by wind, rain or animals.
Flagellated spores and gametes are absent in this division as well as in the
remaining taxa of terrestrial fungi.
Sexual reproduction occurs with the fusion of two multi-nucleate isogametangia or
anisogametangia to produce a zygote.
The zygote later develops into a thick-walled zygospore, the diagnostic feature of
this division.
Because of this the fungi of the class zygomycetes are also known as conjugation
fungi.
Two classes are recognized in this division :-
1. Trichomycetes
2. Zygomycetes.

Zygomycetes :- The members of this class are mostly saprobic(saprophytes), though a


few may become weakly parasite on plants.The common examples are black bread
moulds or pin moulds (the genera Rhizopus and Mucor). In the class, Zygomycetes there
are three orders:-

Mucorales
Entomophthorales
Zoopagales
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Order :- Mucorales

Rhizopus (common bread mold)

image1411

This species is one of the most common


members of this class.
Vegetative thallus is eucarpic, branching,
coenocytic (multinucleate) and aseptate (not
divided by cross walls into cells or
compartments) mycelium and haploid (n).
The mycelium of R. stolonifera is differentiated into three different parts :-
1. Stolons:- The mycelium develop aerially above the substratum is called
"stolons".
2. Rhizoids:- Stolons produce rhizoids in a cluster from below into substratum.
Branching rhizoids behave as roots, anchoring the fungus into its substrate,
releasing digestive enzymes, and absorbing nutrients for the fungus.
3. Sporangiophores: -Sporangia form on the tips of sporangiophores, which are
erect branches formed directly above the nodes of rhizoids.

Internal structure The hyphae cell wall are made up of chitin lined by a thin plasma
membrane. The protoplasm includes typical cell organelles like nuclei, mitochondria,
ribosomes, endoplasmic reticulum etc.

Life cycle of Rhizopus stolonifer

Rhizopus stolonifer Life cycle includes both asexual reproduction and sexual.
Asexual reproduction :- Asexual reproduction during favourable condition.
Asexual reproduction takes place by the formation of sporangiophores with
terminal uni-to-multispored sporangia.
Sporangiophores arise from mycelium.
Each sporangium begins as a swelling into which a number of nuclei flow, and it is
eventually cut off from the sporangiophores (Aplanospores) by the formation of a
septum.
The protoplasm within is cleaved, and a cell wall is in the sporangium. Which
divides sporangium into an outer fertile sporiferous zone and a central sterile
columella.The protoplasts of the sporiferous zone cleave mitotically to form dark-
coloured multinucleate spores.
In this process, nonmotile, single-celled, haploid sporangiospores are formed.
The sporangium becomes black as it matures, giving the mold its characteristic
colour. Each spore, when liberated, can germinate to produce a new mycelium.

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image1409

Sexual reproduction :- R. stolonifer is different from all other fungi because it


reproduces by sexual reproduction via zygospores following gametangial fusion
Sexual reproduction occurs only between different mating strains such fungi are
known as heterothallic.
Which have been traditionally labeled as + and - types. But the mating strains are
morphologically indistinguishable,.
Two hyphae of opposite strains come closer, hormones cause their hyphal tips to
come together.These hyphae are called zygophores.
The tip of zygophores develop into gametangia, which become separated from the
rest of the fungal body by the formation of septa.
The walls between the two touching gametangia dissolve, and the two
multinucleate protoplasts come together.
The + and - nuclei fuse in pairs to form a young zygospore with several diploid
nuclei.
The zygospore which can become dormant for several months.
Meiosis occurs in the diploid nuclei of zygospore.It results in the segregation of
separate '+' and '-' nuclei all the nuclei so formed disintegrate except one.
After a long period of rest, the wall of the zygosporangium cracks. And produces a
sporangium that produces spores same as the asexually produced sporangium,
and the life cycle begins again.

