Characteristics of Fungi - Part I

General Characteristics of True and Allied Fungi –Part I
DC-1
SEM-2
PAPER-3: Mycology and Phytopathology
Lesson: General Characteristics of True and Allied Fungi –

Part I
(Vegetative Structures)
Lesson Author: Dr. Inderjeet Sethi and Dr. Charu Kalra
College/Dept: Sri Guru Tegh Bahadur Khalsa College,

Department of Botany
Deen Dayal Upadhyaya College, Department of Botany
University of Delhi
Institute of Lifelong Learning, University of Delhi 1

Table of Contents
Chapter: General Characteristics of True and Allied Fungi – Part I
(Vegetative Structures)
 Introduction
 Fungal Nutrition
 Saprobes
 Parasites
 Types of Parasites
 Necrotrophic Parasites
 Biotrophic Parasites
 Fungal Parasites on Humans
 Symbionts/Mutualists
 Lichens
 Mycorrhizae
 Endophytes/Ectophytes
 Infection Structures
 Factors Influencing the Development of Infection Structures
 Growth of Fungi
 Growth Forms of Fungi
 Dimorphic Fungi
 Mycelium- The Body of Fungus
 Types of Septa
 Roles of septa
 Chemical Composition of Cell wall

 Architecture of Fungal Cell wall
 Functions of Fungal wall
 The Plasma membrane

 Roles of Plasma Membrane
 Nuclei and associated Structures

 Nuclear Division

 Spindle Pole Bodies

 Nucleolus
 Fungal Organelles
 Organization of Fungal Apex
 Factors affecting Fungal Growth
 Light
 Water
 Aeration
 Temperature
 Hydrogen Ion Concentration
 Summary
 Glossary
 Exercises
 References

Introduction
This chapter explains all characteristics shown by true fungi as well as their allies. As the
majority of the characteristics remain same. Table below (modified from Ainsworth, 1973)
lists major characteristics of such organism and explained later.
Occurrence Diverse
Habitat Commonly present in terrestrial and freshwater habitats and
rarely in the marine environment
Ecology Important ecologically as saprobes, parasites, hyperparasites
or mutualists
Nutrition Heterotrophic, feeding by absorption rather than ingestion
Thallus Unicellular (yeasts) or typically filamentous, non-motile
mycelium of hyphae, on or in the substratum. Hyphae showing
internal protoplasmic streaming and apical growth
Cell Wall Well defined, typically containing glucans and chitin; cellulose
and glucans in Stramenopila
Nuclear status Eukaryotic, uni- or multinucleate, mycelium homo- or
heterokaryotic, haploid, dikaryotic; diploid in Stramenopila
Life cycle Simple, or more commonly, complex
Reproduction Sexual (i.e. nuclear fusion and meiosis) and/or parasexual
(nuclear fusion followed by de-diploidization) and/or asexual
(i.e. only mitotic nuclear division)
Propagules Typically nonmotile, microscopic, spores produced in large
numbers. Motile spores formed in certain groups (e.g. chytrids)
and in Stramenopila
Sporocarps Microscopic or macroscopic with limited tissue differentiation
Nuclear division Mitosis is intranuclear, i.e. it takes place without the

dissolution of nuclear envelope
Reserve food Glycogen, lipids or trehalose; mycolaminarin in Stramenopila
Majority of the fungi grow in warm, moist environments, however, there are many species
which occur in habitats that are extremely cold, hot and dry, or otherwise seemingly

inhospitable. Although, the optimum conditions for growth and reproduction vary widely
with the species, still we find maximum diversity of fungi in tropical areas.
General Characteristics
Fungal Nutrition
Similar to animals, fungi are heterotrophs, i.e. dependent upon already produced organic
compounds and energy sources for survival.
But unlike animals, which ingest and then digest their food, fungi first digest food by the
production of enzymes outside their body (exoenzymes) and then ingest it by absorption.
Exoenzymes (also found in some bacteria) break down large and relatively insoluble
molecules such as carbohydrates, proteins and lipids into smaller and more soluble
molecules, which can easily be absorbed through the wall and cell membrane of fungi.
Presence of free water, however, is essential for the diffusion of exoenzymes and nutrients
across the fungal cell wall and membrane. This might be the reason that actively growing
fungi are mainly restricted to humid places.
Based upon how they obtain nutrients, they have been broadly divided into three
categories:
I. Saprobes
II. Parasites
III. Symbionts/Mutualists
IV. Endophytes/Ectophytes
I. Saprobes (Gr. sapros = rotten; bios = life)
 Also known as saprophytes or saprotrophs.

 Fungi growing and obtaining their nutrition from dead organic materials.
 They can attack almost any existing substance wherever adequate moisture and
temperature are present. Rhizopus, Aspergillus (black mould) and Penicillium (green
mould) are, a few very commonly encountered species on all type of substrata.
 Fungi grow inside the substrata with their absorptive filaments and can invade them
in dark places as well as in lighted places. Such decomposer fungi, which bring about
spoilage of useful products, are known as biodegraders.

 Saprobes along with some bacteria help in recycling of organic matter in natural and
agricultural environment.
II. Parasites (Gr. parasitos = table mate)
 A parasite obtains some or all of its nutrients from the living tissues of another
organism while being in close relationship with it.
 Along with living protoplasm of the host they may be highly specialized to parasitize
a particular species.
 A parasite that causes disease is termed a pathogen. The notable fungal pathogens
of crop plants worldwide are downy mildews (Oomycota), powdery mildews, rusts,
smuts and vascular wilt diseases.
 Majority of the parasitic fungi are also able to grow on dead and decaying organic
matter. The latter can be confirmed by their ability to grow on artificial synthetic
media. Such fungi are known as facultative saprobes/saprotrophs (L. facultas =
ability).
 Similarly, there are saprobic organisms, which are able to infect another living
organism under some conditions. These are known as facultative parasites.
 Those fungi that are unable to grow on artificial media even after the best of efforts
and remain parasitic in nature are known as ecological obligate parasites or
biotrophs.
Value Addition
Why fungi are important as plant parasites in comparison to bacteria?
This is because the hyphal apex has a remarkable penetrating power,

which enables hyphae to invade even intact plant surfaces. Bacteria,
on the other hand, cannot penetrate intact surfaces, but often need
wounds or natural openings to do so.
Types of parasites

Based on the way they obtain their nutrients parasites are of two types: Necrotrophic and
Biotrophic. Both necrotrophs and biotrophs may be parasitic on plants, animals or on other
fungi.
1. Necrotrophic parasites (Gr. nekros = death; trophein = to nourish), are also

known as necrotrophs or perthotrophs. This type kills the living host cells in
advance of its entry, by producing toxins or degradative enzymes and then absorbing
released nutrients and growing between and into dead and dying cells. They can
attack plants (e.g. Botrytis fabae (Ascomycota) on leaves of Vicia faba), insects (e.g.
Lagenidium; Oomycota), Coelomomyces (Chytridiomycota), Cordyceps (Ascomycota)
or other fungi. The latter are known as necrotrophic mycoparasites or
hyperparasites. Some necrotrophs may be host-specific and thus can be used as
biocontrol agents.
Value Addition
DID YOU KNOW?
o Majority of Pythium spp. (Oomycota) are plant pathogens. Only about

six species of Pythium are non-pathogenic to plants but parasitize
other fungi.
o Many Pythium spp. can be used as biocontrol agents of plant
pathogens, especially as seed treatments for control of seedling
pathogens.
2. Biotrophic parasites (Gr. bios = life) are also known as biotrophs. These
parasites obtain some or all of its nutrients from the living cells or tissues of host,
usually by means of special nutrient absorbing structures - haustoria (sing.
haustorium; L. haustor = drinker). Majority of the biotrophs cannot be grown in
axenic culture. Majority of the biotrophs belong to Division Oomycota, Ascomycota,
Basidiomycota and class Trichomycetes. Like necrotrophs, biotrophic fungi can attack
plants (e.g. rust fungi (Puccinia graminis) and powdery mildew (Blumeria graminis),
insects (e.g. Laboulbenia spp), as well as other fungi (e.g. Piptocephalis, Dispira
(Zygomycota) commonly parasitize other members of Zygomycota such as Mucor
and Pilaira). These are known as biotrophic mycoparasites. Like necrotrophs,
biotrophs can also be used as biocontrol agents, e.g. Black scurf of potato caused by

Rhizoctonia solani (Basidiomycota) can be controlled by Verticillium biguttatum

(Deuteromycota).
Value Addition
DID YOU KNOW?
Some pathogens start their parasitic phase as haustorial biotrophs, but then kill
the host tissues and spread within them as necrotrophs. Such pathogens are
known as hemibiotrophs. Example: Phytophthora infestans on potato and P.
sojae on soybean (Oomycota).
3. Fungal parasites of Humans
 A few fungi parasitize humans and most of those infecting humans are
opportunistic parasites.
 These grow more commonly as saprobes in soil, composts, bird excreta, etc., but
can infect through wounds or through lungs.
 Most of the spores of such parasites are air borne, e.g. Aspergillus fumigatus,
Histoplasma capsulatum, and Blastomyces dermatitidis.
 Generally, all these pathogens pose a threat to debilitated individuals suffering
from life threatening diseases or to transplant patients. For example, fungus
Pneumocystis carinii is the most common cause of death of acquired immune
deficiency syndrome (AIDS) patients in several parts of the world.
III. Symbionts or Mutualists
In this kind of association, the fungi absorb nutrients from other organisms but are not
detrimental to them. Such fungi are known as mutualistic fungi. The association can be:
(i) Lichens. This is an association of fungi (mycobiont) with algae and cyanobacteria
(photobiont). Although lichens may seem infrequent in polluted cities, they can
form the dominant vegetation in unfavourable environments. Within this
symbiosis, the mycobiont gets carbohydrates from algae and cyanobacteria and
fixed nitrogen from cyanobacteria; the photosynthetic organisms apparently
receive nutrients, protection and a receptive substrate for growth from the
surrounding fungus.