Section "E"

Sub- division :- Ascomycotina(sac fungi)

Members of the Ascomycotina are known as the Sac Fungi. The majority of fungi that
lack morphological evidence of sexual reproduction are placed here. Examples of sac
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fungi are yeasts, morels, truffles, and Penicillium. This includes the salmon-coloured
bread mold Neurospora sp., which has played an important role in the development of
modern genetics. Important features of Ascomycotina:-
These fungi posses well-developed, profusely branched mycelium except the
unicellular yeasts.
Hyphae with regular cross-walls called septa and haploid.Which are centrally
perforated to allow movement of cytoplasm, and sometimes nuclei, between
compartments.
The hyphal cells of the vegetative mycelium may be either uninucleate or
multinucleate.
Cell walls are composed mostly of chitin.
All produce an ascus (sac-like structure) that contains haploid (n) ascospores after
meiosis .
Plasmogamy is separated from karyogamy in time so that a dikaryotic phase is
produced – the ascogenous hyphae represent the dikaryotic hyphae(or at least in
those which produce a teleomorph).
Sexual life cycle is basically similar – haploid-dikaryotic life cycle.
Some species of Ascomycota can self-fertilize and produce sexual structures from
a single genetic strain; others require a combination of + and - strains.

Nutrition and Symbioses :-Ascomycotina are heterotrophs and obtain nutrients from
dead or living organisms :- As saprotrophs they can consume almost any carbonaceous
substrate such as cellulose (found in plant cell walls), lignin (found in wood) and recycled
dead plant material .

As parasites, ascomycetes account for most of plant pathogens including powdery


mildews that attack fruits, chestnut blight, and Dutch Elm disease (caused by Ceratocytis
ulmi).
Lichens- Symbiotic association between a sac fungus and a photosynthetic green algae
or cyanobacteria
Mycorrhizae :- Symbiotic association of a sac fungus living on plant roots.
Endophytes:- Symbiotic association of a sac fungus with the leaves and stems of
plants.
Reproduction and life cycle Fungi are Holomorphs.

Holomorph a fungus that is characterized both by sexual and asexual reproductive


states.
Teleomorph :-the fungus when reproducing sexually.
Anamorph :- the fungus when reproducing asexually.

Asexual reproduction :-
Yeasts reproduces by budding or fission.
Asexual reproduction in the majority of the Ascomycetes occurs by the
formation of specialized spores, known as conidia.
Which are formed on tips of modified hyphae called conidiophores.

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Conidia are formed in longitudinal chains on the conidiophores.
Each conidium contains one or more nuclei.
Conidia form on the surface of conidiophores in contrast to spores that form
within sporangia in Rhizopus.
When mature, conidia are released in large numbers and germinate to
produce new organisms.

image1413

Sexual reproduction :-In these fungi sexual reproduction occurs by the formation
of multinucleate gametangia called :-
Male gametangia may be an antheridium or conidium-like structure –
spermatium.
Female gametangium - ascogonium, may have a long projection, the
trichogyne
Asci formation occur on the same mycelia that produce conidia.
The male nuclei of the antheridium pass into the ascogonium or an ascogonium
and a spermatium ,through the trichogyne.
Plasmogamy or the fusion of the two cytoplasms, has now taken place.
The male nuclei then pair with the genetically different female nuclei within the
common cytoplasm but do not fuse.
Hyphae now begin to grow out of the ascogonium.
As the hyphae develop, pairs of nuclei migrate into them and simultaneous mitotic
divisions occur in the hyphae and ascogonium -dikaryotic cells.
The pairing of two genetically (+ and -) different types of nuclei followed by
septation results in the formation of a number of dikaryotic cells (i.e. containing
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two haploid nuclei, one from each strain).
Sterile haploid hyphae known as peridium envelope the ascogonium to form the
multicellular ascoma or ascocarp(fruiting body).
Many ascogenous hyphae produced inside the developing ascoma.
Tips of ascogenous hyphae form croziers (hooks) before developing into an ascus.
Ascoma contains two types of hyphae :-
Ascogenous hyphae – dikaryotic, form asci through crozier formation.
Sterile hyphae – haploid, form bulk of ascoma.
The two nuclei in the terminal cell (ascus) of the dikaryotic hyphae then fuse into a
single diploid nucleus ("karyogamy"). The zygote is only diploid phase in life cycle.
The ascus then elongates and the diploid nucleus divides by meiosis, forming 4
haploid nuclei.
Each haploid nucleus usually divides again by mitosis, generally resulting in a total
of 8 haploid nuclei but this may vary.
These haploid nuclei are then cut off in segments of the cytoplasm to form
ascospores.
In most Ascomycotina, the ascus becomes turgid at maturity and finally bursts,
sending its ascospores explosively into the air.
Within the ascocarp, there is usually a layer containing the asci, and sterile cells
known as paraphyses. This layer is the hymenial layer.