(ii) Mycorrhizae. This is an association of fungi with roots of higher plants. The
fungal symbiont in a mycorrhiza gets carbohydrates from plant host, and the
plant gets minerals from the fungus.
IV. Endo or Ectophytes
These are fungi present in or on the leaves & stem of healthy plants and animals. Some
such endophytic fungi may protect both conifers and grasses against insect herbivory,
influence flowering and many of these fungi produce secondary metabolites, which may
be antibacterial or antifungal. Endophytes may also be present on woody perennials,
mosses, hepatics and pteridophytes. Alternaria alternata and Cladosporium
cladosporioides are commonly found as ectophytes, but are also capable of internal
colonization of healthy tissue.
Fungi, which are present within or on host animals, may or may not cause any damage
to the animals. Many fungi belonging to Ascomycota and Basidiomycota, are associated
symbiotically with arthropods, wood wasps, ambrosia and bark beetles, leaf cutting ants
and termites.
Value Addition
INTERESTING FACTS
1. Ants have been associated with fungi from about 50 million years. Attine ants take green
leaves inside their nest, finely shred these leaves and neatly stack them. Development of
mycelium on this organic substratum makes up the food of the ants.
When the ants move to new areas, the fungus is carried along, thus ensuring its dispersal.
The dependence of the fungus and the ants is mutual; the ants can't survive without the fungus
as it is a direct source of nutrients and also the fungal cellulases make the ingested plant
materials easily digestible.

Figure: Ant-fungus relationship

http://upload.wikimedia.org/wikipedia/commons/c/cb/Atta_colombica_queen.jpg
2. Ambrosia beetles are found in the tunnels of ambrosia trees in association with yeasts known
as 'Ambrosia fungi'.
These fungi besides providing nutrients also provide ergosterol that is essential for larvae to
develop into adults.
They are carried in specialized pouches that range from small depressions in the insect cuticle to
larger glandular structures that produce secretions probably nourishing the fungal cells.
These fungus filled pouches are known as mycangia.
Figure: Pupae of Ambrosia beetles, Xylosandrus crassiusculus with black fungi
Source: http://upload.wikimedia.org/wikipedia/en/7/76/Xylosandrus_crassiusculus_galleryR.jpg

Infection Structures
Infection structures are specialized structures formed in parasitic fungi at the tips of
germ tubes or hyphae on the outside of the host. The simplest pre-penetration
structures are terminal swellings on germ tubes; called appressoria (sing.
appressorium). Appressoria produced by rust fungi, and those of Magnaporthe grisea,
the fungus that causes rice blast disease have been studied most extensively.
Infection structures are formed when the spore (s) germinates; producing a germ tube
(GT), which align itself, locate a stomatal ridge and stops growing. Then the apex swells
to form an appressorium (A). A septum separates the appressorium from the germ tube.
Appressorium adheres the fungus to the host surface, usually by secretion of a
mucilaginous matrix.
After about 2 hrs of its contact with the stoma, a penetration peg (PE) grows into the
substomatal cavity. Appresorium may secrete enzymes like cutinase helping penetration
peg enter the host through epidermal cell wall directly or through stomatal opening.
Figure: Obligate biotrophic Uromyces spp. A germ tube (GT) emerges from an urediospore
(S) attached to the host by an adhesion pad. After recognition of the guard cell lip, an
appressorium (A) develops over the stomatal pore. The penetration hypha (PE) penetrates
into the substomatal chamber and elongates into an infection hypha (IH). When the tip of
the infection hypha contacts a host cell wall, a haustorial mother cell (HM) is formed from
which the haustorium (H) invades the host cell. Unique features of the dikaryotic
haustorium are the dark-staining neckband (NB) around the haustorial neck and the
interfacial, extrahaustorial matrix (yellow) surrounded by the extrahaustorial membrane

(EHM). After forming the first haustorium, the infection hypha branches and further
intercellular hyphae, haustorial mother cells (HM) and haustoria are formed.
Source: http://www.uni-konstanz.de/FuF/Bio/AGMendgen/Biologie/ralf_background.htm
Penetration peg further produce a substomatal vesicle and an infection hypha (IH)
develops from this to produce a haustorium mother cell (HM) on the parenchyma of
the leaf.
Formation of haustorium - a specialized nutrient absorbing structure - completes the
infection process. The name haustorium (L. haustor = the pail) was introduced by the
German physician Heinrich Anton de Bary. Unique features of the dikaryotic haustorium
are the dark-staining neckband around the haustorial neck and the interfacial,
extrahaustorial matrix enclosed by the extrahaustorial membrane.

Figure: Haustorial complex, a specialized feeding organ of biotrophic fungal parasites of

plants. To move from host cell to fungus, nutrients must traverse the extrahaustorial
membrane, the extrahaustorial matrix, the haustorial wall, and the haustorial plasma
membrane. A neckband seals the extrahaustorial matrix from the plant cell wall region so
that the matrix becomes a unique, isolated, apoplast-like compartment. The haustorium
connects to intercellular fungal hyphae by way of a haustorial mother cell. A: Transmission
electron micrograph image of a haustorium (Szabo & Bushnell. 2001. PNAS USA 98:7654-5;
B: Diagrammatic view showing ultrastructure.http://www.uni-
konstanz.de/FuF/Bio/AGMendgen/Biologie/ralfbackground_image21.jpg
Factors Influencing the Development of Infection Structures
Development of the appressorium have been found to be influenced by physical and

chemical factors, but one absolute requirement is contact with a surface of sufficient
physical hardness, such as leaf cuticle, insect cuticle or artificial surfaces. So, it can be
concluded that it is a thigmotropic kind of response or contact sensing.
The formation of haustorial mother cell usually depends on chemical recognition of a
leaf cell wall showing that most of this developmental process is programmed and it
requires only an initial leaf surface signal.
Growth of fungi

Fungal germ tubes and hyphae grow almost exclusively at their tips, i.e. they show
apical growth. The term germ tube refers to the young hypha that emerges from a
germinating spore and branches.
The original hypha and the first formed branches repeatedly branch behind their tips.
These branches separate from each other until finally the colony develops a typical
circular outline.
Figure: Stages in the development of a fungal colony from a germinating spore. The
broken lines in (e) represent narrow anastomosing hyphae near the centre of the colony.
Source: http://archive.bio.ed.ac.uk/jdeacon/FungalBiology/chap3a.htm
In the older parts of the colony where nutrients have been depleted, mycelia grow
towards one another and fuse by tip-to-tip contact producing narrow hyphal branches.
This leads to localized breakdown of their walls and continuity of the cell cytoplasm. This
process is known as hyphal anastomosis leading to the development of typical
interconnected network of hyphae observable in higher fungi.

Figure: Anastomoses in fungi
Source: http://www.uoguelph.ca/~gbarron/MISCE2002/anast01.jpg
Figure: Videotaped sequence of anastomosis (successful hyphal fusion) of two

compatible hyphae of Rhizoctonia solani. The times shown are minutes after the start of
recording. The upper hypha had stopped growing at t 0 but began to produce a branch
(arrowhead) at t6 in response to the approaching hyphal tip. The hyphal tips met at t 14.
Dissolution of the tip walls and complete hyphal fusion was achieved at t 32.
Source: archive.bio.ed.ac.uk/jdeacon/FungalBiology/chap3a.htm (Courtesy of P.
McCabe, M.P.Gallagher & J.W.Deacon, The University of Edinburgh)
In lower fungi the vegetative hyphae do not anastomose.

Diffused (intercalary) type of growth common in plant tissues is rare in fungi and
seems to be limited to hyphae involved in lifting reproductive structures into the air for
spore dispersal.