The form of the ascocarp may be one of four types :-

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image1415

image1412

There are four basic types of ascus:-

Unitunicate-operculate ascus has


single wall, operculum (lid/) at the
top. The operculum pops open when
the ascus becomes mature to release
ascospores. - found in apothecia
Unitunicate-inoperculate ascus has no operculum but instead contains a special
elastic ring at the top. On ripening it briefly expands and so lets the spores shoot
out - found in perithecia and some apothecia.
Prototunicate asci are mostly spherical in shape and they have no active dispersal
mechanism at all. The ripe ascus wall simply dissolves so that the spores can
escape, or it is broken open by other influences such as animals. Found in
cleistothecia, some perithecia, and hypogeous ascomata.
Bitunicate ascus is enclosed in a double wall. This consists of a thin brittle outer
shell and a thick elastic inner wall. At maturity the thin outer wall splits, and the
thick inner wall absorbs water and expands upward, carrying the ascospores with it
. This 'Jack-in-a-box' design allows the ascus to stretch up into the neck of the
ascoma to expel its spores. Bitunicate asci occur only in pseudothecia.

Section "F" Sub-section "1"

Sub division - Basidiomycotina

Basidiomycotina are mostly terrestrial and saprophytic or parasitic. It includes some


25,000 described species. This group includes most of the mushrooms , toadstools,
stinkhorns, puffballs, and shelf fungi. And also contains important obligate parasites,
two important plant pathogens the rusts and smuts.

This sub division shares many features in common with the Ascomycotina:-

Well-developed, branched, regularly septate hyphae.


Hyphae are initially uninucleate but soon become dikaryotic.
Presence of yeast stage and presence of macroscopic fruiting bodies, in some taxa.
Few species form blastic conidia.
Cell walls are usually chitinous.
As in the Ascomycota and Zygomycota, the most important feature is the
production of basidiospores (sexual spore).
The basidiospores that are typically borne, exogenously, on horn-like sterigmata
(singular-sterigma) of basidia (singular- basidium).

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General characters of Basidiomycotina :-The mycelium of the Basidiomycotina in
most species have three distinct phases during the life cycle of the fungus:-

Primary mycelium :- When it germinates, a basidiospore produces haploid primary


mycelium . Initially the mycelium may be multinucleate, but septa soon form and the
mycelium is divided into monokaryotic (uninucleate) cells. This septate mycelium grows
by division of the terminal cell.
Dolipore septa with parenthesome are present in most of the genera.
Allows cytoplasmic movement but prevents nuclear migration from one compartment
to the next.
Secondary mycelium :-Commonly a secondary mycelium forms upon conjugation of
two sexually compatible hyphae ( heterokaryotic).
The dominant phase of the life cycle in most Basidiomycotina is a dikaryon, in which
the two nuclei brought together in mating exist side-by-side in each cell.
But some times, septa are not formed after a mitosis in the terminal cell of primary
hyphae as a results secondary mycelium is formed. Therefore all subsequent cells of the
hyphae are binucleate (dikaryotic).
In both cases, the dikaryotic (binucleate) mycelium is formed, since karyogamy (the
fusion of the gametic nuclei) dose not immediately follow plasmogamy (fusion of the
protoplasts).
As the dikaryotic mycelium grows, the cells divide and more septa are formed
between the new cells.
Each of the new cells in the secondary mycelium has one haploid nucleus from each
parent. This is due to clamp connections, specialized structures unique to the
Basidiomycotina.
>

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These are loop-like hyphae which connect the cytoplasm of adjacent cells and through
which nuclei move during cell division.
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During cell division, one nucleus divides directly into the newly formed cell. The other
nucleus divides inside the clamp connection and the two daughter nuclei migrate
through the clamp connection in opposite directions to the two daughter cells,
reestablishing the dikaryotic condition.
The tertiary mycelium which is also dikaryotic, arises directly from the secondary
mycelium forms the fruiting bodies -basidiocarps.
Reproduction and life-cycle of Basidiomycotina :- Like all fungi, Basidiomycotina can
undergo both asexual and sexual reproduction.