Growth Forms of Fungi
There are three major growth forms of fungi:
(i) The thallus or the body of the fungus typically consists of microscopic, tubular,
threadlike hyphae that branch in all directions, spreading over or within whatever
substrate the fungus uses for food. The collection of hyphae is known as mycelium
and such thalli are commonly known as 'moulds'.
(ii) Some of the more primitive fungi, such as the Chytridiomycota, often have single
rounded cells or dichotomously branched chains of cells, attached to a food source
by tapering rhizoids.
(iii) Many other fungi grow as unicellular yeasts and produce daughter cells either by binary
fission, e.g. Schizosaccharomyces pombe, or budding, e.g. Saccharomyces cerevisiae.
Intermediate stages between yeast cells and true hyphae also occur and are known as
pseudohyphae. The term yeast is a morphological term and does not refer to a
particular taxonomic group.
Source: http://www.fungionline.org.uk/images/1intro/yeast1.JPG
Dimorphic Fungi

 Certain fungi can exist as either mycelial (M) or yeast-like (Y) phase and are said to
be dimorphic. This ability to switch between these two phases is termed
dimorphism. Important pathogens of humans, higher animals and some plants
exist as dimorphic fungi. They grow as yeasts in body fluids or in water films, but
convert to hyphae for invasion of the tissue or outside their hosts. For example,
Candida albicans grows in yeast form on the moist mucosal membranes of humans,
but changes to hyphae for invasion of host tissues.
Figure: Candida albicans a dimorphic fungus that normally grows as budding yeast
and under appropriate conditions produces mycelia.
Source: http://archive.bio.ed.ac.uk/jdeacon/FungalBiology/chap1_im.htm#Fig.1.4a
 Similarly, insect pathogens, Metarhizium and Beauveria, penetrate the insect cuticle
by hyphae, but then form single cells in the circulating fluids of the host. The
vascular wilt pathogens of plants (e.g. Fusarium oxysporum) penetrate plants by
hyphae but then spread as yeast-like form in the vessel elements.
 The reason for such behaviour is that yeasts have little or no ability to degrade
cellulose, and also do not have any penetrating power. However, the mycelial fungi
commonly have these abilities.
 Various physical factors (e.g. temperature, oxygen, carbon dioxide) and chemical
factors (calcium, nutrients) often influence the conversion from mycelial (M) to
yeast-like (Y) phase and vice versa.
 Dimorphism may also be genetically controlled. For example, Ustilago maydis and
other plant pathogenic smut fungi are in yeast phase when monokaryotic but
become hyphal in the dikaryotic phase.

Mycelium - the body of the fungus
As already noted, the body of the fungus is composed of hyphae and the mass of branched,
tubular filaments (hyphae) constitute mycelium.
 A thin, usually transparent, tubular wall lines a fungal hypha. The interior of the
hypha is filled or lined with a layer of protoplasm.
 Hyphae of majority of the species are interrupted at some places by partitions, or
cross-walls, called septa (sing. septum). Such hyphae are known as septate and
are present in Ascomycota, Basidiomycota and Anamorphic fungi.
 The septate fungi commonly have several nuclei in the apical compartment, but often
only one or two in each compartment behind the apex. In some species septa are
produced at more or less regular intervals along the length of a hypha, dividing it
into individual compartments or cells. Individual compartment may contain one
nucleus (monokaryon), or two nuclei (dikaryon), or many nuclei (multinucleate).
 In other species, the vigorously growing portions of hyphae lack regularly spaced
septa. Such mycelia are known as aseptate/nonseptate/coenocytic and are
observed in Zygomycota and Oomycota. In such mycelia, septa are present only at
the bases of reproductive structures and in mature, highly vacuolate portions of
hyphae.
http://science.kennesaw.edu/~jdirnber/Bio2108/Lecture/LecBiodiversity/31_Labeled_Image
s/31_03-HyphaeTwoForms-L.jpg
Types of Septa

Basically, there are two types of septa - simple and complex - both types appear to form
by the centripetal growth from the hyphal wall inward.
(i) Simple septum can be of two types:
a) Complete septum: Such septa are present at the bases of asexual and
sexual reproductive structures and in older, highly vacuolated portions of
aseptate hyphae.
http://www.fungionline.org.uk/images/3hyphae/septa1.gif
b) Incomplete septum: This kind of septum possesses a single large central

pore of 0.05-0.5µm in diameter, which allows the passage of cytoplasmic
organelles and even nuclei, showing that the protoplasts of adjacent hyphal
compartments are continuous. Typically, the Ascomycota and Anamorphic
fungi have a septum of this type. Variations of this can be observed in
individual fungi; for example, Geotrichum candidum and several unrelated
fungi, have septa with multiple, small micropores or plasmodesmata-like
channels.

(ii) Complex septum: This kind of septum is characteristics of Basidiomycota. It has

a narrow central channel of about 100-150 nm diameter and is bounded by two
flanges of amorphous wall material. On either side of this septum there are
bracket-shaped membranous structures termed septal pore cap or
parenthosome, which have pores that allow cytoplasmic continuity but prevent
the movement of major organelles. This type of septum is known as dolipore
septum (L. dolium = a large jar or cask, i.e. barrel).
This helps to ensure the regular distribution of nuclei in dikaryotic hyphae of the
Basidiomycota. These septa are selectively degraded when the fungus begins to
form a fruiting body, enabling the mass movement of materials to support the
development of these large structures. Depending upon the species involved, the
septal pore cap may be perforate or imperforate.
Roles of Septa
The roles of septa have been debated for many years and are still not completely
understood.
(i) They might help to provide structural support to hyphae, especially in

relatively dry conditions, and in this respect the septate fungi in general
are much more tolerant of water stress than are aseptate fungi.
(ii) Septa also can help to defend against damage, because Woronin bodies or
other materials plug the septal pores if hyphae are physically disrupted or
are damaged by other organisms.

Source: http://www.fungionline.org.uk/images/3hyphae/septa2.gif
(iii) The main significance of septa is thought to be in differentiation. If a

septum is plugged it enables a fungus to transform from a continuous
series of compartments to a number of independent cells or regions that
can undergo separate development by differential gene expression or
biochemical activity, whereas, this would be difficult to achieve if the
protoplasm were continuous.
 Most complex and highly differentiated sporulating structures are

produced by fungi belonging to the Basidiomycota, which possess
the most complex types of septa.
 The aseptate fungi also produce complete cross-walls to isolate old
parts of the hyphae or to produce asexual or sexual structures.
Chemical Composition of Fungal Cell Walls
Chemical analysis of fungal walls can vary considerably between and within different groups
of fungi, however, the basic components remains the same. The major wall components can
be categorized basically into two types:
(i) The structural (fibrillar) polymers, consisting predominantly of straight-chain

molecules, which are cross-linked, providing structural rigidity. It forms the inner
layer of the wall.
 The structural components typically have chitin, chitosan and glucans (polymers of
glucose) as their major wall polysaccharides.
 Chitin consists of long, straight chains of -1,4-linked homopolymer of N-
acetylglucosamine residues occurring in a microcrystalline condition.

Figure: Chitin
Source: http://en.wikipedia.org/wiki/File:Chitin.svg
 Chitosan is a poorly acetylated or nonacetylated form of chitin.
Figure: Chitosan
Source: http://en.wikipedia.org/wiki/File:Chitosan2.jpg
 Chitin is the most important component of cell wall of Chytridiomycota,

Ascomycota and Basidiomycota. It is also a major component of the exoskeleton of
insects and other arthropods.
 It is synthesized in a growing hypha at the hyphal tip, at branching points and in
developing septa.
 Chitin along with chitosan (a major component) is present in the hyphal walls of
Zygomycota.
 However, in Oomycota, cellulose instead of chitin is the main constituent of the
primary wall. Small amount of chitin has been reported in some genera of
Oomycota. For details, readers are referred to see chapter on Oomycota.

(ii) The matrix components, consisting of amorphous or gel-like or crystalline material

found predominantly in the outer layer, covering and embedding the structural (fibrillar)
polymers.
 The amorphous component typically includes proteins, mannans and glucans.