Asexual Reproduction :- Basidiomycotina reproduce asexually by either budding or


asexual spore formation.
Budding occurs when an outgrowth of the parent cell is separated into a new
cell. Any cell in the organism can bud.
The usual asexual spores formed by these fungi are arthrospores, sometimes
called oidia.
Few species form blastic conidia. Conidia (asexual spore) formation, takes
place at the ends of specialized structures called conidiophores. The septae
of terminal cells become fully defined, dividing a random number of nuclei
into individual cells. The cell walls then thicken into a protective coat. The
protected spores break off and are disbursed and germinate.
Oidia are formed on specialised, erect hyphal branches known as
oidiophores. Oidia can have two functions. They may germinate to form
mycelium or they may function in the mating process. Asexual spores are
usually haploid and the hyphae that form after germination are also haploid.
Sexual reproduction :-
No specialized sex organs form in the Basidiomycotina except in the rusts.
The sexuality in Basidiomycotina gets progressively subdued.
The specialized nature of sex organs is lost and the whole process is
ultimately confined to copulation between vegetative mycelia (somatogamy).
The tertiary mycelium, which is also dikaryotic, arises directly from the
secondary mycelium, and forms the basidiocarp. The spore forming
basidia(basidia are not produced by asexual Basidiomycota) are produced by
the terminal cell on millions of dikaryotic hyphae.
Basidiocarps vary greatly in appearance in different genera, but all bear
basidia which are usually arranged in a layer of hymenium.
The basidia and basidiospores are produced on even sufaces or on
hymenophores in form of teeth, tubes or leaf-like structures (gills) at the
inferior side of caps or consoles (crustothecium, pileothecium).
In special parts of the fruit body (hymenia, hymenophores if existing)
karyogamy occurs between the two haploid nuclei within a developing
basidium. Then, the diploid nucleus undergoes meiosis to produce four
haploid nuclei.
Basidiospore formation on the basidium is exogenic (external) which means
that the 4 basidiospores appear on small appendages (sterigma) on top of
each basidium, in contrast to that, ascospore formation is endogenic.
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Basidia of Hymenomycetidae are formed in layers called hymenia.
The hymenia are exposed on special surface enlarging structures
(hymenophores) for spore discharge.
One of the most important characteristics of Basidiomycotina is the
production of forcibly discharged ballistospores which are propelled into the
air from the sterigma.
Ballistospores are sexual or asexual, and produced by basidia, hyphae, yeast
cells, or even other ballistospores.
This type of spore discharge have been evolved very early in the evolutionary
history of the Basidiomycotina as it is found in members of the earliest
diverging lineages within the group.
Ballistospory is associated with forms that disperse their spores directly into
the air. Most aquatic Basidiomycotina and forms that produce spores inside
the fruiting body, such as puffballs, have lost ballistospory.
Basidiocarps is generally divided into 3 parts :-
Hymenium The typical hymenium consists of parallel arranged fertile
elements: young basidia (basidioles), basidiospore forming basidia in
various stages of developement and also sterile elements (cystidia).
Subhymenium – Below the hymenium a layer of isodiametric cells is
present which is the subhymenium
Trama – The layer on which the subhymenium and the hymenium is
found to be located is the trama. The trama is the main hyphal
structure within the hymenophore (hymenophoral trama) and within
the whole fruit body.
Fruit bodies (Basidiocarps) of the different species show a great variety of different
shapes :-

Holothecium:- No clear delimination of spore producing and sterile parts in


the fruit body which is cushion-like, club-shaped (clavate), coralliform
(coraloid) or irregularily lobed.
Crustothecium : -A fruit body which is more or less completely adnate to the
substrate.
Pilothecium :-a fruit body which is differentiated in a stipe and a cap.