 Proteins have two kinds of roles, as enzymes, they are involved in the synthesis
or lysis of the cell wall or in extracellular digestion, and also function as structural
proteins.
 Glycoproteins, especially mannoproteins, galactomannoproteins and
xylomannoproteins may be present as the major amorphous constituent in many
fungal walls. These may act as important surface antigens in Aspergillus flavus
and Candida albicans (pathogenic fungi).
(iii) The miscellaneous components that may be present in the cell walls of true fungi
include lipids, melanins, sporopollenins, D-galactosamine polymers, and polyuronids.
 Lipids are often deposited on the inner surface of the wall and may help to
prevent desiccation.
 Melanins, dark brown to black pigments, are produced by the oxidation and
polymerization of phenolic compounds. They get incorporated into the existing
wall layers or get deposited as a surface coating on spores, vegetative hyphae or
fruiting bodies of many fungi.
 Sporopollenin, an aromatic polymer is a characteristics component of pollen
grains and is also present in some fungal spore walls. It is very resistant to
physical and chemical attack.
Value addition
 Many call wall proteins are modified by glycosylation, i.e. the attachment
of short chains of polysaccharides to the polypeptide.
 In Saccharomyces cerevisiae, the degree of glycosylation is quite high,
and the mannose is the sugar involved in the process, thus the proteins
are often called mannoproteins or mannans.
 Proteins, which are located close to the external wall surface, determine
the pore size of the cell wall as well as the surface properties (adhesion
and recognition).
 Zygomycota typically have a mixture of fully acetylated chitin, partially
acetylated and nonacetylated chitin to produce  -1, 4 glucosamine

polymers, called chitosan.

 These chitosan fibres are cross-linked by polymers of uronic acids such
as glucuronic acid and various neutral sugars. The cell wall matrix
comprises glucans and mannoproteins as it does in members of the
other fungal groups.
Value addition
 Fungal wall composition is not fixed. It can change substantially at
different stages of the life cycle, demonstrating that the wall structure
and composition reflect the functional needs of a fungus.
 Mucor rouxii (Zygomycetes), which is a dimorphic fungus, the yeast cells
have considerably more mannan, probably in the form of
mannoproteins, compared with the hyphae.
 As another example, many yeasts such as S. cerevisiae have a high
content of mannans but little chitin in their walls, the yeast forms of
Basidiomycetes also have a high mannan content but correspondingly
less glucan.
 The Major Components of Cell Walls of Selected Groups of Fungi.
Taxonomic group (with Structural components (fibrous) Matrix component

example) (gel-like)
Chytridiomycota Chitin Protein

(Allomyces)
Glucan Lipid
Zygomycota Chitin and Chitosan Polyglucuronic acid
(Mucor) Gluronomannoproteins
Lipids

Polyphosphate
Ascomycota Chitin  -1,3 and  -1,4 glucans

(Saccharomyces)
 -(1, 3),  -(1, 6) glucan Galactomannoproteins
Lipid
Basidiomycota
(Schizophyllum) Chitin Xylomannoproteins
 -1, 3,  -1, 6 glucan  -1, 3 glucan
Lipids
(Coprinus) Chitin Mannoproteins
 -1, 3 or b -1, 6 glucan  -1, 3 glucan
Lipids
Oomycota Cellulose Protein
(Phytophthora)  -1, 3,  -1, 6 glucan Lipid
 -1, 3 glucan
Architecture of Fungal Cell Wall

Wall architecture represents the bricks and mortar of the wall. The major wall components
can be revealed by chemical analysis. General pattern of wall architecture was studied in
Neurospora crassa and was found to be quite constant for other fungi also and consist of the
following layers:
a) An outermost layer of amorphous glucans (non-fibrillar or matrix), predominantly

with  -1, 3- and  -1, 6-linkages.

b) A reticulum of glycoprotein embedded in a protein matrix. This layer is not

commonly found in several other fungi studied except in Neurospora.
c) A more or less discrete layer of protein.
d) Chitin or cellulose micro fibrils embedded in protein forming the innermost layer.

Figure: A. Diagram to illustrate the wall architecture in a ‘mature’ (subapical) region of a

hypha of Neurospora crassa as evidenced by sequential enzymatic digestion. (a) Outermost
layer of amorphous ß-1,3-glucans and ß-1,6-glucans. (b) glycoprotein reticulum embedded
in protein. (c) a more or less discrete protein layer, (d) chitin microfibrils embedded in
protein, (e) plasma membrane. Based on Hunsley & Burnett, 1970
B. Diagrammatic representation of a typical fungal wall
Source:http://archive.bio.ed.ac.uk/jdeacon/FungalBiology/chap3a.htm#Fig3.9
http://upload.wikimedia.org/wikipedia/commons/f/fc/Cell_wall_structure_of_Fungi.gif
 Both inner microfibrillar component (cellulose, chitin) and outer amorphous (non-
fibrillar or matrix) component (glucan, proteins, mannan) are bonded to each other
and amongst themselves by covalent bonds thus strengthening the wall.
 The total wall thickness is about 125nm in the mature region of the hyphae.
 However, at the growing tip the wall is much thinner, about 50nm, and simpler too
consisting of only two wall layers - inner layer of chitin and outer layer of protein.
This observation suggests a much complex as well as stronger wall at maturity.
Functions of Fungal Wall
The wall is the structure that gives fungi most of their unique features and serve many
important roles:
1. It determines the shape of the fungi, because removal of the wall by enzymic
treatments leads to the release of spherical protoplasts. So, the way in which a
fungus grows, whether as hyphae or yeasts, is determined by the wall components.
2. It acts as a molecular sieve regulating the passage of large molecules through the
wall pore space. The wall's ability to safely limit turgor pressure appears to be a
primary reason for the survival and evolution of fungi.
3. It functions in the recognition of events associated not only with sexual reproduction,
but also with various interactions of fungi with potential plant and animal symbionts.
4. It acts as an interface between a fungus and its environment protecting it against

osmotic lysis.
5. Presence of pigments such as melanin in the wall can protect the cells against
ultraviolet radiation or the lytic enzymes of other organisms.

6. The wall can also have several physiological roles. It can act as binding site for
enzymes, such as invertase (which degrades sucrose to glucose and fructose) and b-
glucosidase (which degrades cellobiose to glucose in the final stages of cellulose
breakdown).
7. It can also have antigenic properties that mediate the interactions of fungi with other
organisms.
8. The mechanical strength provided by the hyphal wall enables fungi to assume a
variety of forms.
Value Addition
Extrahyphal Matrix
 A diffuse layer of polysaccharide or glycoprotein surrounding some yeasts
cells and hyphae. This layer is present in addition to the main structural
components of the wall.
 Production often influenced strongly by growth conditions. For example,
pullulan, (β-1,4-glucan) an extracellular sheath on Aureobasidium pullulans is
favoured by an abundant sugar supply in nitrogen-limiting growth conditions.
 This polymer is used in Japan to make a film-wrap for food.
 Besides pullulan, some other gel-like materials around fungi also have found
potential commercial roles. Scleroglucan from Sclerotium rolfsii
(Basidiomycota) binds to the surface of some tumour cells and has been
investigated for a possible therapeutic role.
 These extracellular matrix materials can also have important roles in the
interactions of fungi with other organisms.
 For example, Cryptococcus neoformans (yeast) is an opportunistic pathogen
of humans. The presence of a polysaccharide capsule masks the antigenic
components of the cell wall so that the fungus is not engulfed by white blood
cells and thus can proliferate in the tissues.
The Plasma Membrane

The plasma membrane of fungi is similar to that of other eukaryotes, consisting of a

phospholipid bilayer and associated proteins and sterols. It is intimately bound to the cell
wall and is also known as plasmalemma or cell membrane.
 However, the fungal membrane is unique in one respect: the main membrane
sterol is ergosterol, in contrast to cholesterol in animals and the
phytosterols, such as β-sitosterol in plants.
Figure: Ergosterol
Source: http://en.wikipedia.org/wiki/File:Ergosterol_structure.svg
 This difference is important in practice to control human mycoses and plant-

pathogenic fungi, because some of the most effective antifungal antibiotics
and fungicides act specifically on ergosterol or on its biosynthetic pathway.
Role of Plasma Membrane
The main role of the plasma membrane is:
1. It regulates the uptake and release of materials.