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Section "F" Sub-section "2"

Rusts:- Rusts (Pucciniales, previously


known as Uredinales) shows greatest
complexity in their life-cycle. Five
different types of spores are formed on
two different hosts in two unrelated
host families. Such rusts are
heteroecious (requiring 2 hosts) and
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macrocyclic. Autoecious rusts complete their life-cycles on one host intead of two.
In order to complete its life history, this species produces the following five spore
stages:-

1. Spore Stage 0: Spermogonia (sing.=spermogonium) (consists of Receptive


hyphae and Spermatia [sing.=spermatium]) :-Generally , basidiospores infect
the alternate host the barberry , the mycelium forms pycnidia, called
spermagonia. which are miniature, flask-shaped, hollow, submicroscopic
bodies embedded in host tissue (upper surface of a leaf). This stage,
numbered "0", produces single-celled, minute spores that ooze out in a sweet
liquid and that act as nonmotile spermatia, and also protruding receptive
hyphae. Insects and probably other vectors such as rain carry the spermatia
from spermagonia to spermagonia, for crossing between the mating types.

image1418

2. Spore Stage I: Aeciospores in aecia (sing.=aecium) :-After crossing , the


dikaryons are formed and a second spore stage is formed, numbered "I" and
called aecia. The aecia with the dikaryotic aeciospores are formed in dry
chains in inverted cup-shaped bodies embedded in the lower surface of the
barberry leaves. These aeciospores then infect the second host genus and
cannot infect the host on which they are formed (in macrocyclic rusts).
3. Spore Stage II: Urediospores in uredia (sing.=uredium) :- During the early
spring ,the aeciospores infect the wheat plant. Infection of the wheat occurs
in both the stem and leaves. Entry into the host occurs when the spores
germinates and enters the plant through openings called stomata. After
germination of spores, the mycelium in the host plant gives rise to clusters of
the uredospores in dry pustules called uredinia. On the second host a
repeating spore stage is formed, numbered "II", As the urediospores develop,

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they will burst the epidermis, exposing the characteristic, rusty-colored
urediospores on the surface of the plant . The urediospore can infect other
wheat plants throughout the spring and early summer.
4. Spore Stage III: Teliospores in telia (sing.=telium) :- During late summer, the
uredium gradually converts into the telium and begin to produce the two-
celled, thick-walled teliospores a fourth spore type . The rusty-brown uredium
becomes black as the teliospores are borne.The teliospore stage, with its
thick wall is the over winter stage and will remain dormant for the winter.
5. Spore Stage IV: Basidiospores on basidia (sing.=basidium) :- Under favorable
condition, each cell is capable of germinating to produce basidia and
basidiospores.In the Pucciniales, the basidia are cylindrical and become 3-
septate (four celled) after meiosis, with each of the 4 cells bearing one
basidiospore each. The basidospores disperse and start the infection process
on host first again.
On the basis of life cycle patterns, the rusts are of three types :-

Macrocyclicrusts -Producing all 5 spores types.


Demicyclic rusts in which uredial stage absent
Microcyclic rusts Only two types of spores are formed -teleutospores and
basidiospores)
Section "G"

Sub- division - Deuteromycotina (Fungi Imperfecti)

The Deuteromycota are characterized by a well-developed, septate mycelium,


some are siphonaceous.
Cell walls: Usually chitin and glucan.
Asexual reproduction is by means of conidia (sing.=conidium) or may be
lacking.
Sexual reproduction is not known; thus these are the "imperfect Fungi."
This includes unrelated fungi from the Ascomycotina, Basidiomycotina and
even the Zygomycotina. Members of different fungal division may produce
conidial states which are morphologically very similar and classified in the
same genus. Because any such genus may include such unrelated taxa, it is
called a "form-genus."
Many forms of Deuteromycota are pathogenic, affecting man, animals, or
plants. Example Alternaria tenuis, Sporobolomyces.

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