2. It anchors some enzymes, in fact, the two main wall synthetic enzymes, chitin
synthase and glucan synthetase, are integral membrane proteins; they are anchored
in the membrane in such a way that they produce polysaccharide chains from the
outer membrane face.
3. It relays signals from the external environment to the cell interior. This process is
known as signal transduction.
Nuclei and Associated Structures

 The hyphae of fungi almost invariably contain large numbers of nuclei. In aseptate
fungi, nuclei generally appear to be distributed randomly throughout the cytoplasm
of an actively growing hypha.
 In septate forms, depending upon the species involved and the phase of the life cycle
examined, individual hyphal compartments may contain one, two, or many nuclei.
 Some species possess special mechanisms that ensure that only two genetically
compatible nuclei are present in each compartment of a hypha.
 Most fungi are haploid with chromosome numbers ranging from about 6 to 20, but
the Oomycota are diploid, and some other fungi can alternate between haploid and
diploid generations. For example, Saccharomyces cerevisiae can grow as either
haploid or diploid yeast.
 Interestingly, the peculiar organization of fungal hyphae, with several nuclei in a
common cytoplasm, enables haploid fungi to exploit the advantages of both haploidy
and diploidy.
 Haploidy in fungi is one of the most remarkable features. But, it seems that mycelial
fungi have remained haploid because they can exploit the advantages of a diploid
lifestyle in any case.
 Diploidy predominates in the rest of the eukaryotic world (plants, animals,
Oomycota, and protists) and even in yeasts such as Candida and Saccharomyces.
 The nuclei of most fungi are usually small (1-2 µm diameter), but exceptionally large
nuclei up to 20-25µm diameter can be observed in some genera (e.g. Basidiobolus
ranarum, Chytridiomycota).
 The shape of nuclei is generally spherical to ovoid.
 They are extremely plastic and capable of squeezing through tiny septal pores as
well as through narrow structures at the tips of which various types of spores are
produced.
 They also have a tendency to become thin and elongated or teardrop shaped while
moving into germ tubes arising from germinating spores.
 As in other eukaryotes, they are surrounded by a double nuclear membrane with
pores, and they contain a ribonucleic acid (RNA)-rich nucleolus.
 They are difficult to be observed by light microscopy because of their small size and
because of the same optical properties as cytoplasm
 Fungal nuclei can be observed under light microscope with stains such as Giemsa,
iron-hematoxylin, acetoorcein, acetocarmine and with fluorescent stains such as 4',
6' -diamidino-2-phenylindole (DAPI) and mitramycin.

Nuclear Division
 Nuclear divisions in the fungi are basically intranuclear, i.e. the nuclear membrane
and the nucleolus usually remain intact during most stages of mitosis.
 Nuclear membrane breaks in the interzonal region and then re-forms around the
daughter nuclei.
 There is no clear metaphase plate; the chromosomes appear to be dispersed
randomly.
 There has been considerable interest in ultrastructural details of mitosis and meiosis
in fungi in recent years because of their value as phylogenetic characters.
Spindle-Pole Bodies (SPB)
 Generally, typical centrioles are not present in fungi. But they possess nucleus-
associated bodies, known as spindle-pole bodies (SPB).
 Species of fungi that produce flagellate cells lack SPBs, possessing instead a pair of
centrioles that are associated with the nuclear envelope.
 Chemical composition of SPB is not clear, but evidence indicates that these
structures functions like a centriole, as a centre for microtubule assembly during
mitotic and meiotic divisions; therefore they are also known as microtubule
organizing centres.
 The morphology of SPB varies depending upon the type of fungus and the exact
phase of nuclear division studied.
 It may be a small, electron-dense disc-shaped structure lying just outside the
nuclear envelope in Ascomycota and anamorphic fungi or is often composed of two
globular ends connected by a bridge in Basidiomycota.
 Duplication of SPB takes place during prophase. This duplicated SPB then separates
into two identical halves that eventually become positioned at opposite poles of the
dividing nucleus.
 Depending upon the species involved, SPBs may remain outside the nuclear envelope
during nuclear division or become inserted in the nuclear envelope.
Nucleolus

The typical fungal nucleus usually contains a centrally positioned, prominent nucleolus.
Depending upon the species involved, the nucleolus can have one of three fates during
division:
i. It may persist and eventually undergo fission.

ii. It may become dispersed and no longer be visible in a dividing
nucleus.
iii. It may be discarded from the dividing nucleus into cytoplasm as
an intact entity.
Value Addition
How to observe fungal chromosomes?
 Fungal chromosomes are usually quite small and difficult to observe in squashed
and stained preparations. As a result, direct chromosome counts are difficult to
make.
 A new approach to karyotype analysis called pulsed-field gel electrophoresis
(PFGE) has proved to be extremely valuable for determining chromosome
numbers in fungi.
 In this technique, chromosomes in an agarose gel are exposed to a pulsed
electrical field that causes them to move at different velocities depending on their
sizes and shapes.
 Then it is stained with ethidium bromide and exposed to ultraviolet light. The
chromosomes can be observed as distinct countable bands.
Fungal organelles
 Beside nuclei, the most noticeable fungal organelle is the mitochondrion. There are
numerous mitochondria in hyphae. Under the electron microscope, they commonly
appear as electron-dense, branched or lobed structures and less commonly as
thread-like, extremely long structures. The mitochondrial cristae of the true fungi are
flattened and plate-like in contrast to the tubular cristae found in many organisms
including Oomycota.

 Fungal cytoskeleton is composed of ribosomes, endoplasmic reticulum, Golgi

bodies, microbodies, filasomes, multivesicular bodies, microtubules,
microfilaments vacuoles, lipid bodies and glycogen storage particles.
 Fungi have a secretory system, consisting of the endoplasmic reticulum (ER), Golgi
bodies and membrane-bound vesicles. Typically the Golgi bodies or dictyosomes
in a true fungus are very simple unlike other organisms including Oomycota. It
consists of a single cisternal element instead of multiple stacked cisternae. Moreover,
there is no differentiation into proximal and distal faces. Because of their simplicity,
these structures are sometimes referred to as Golgi equivalents. However, their
functionality remains the same as that of complex Golgi bodies, i.e. packaging and
transfer of materials in vesicles. These vesicles may collect at the hyphal tip or may
coalesce to form larger vesicles, are known as apical vesicle cluster (AVC).
 Another true organelles known as Woronin bodies (named after Michael S Woronin,
a Russian scientist) are also present in certain types of fungi and are typically
associated with septal pores. They are spherical structures, having a diameter of 0.1-
0.75 µm and are bound by a single bilayer membrane. They are only present in
filamentous, septate fungi belonging to Ascomycota and Anamorphic fungi.
Figure: TEM of L.S. septa showing woronin bodies (arrow)

Source: http://www.mycologia.org/content/94/2/260/F1.expansion
 Lomasomes are the membranous structures that have been observed between the
plasma membrane and the hyphal wall in a variety of different types of fungi. They
are common in samples prepared for study with traditional chemical fixation
procedures.
 Vacuoles are often seen as conspicuous, rounded structures in the older regions of
hyphae, but recent work has shown that there is also a tubular vacuolar system

extending into the tip cells. It is an extremely dynamic system with several
important functions, rather than being an inert depository for storage of water,
nutrients or waste materials.
http://www.fungionline.org.uk/images/3hyphae/hypha.gif
The important functions performed by vacuolar system in fungi:
(i) Storage of compounds, e.g. the vacuoles of several fungi, including mycorrhizal
species, accumulates phosphates in the form of polyphosphate. Vacuoles also seem
to be major sites for storage of calcium, which can be released into the cytoplasm as
part of the intracellular signaling system. They also store volutin in the yeasts.
(ii) Vacuoles contain a range of lytic enzymes such as nucleases, phosphatases,

trehalases and proteases, which may serve in the recycling of cellular metabolites
during growth and development, and in the autolysis of cell components in senescent
cells.
(iii) Vacuoles provide turgor needed for cell growth and maintenance of the cell
shape.
(iv) They also have a role in regulation of cellular pH.

 Although the production of crystals by fungi is known since the time of de

Bary (1887), little is known about their development and exact function. It
has been suggested that crystals may be involved in calcium regulation and
may also function to strengthen the walls of certain structures. It is possible
that they also may protect hyphae from fungus-grazing arthropods. Hyphae
of some fungi excrete oxalic acid, which unites with calcium ions in the
environment. Crystals of calcium oxalate form on the hyphae and host tissues
of the plant pathogenic fungus Sclerotium rolfsii.
Organization of the Hyphal Apex
 The hyphal apex is of special interest as the site of growth, and for many
years it has been recognized as the key to understand the behaviour of fungi.
 Under transmission electron microscope the apex of the growing hypha is
observed to be packed with two types of membrane-bound vesicles –
macrovesicles (diameter >100nm) and microvesicles (diameter <100nm).
These clusters of vesicles at the tip are called as the apical vesicle cluster
(AVC), which disappear and reappear with the cessation and start of the
growth, respectively.
 In majority of the true fungi there is a third structure visible with these
vesicles called the Spitzenkorper. This structure also disappears when
growth stops and reappears when growth restarts and it changes position in
the apex, when a hypha changes direction of growth. Exclusion of other major
organelles occurs with it in the growing tip, and so it was regarded as a
possible 'apical growth organelle'.
 Macrovesicles present in the hyphal tips may act secretory vesicles that
contain enzymes and preformed polymers used for the production of
amorphous matrix of fungal cell wall.
 Some of the apical vesicles, especially on the shoulders of the apex and lining
the lateral walls have a distinctly filamentous appearance, so they have been
termed filasomes.
 Most of the vesicles in hyphal tips have not been characterized chemically,
but some of the microvesicles have been termed chitosomes as they are
involved in the movement of the enzyme chitin synthase through the
cytoplasm to the plasma membrane at the hyphal apex.

 These are small spheroidal bodies, 40-70nm in diameter. A ‘shell’ about 7nm
thick surrounds each. They are self-assembling aggregates of the enzyme
chitin synthase, each particle containing sufficient enzyme molecules to
produce a chitin microfibril, in which the individual chitin chains coil round one
another.
Factors Affecting Fungal Growth
The growth of fungi is affected by nutrients, light, water, aeration, temperature and
Hydrogen ion (H+) concentration.
Light
 Many fungi grow well in dark, but some prefer daylight or alternate light and
darkness to produce spores.
 Visible light (380-720nm) has little effects on somatic growth but it can affect
formation of fruit bodies of many Basidiomycota or other differentiation
events.
 Sporangiophores of Zygomycota and ascal tips of some Ascomycota show
phototropism.
Water
 All fungi require moisture for growth but the amount required varies widely.
Water is essential for diffusion of nutrients into cells. It helps release
extracellular enzymes. It also maintains the turgidity of cytoplasm.
 Those capable of growing at very low water activity are referred to as
xerophilic, e.g. Aspergillus, Penicillium etc.
 Those that are capable of growing at very high water activity are known as
hydrophilic, e.g. Chaetomium, Stachybotrys etc.
Aeration
Most fungi are strictly aerobic. They require oxygen throughout their life cycle in
some stage or the other.
Temperature
 Majority of fungi are mesophilic, i.e., they grow at temperature ranging from
10-35oC. Optimum range for growth may be between 15 and 30oC.

 However, some fungi are heat-loving and grow only at temperature above 45oC
or higher and fail to grow below 20oC. Such fungi are known as thermophilic
such as, Chaetomium thermophilum, Rhizomucor meihei etc.
 Some fungi are psychrophilic i.e., they are able to grow below 10oC but are
unable to grow above 20oC. Quite a few fungi are psychrotolerant and are
able to show growth both at freezing point and at room temperature.
Examples: Mucor, Rhizopus, Penicillium, Cladosporium etc.
Hydrogen ion (H+) concentration
 Fungi differ in their pH requirements. Majority will grow well over a pH range of
3 -7.
 Some fungi can tolerate pH of 2 and below. Such fungi are known as
acidophilic, e.g. Penicillium funiculosum, Aspergillus niger etc.
 In contrast, alkalophilic fungi pH optima between 9-10, e.g. Aspergillus oryzae.
Summary
Majority of the fungi grow in warm, moist environments, however, there are many species
which occur in habitats that are extremely cold, hot and dry, or otherwise seemingly
inhospitable. Although, the optimum conditions for growth and reproduction vary widely
with the species, still we find maximum diversity of fungi in tropical area.
Similar to animals, fungi are heterotrophs, i.e. dependent upon already produced
organic compounds and energy sources for survival. But unlike animals, which ingest and
then digest their food, fungi first digest food by the production of enzymes outside their
body (exoenzymes) and then ingest it by absorption. Exoenzymes break down large and
relatively insoluble molecules such as carbohydrates, proteins and lipids into smaller and
more soluble molecules, which can easily be absorbed through the wall and cell membrane
of fungi. Presence of free water, however, is essential for the diffusion of exoenzymes and
nutrients across the fungal cell wall and membrane. This might be the reason that actively
growing fungi are mainly restricted to humid places.Based upon how they obtain nutrients,
they have been broadly divided into categories of saprobes, parasites,
symbionts/mutualists, and endophytes/ectophytes. Saprobes are also known as saprophytes
or saprotrophs are fungi growing and obtaining their nutrition from dead organic materials.
They can attack almost any existing substance wherever adequate moisture and
temperature are present. Rhizopus, Aspergillus (black mould) and Penicillium (green mould)
are, a few very commonly encountered species on all type of substrata. Saprobes along with

some bacteria help in recycling of organic matter in natural and agricultural environment.
Parasite obtains some or all of its nutrients from the living tissues of another organism while
being in close relationship with it. A parasite that causes disease is termed a pathogen.
Parasitic fungi are also able to grow on dead and decaying organic matter. The latter can be
confirmed by their ability to grow on artificial synthetic media. Such fungi are known as
facultative saprobes/saprotrophs. Similarly, there are saprobic organisms, which are
able to infect another living organism under some conditions. These are known as
facultative parasites. Those fungi that are unable to grow on artificial media even after
the best of efforts and remain parasitic in nature are known as ecological obligate
parasites or biotrophs. Based on the way they obtain their nutrients parasites are of two
types: Necrotrophic and Biotrophic. Both necrotrophs and biotrophs may be parasitic on
plants, animals or on other fungi. Necrotrophic parasites are also known as necrotrophs
or perthotrophs. This type kills the living host cells in advance of its entry, by producing
toxins or degradative enzymes and then absorbing released nutrients and growing between
and into dead and dying cells. Biotrophs are also known as biotrophic parasites. These
parasites obtain some or all of its nutrients from the living cells or tissues of host, usually by
means of special nutrient absorbing structures - haustoria. A few fungi parasitize humans
and most of those infecting humans are opportunistic parasites. These grow more
commonly as saprobes in soil, composts, bird excreta, etc., but can infect through wounds
or through lungs. Most of the spores of such parasites are air borne, e.g. Aspergillus
fumigatus, Histoplasma capsulatum, and Blastomyces dermatitidis. Generally, all these
pathogens pose a threat to debilitated individuals suffering from life threatening diseases or
to transplant patients. For example, fungus Pneumocystis carinii is the most common cause
of death of acquired immune deficiency syndrome (AIDS) patients in several parts of the
world. Symbionts or mutualists are kind of association, where the fungi absorb nutrients
from other organisms but are not detrimental to them. Such fungi are known as
mutualistic fungi. The association can be: lichens and mycorrhizae. Lichens are
associations of fungi and algae and the mycorrhizae are associations of fungi with roots of
higher plants. The fungal symbiont in a mycorrhiza gets carbohydrates from plant host, and
the plant gets minerals from the fungus. Endo or ectophytes are fungi present in or on the
leaves & stem of healthy plants and animals. Some such endophytic fungi may protect both
conifers and grasses against insect herbivory, influence flowering and many of these fungi
produce secondary metabolites, which may be antibacterial or antifungal. Infection
structures are specialized structures formed in parasitic fungi at the tips of germ tubes or
hyphae on the outside of the host. The simplest pre-penetration structures are terminal

swellings on germ tubes; called appressoria. Infection structures are formed when the
spore (s) germinates; producing a germ tube (GT), which align itself locate a stomatal ridge
and stops growing. Then the apex swells to form an appressorium (A). Appresorium may
secrete enzymes like cutinase helping penetration peg enter the host through epidermal cell
wall directly or through stomatal opening.
Fungal germ tubes and hyphae grow almost exclusively at their tips, i.e. they show
apical growth. The original hypha and the first formed branches repeatedly branch behind
their tips. These branches separate from each other until finally the colony develops a
typical circular outline. In the older parts of the colony where nutrients have been depleted,
mycelia grow towards one another and fuse by tip-to-tip contact producing narrow hyphal
branches. This leads to localized breakdown of their walls and continuity of the cell
cytoplasm. This process is known as hyphal anastomosis leading to the development of
typical interconnected network of hyphae observable in higher fungi. In lower fungi the
vegetative hyphae do not anastomose. Diffused (intercalary) type of growth common in
plant tissues is rare in fungi and seems to be limited to hyphae involved in lifting
reproductive structures into the air for spore dispersal.
There are three major growth forms of fungi. The thallus or the body of the fungus
typically consists of microscopic, tubular, threadlike hyphae that branch in all directions,
spreading over or within whatever substrate the fungus uses for food. The collection of
hyphae is known as mycelium and such thalli are commonly known as 'moulds'. Some of the
more primitive fungi, such as the Chytridiomycota, often have single rounded cells or
dichotomously branched chains of cells, attached to a food source by tapering rhizoids.
Many other fungi grow as unicellular yeasts and produce daughter cells either by binary
fission, e.g. Schizosaccharomyces pombe, or budding, e.g. Saccharomyces cerevisiae.
Intermediate stages between yeast cells and true hyphae also occur and are known as
pseudohyphae. The term yeast is a morphological term and does not refer to a particular
taxonomic group.
Certain fungi can exist as either mycelial (M) or yeast-like (Y) phase and are said to be
dimorphic. This ability to switch between these two phases is termed dimorphism.
Important pathogens of humans, higher animals and some plants exist as dimorphic fungi.
For example, Candida albicans grows in yeast form on the moist mucosal membranes of
humans, but changes to hyphae for invasion of host tissues.

A thin, usually transparent, tubular wall lines a fungal hypha. The interior of the hypha is
filled or lined with a layer of protoplasm. Hyphae of majority of the species are interrupted
at some places by partitions, or cross-walls, called septa. Such hyphae are known as
septate and are present in Ascomycota, Basidiomycota and Anamorphic fungi. The septate
fungi commonly have several nuclei in the apical compartment, but often only one or two in
each compartment behind the apex. In some species septa are produced at more or less
regular intervals along the length of a hypha, dividing it into individual compartments or
cells. Individual compartment may contain one nucleus (monokaryon), or two nuclei
(dikaryon), or many nuclei (multinucleate). In other species, the vigorously growing
portions of hyphae lack regularly spaced septa. Such mycelia are known as
aseptate/nonseptate/coenocytic and are observed in Zygomycota and Oomycota. In
such mycelia, septa are present only at the bases of reproductive structures and in mature,
highly vacuolate portions of hyphae.
Basically, there are two types of septa - simple and complex - both types appear to
form by the centripetal growth from the hyphal wall inward. Simple septum can be of two
types, complete and incomplete. Complete septa are present at the bases of asexual and
sexual reproductive structures and in older, highly vacuolated portions of aseptate hyphae
and incomplete septum possesses a single large central pore, which allows the passage of
cytoplasmic organelles and even nuclei, showing that the protoplasts of adjacent hyphal
compartments are continuous. Typically, the Ascomycota and Anamorphic fungi have a
septum of this type. Complex septum is kind of septum characteristics of Basidiomycota.
It has a narrow central channel and is bounded by two flanges of amorphous wall material.
On either side of this septum there are bracket-shaped membranous structures termed
septal pore cap or parenthosome, which have pores that allow cytoplasmic continuity but
prevent the movement of major organelles. This type of septum is known as dolipore
septum. The roles of septa have been debated for many years and are still not completely
understood. They might help to provide structural support to hyphae, especially in relatively
dry conditions. Septa also can help to defend against damage, because Woronin bodies or
other materials plug the septal pores if hyphae are physically disrupted or are damaged by
other organisms. The main significance of septa is thought to be in differentiation. If a
septum is plugged it enables a fungus to transform from a continuous series of
compartments to a number of independent cells or regions that can undergo separate
development. The aseptate fungi also produce complete cross-walls to isolate old parts of
the hyphae or to produce asexual or sexual structures.

Chemical analysis of fungal walls can vary considerably between and within different
groups of fungi, however, the basic components remains the same. The major wall
components can be categorized basically into two types. The structural (fibrillar)
polymers, consisting predominantly of straight-chain molecules, are cross-linked, providing
structural rigidity. It forms the inner layer of the wall. The structural components typically
have chitin, chitosan and glucans as their major wall polysaccharides. Chitin is the most
important component of cell wall of Chytridiomycota, Ascomycota and Basidiomycota. It is
also a major component of the exoskeleton of insects and other arthropods. Chitin along
with chitosan (a major component) is present in the hyphal walls of Zygomycota. However,
in Oomycota, cellulose instead of chitin is the main constituent of the primary wall. Small
amount of chitin has been reported in some genera of Oomycota. For details, readers are
referred to see chapter on Oomycota. The second component is matrix components,
consisting of amorphous or gel-like or crystalline material found predominantly in the outer
layer, covering and embedding the structural (fibrillar) polymers. The amorphous
component typically includes proteins, mannans and glucans. Some miscellaneous
components that may be present in the cell walls of true fungi include lipids, melanins,
sporopollenins, D-galactosamine polymers, and polyuronids.
The wall is the structure that gives fungi most of their unique features and serve many
important roles, it determines the shape of the fungi, because removal of the wall by
enzymic treatments leads to the release of spherical protoplasts. It acts as a molecular
sieve regulating the passage of large molecules through the wall pore space. The wall's
ability to safely limit turgor pressure appears to be a primary reason for the survival and
evolution of fungi. It functions in the recognition of events associated not only with sexual
reproduction, but also with various interactions of fungi with potential plant and animal
symbionts. It acts as an interface between a fungus and its environment protecting it
against osmotic lysis. Presence of pigments such as melanin in the wall can protect the cells
against ultraviolet radiation or the lytic enzymes of other organisms. The wall can also have
several physiological roles. It can act as binding site for enzymes, such as invertase (which
degrades sucrose to glucose and fructose) and b-glucosidase (which degrades cellobiose to
glucose in the final stages of cellulose breakdown). It can also have antigenic properties
that mediate the interactions of fungi with other organisms. The mechanical strength
provided by the hyphal wall enables fungi to assume a variety of forms.
The plasma membrane of fungi is similar to that of other eukaryotes, consisting of a

phospholipid bilayer and associated proteins and sterols. It is intimately bound to the cell

wall and is also known as plasmalemma or cell membrane. However, the fungal membrane
is unique in one respect: the main membrane sterol is ergosterol, in contrast to cholesterol
in animals and the phytosterols, such as β-sitosterol in plants.
The hyphae of fungi almost invariably contain large numbers of nuclei. In aseptate fungi,
nuclei generally appear to be distributed randomly throughout the cytoplasm of an actively
growing hypha. In septate forms, depending upon the species involved and the phase of the
life cycle examined, individual hyphal compartments may contain one, two, or many nuclei.
Some species possess special mechanisms that ensure that only two genetically compatible
nuclei are present in each compartment of a hypha. Most fungi are haploid but the
Oomycota are diploid, and some other fungi can alternate between haploid and diploid
generations. Interestingly, the peculiar organization of fungal hyphae, with several nuclei in
a common cytoplasm, enables haploid fungi to exploit the advantages of both haploidy and
diploidy. Haploidy in fungi is one of the most remarkable features. But, it seems that
mycelial fungi have remained haploid because they can exploit the advantages of a diploid
lifestyle in any case. The nuclei of most fungi are usually small, their shape is generally
spherical to ovoid and they are extremely plastic and capable of squeezing through tiny
septal pores. They are surrounded by a double nuclear membrane with pores, and they
contain a ribonucleic acid (RNA)-rich nucleolus. They are difficult to be observed by light
microscopy because of their small size and because of the same optical properties as
cytoplasm. Fungal nuclei can be observed under light microscope with special stains.
Nuclear divisions in the fungi are basically intranuclear, i.e. the nuclear membrane and the
nucleolus usually remain intact during most stages of mitosis. There is no clear metaphase
plate; the chromosomes appear to be dispersed randomly.
Generally, typical centrioles are not present in fungi. But they possess nucleus-
associated bodies, known as spindle-pole bodies (SPB). Species of fungi that produce
flagellate cells lack SPBs, possessing instead a pair of centrioles that are associated with the
nuclear envelope. SPBs may be a small, electron-dense disc-shaped structure lying just
outside the nuclear envelope in Ascomycota and Anamorphic fungi or is often composed of
two globular ends connected by a bridge in Basidiomycota. Duplication of SPB takes place
during prophase. Depending upon the species involved, SPBs may remain outside the
nuclear envelope during nuclear division or become inserted in the nuclear envelope.
The typical fungal nucleus usually contains a centrally positioned, prominent nucleolus.
Beside nuclei, the most noticeable fungal organelle is the mitochondrion. There are
numerous mitochondria in hyphae. Under the electron microscope, they commonly appear

as electron-dense, branched or lobed structures and less commonly as thread-like,

extremely long structures. The mitochondrial cristae of the true fungi are flattened and
plate-like in contrast to the tubular cristae found in many organisms including Oomycota.
Fungal cytoskeleton is composed of ribosomes, endoplasmic reticulum, Golgi bodies,
microbodies, filasomes, multivesicular bodies, microtubules, microfilaments vacuoles, lipid
bodies and glycogen storage particles. Fungi have a secretory system, consisting of the
endoplasmic reticulum (ER), Golgi bodies and membrane-bound vesicles. Typically the Golgi
bodies or dictyosomes in a true fungus are very simple unlike other organisms including
Oomycota. It consists of a single cisternal element instead of multiple stacked cisternae.
Moreover, there is no differentiation into proximal and distal faces. However, their
functionality remains the same as that of complex Golgi bodies, i.e. packaging and transfer
of materials in vesicles. These vesicles may collect at the hyphal tip or may coalesce to form
larger vesicles, are known as apical vesicle cluster (AVC). Another true organelles known as
Woronin bodies are also present in certain types of fungi and are typically associated with
septal pores. They are spherical structures, having a diameter of 0.1-0.75 µm and are
bound by a single bilayer membrane. They are only present in filamentous, septate fungi
belonging to Ascomycota and Anamorphic fungi. Lomasomes are the membranous
structures that have been observed between the plasma membrane and the hyphal wall in a
variety of different types of fungi. They are common in samples prepared for study with
traditional chemical fixation procedures. Vacuoles are often seen as conspicuous, rounded
structures in the older regions of hyphae, but recent work has shown that there is also a
tubular vacuolar system extending into the tip cells. It is an extremely dynamic system with
several important functions.
The hyphal apex is of special interest as the site of growth, and for many years it has
been recognized as the key to understand the behaviour of fungi. Under transmission
electron microscope the apex of the growing hypha is observed to be packed with two types
of membrane-bound vesicles – macrovesicles and microvesicles . These clusters of vesicles
at the tip are called as the apical vesicle cluster (AVC), which disappear and reappear
with the cessation and start of the growth, respectively. In majority of the true fungi there
is a third structure visible with these vesicles called the Spitzenkorper. This structure also
disappears when growth stops and reappears when growth restarts and it changes position
in the apex, when a hypha changes direction of growth. Exclusion of other major organelles
occurs with it in the growing tip, and so it was regarded as a possible 'apical growth
organelle'. Most of the vesicles in hyphal tips have not been characterized chemically, but
some of the microvesicles have been termed chitosomes as they are involved in the

movement of the enzyme chitin synthase through the cytoplasm to the plasma membrane
at the hyphal apex. These are small spheroidal bodies, 40-70nm in diameter. A ‘shell’ about
7nm thick surrounds each. They are self-assembling aggregates of the enzyme chitin
synthase, each particle containing sufficient enzyme molecules to produce a chitin
microfibril, in which the individual chitin chains coil round one another.
The growth of fungi is affected by nutrients, light, water, aeration, temperature and
Hydrogen ion (H+) concentration. Many fungi grow well in dark, but some prefer daylight or
alternate light and darkness to produce spores. Visible light (380-720nm) has little effects
on somatic growth but it can affect formation of fruit bodies of many Basidiomycota or other
differentiation events. All fungi require moisture for growth but the amount required varies
widely. Water is essential for diffusion of nutrients into cells. It helps release extracellular
enzymes. It also maintains the turgidity of cytoplasm. Those capable of growing at very low
water activity are referred to as xerophilic, e.g. Aspergillus, Penicillium etc. Most fungi are
strictly aerobic. They require oxygen throughout their life cycle in some stage or the other.
Majority of fungi are mesophilic, i.e., they grow at temperature ranging from 10-35oC.
Optimum range for growth may be between 15 and 30 oC. However, some fungi grow only
at temperature above 45oC or higher. Such fungi are known as thermophilic such as,
Chaetomium thermophilum, Rhizomucor meihei etc. Some fungi are psychrophilic i.e.,
they are able to grow below 10oC but are unable to grow above 20oC. Quite a few fungi are
psychrotolerant and are able to show growth both at freezing point and at room
temperature. Examples: Mucor, Rhizopus, Penicillium, Cladosporium etc. Fungi differ in their
pH requirements. Majority will grow well over a pH range of 3 -7. Some fungi can tolerate
pH of 2 and below. Such fungi are known as acidophilic, e.g. Penicillium funiculosum,
Aspergillus niger etc.
Glossary
Anastomosis: a process where the hyphal branches are capable of undergoing fusions with
one another.
Halophile: a salt loving organism that requires moderate to large concentrations of salt.
Biotroph: used in reference to a plant pathogenic fungus that requires living host cells.
Hemibiotroph: a plant pathogenic fungus that initially requires living host cells but that
eventually kills host cells and grows on the remains of dead and dying cells.
Necrotroph: a pathogenic fungus that kills host cells in advance of its hyphae and then
lives on the dying and dead cells.

Facultative parasite: a saprobe capable of infecting another living another organism under
some conditions.
Facultative saprotroph: a parasitic organism capable of growing on dead organic matter
under some conditions.
Mycoparasite: a fungus that parasitizes another fungus
Endophyte: fungi that live inside the leaves and or stems of apparently healthy plant.
Appressorium: a flattened hyphal organ from which a minute hyphal peg grows and enters
the host.
Teleomorph: the sexual stage in the life cycle of a fungus.
Anamorph: the asexual stage in the life cycle of a fungus.
Holomorph: a fungus in all its possible forms either latent or expressed.
Dimorphism: a process of the production of two different types of zoospores or a fungus
able to occur in yeast form or in mycelia form.
Dolipore septum: a septum with a central pore surrounded by a barrel-shaped swelling of
the septal wall and covered on both sides by a perforated membrane termed either the
septal pore cap or the parenthosomes.
Haustorium: an absorbing organ originating on a hypha of a parasite that penetrates the
host cell wall and invaginates the host cell plasma membrane; most often formed by
obligate parasites.
Germ tube: the hyphal structure that first emerges from a germinating spore in most fungi.
Mycangium: a specialized fungus, containing pouch found in some beetles.
Septum: a cross wall in a hypha that develops centripetally.
Exercises
1.1 Fill in the blanks:
1. ………..… which is present in Oomycota is generally absent in most of the true

fungi.
2. Presence of ……………. is essential for the diffusion of exoenzymes and nutrients
across the fungal cell wall and membrane.
3. Terminal swellings of germ tube are called……………..
4. Fungal membrane differs from those of plants in having……………instead of beta-
sitosterol.

5. Dark phase visible under light microscope at the tips of fungal hyphae is termed
as…………………………
1.2 Match the following:
1 Aternaria alternata A Dimorphic fungi

2 Ambrosia beetles B Pullulan
3 Candida albicans C Ergosterol
4 Septal pores D Woronin bodies
5 Film wrap for food E Common ectophyte
1.3 Differentiate between:
1. Necrotrophs and Biotrophs

2. Septate and Aseptate mycelium
3. Facultative parasite and Facultative saprophyte
4. Endophyte and Ectophyte
5. Simple and Complex septum
6. Thermophilic and Psychrophilic
1.4 Define the following:

1. Ecological obligate parasite
2. Anastomosis
3. SPB
4. Dimorphic fungi
5. Lomasomes
6. Haustorium
7. Mycorrhizae
8. AVC
9. Mycoparasite
10. Chitosomes
Answers:
1.1 Fill in the blanks:

(1) Cellulose (2) Water (3) Appressoria (4) Ergosterol (5) Spitzenkorper
1.2 Match the following:
1-E, 2-C, 3-A, 4-D, 5-B
References
1. Alexopoulos, C.J., Mims, C.W., Blackwell, M. (1996). Introductory Mycology, John Wiley &
th
Sons (Asia) Singapore. 4 edition.
2. Webster, J. and Weber, R. (2007). Introduction to Fungi, Cambridge University Press,

rd
Cambridge. 3 edition.
3. Sethi, I.K. and Walia, S.K. (2011). Text book of Fungi and Their Allies, Macmillan
Publishers India Ltd.

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General Characteristics of True and Allied Fungi –Part I

PAPER-3: Mycology and Phytopathology

Lesson: General Characteristics of True and Allied Fungi –

Lesson Author: Dr. Inderjeet Sethi and Dr. Charu Kalra

College/Dept: Sri Guru Tegh Bahadur Khalsa College,

Deen Dayal Upadhyaya College, Department of Botany

Institute of Lifelong Learning, University of Delhi 1

Chapter: General Characteristics of True and Allied Fungi – Part I

 Chemical Composition of Cell wall

 The Plasma membrane

 Nuclei and associated Structures

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 Spindle Pole Bodies

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Nuclear division Mitosis is intranuclear, i.e. it takes place without the

Reserve food Glycogen, lipids or trehalose; mycolaminarin in Stramenopila

Institute of Lifelong Learning, University of Delhi 4

I. Saprobes (Gr. sapros = rotten; bios = life)

 Also known as saprophytes or saprotrophs.

Institute of Lifelong Learning, University of Delhi 5

II. Parasites (Gr. parasitos = table mate)

Why fungi are important as plant parasites in comparison to bacteria?

This is because the hyphal apex has a remarkable penetrating power,

Institute of Lifelong Learning, University of Delhi 6

1. Necrotrophic parasites (Gr. nekros = death; trophein = to nourish), are also

DID YOU KNOW?

o Majority of Pythium spp. (Oomycota) are plant pathogens. Only about

Institute of Lifelong Learning, University of Delhi 7

Rhizoctonia solani (Basidiomycota) can be controlled by Verticillium biguttatum

3. Fungal parasites of Humans

III. Symbionts or Mutualists

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IV. Endo or Ectophytes

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Figure: Ant-fungus relationship

Figure: Pupae of Ambrosia beetles, Xylosandrus crassiusculus with black fungi

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Figure: Haustorial complex, a specialized feeding organ of biotrophic fungal parasites of

Factors Influencing the Development of Infection Structures

Development of the appressorium have been found to be influenced by physical and

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Figure: Anastomoses in fungi

Figure: Videotaped sequence of anastomosis (successful hyphal fusion) of two

McCabe, M.P.Gallagher & J.W.Deacon, The University of Edinburgh)

In lower fungi the vegetative hyphae do not anastomose.

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Growth Forms of Fungi

There are three major growth forms of fungi:

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Mycelium - the body of the fungus

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(i) Simple septum can be of two types:

b) Incomplete septum: This kind of septum possesses a single large central