EGERTON UNIVERSITY

BOTA 233: MYCOLOGY

LECTURE NOTES

TOPIC ONE: INTRODUCTION TO MYCOLOGY AND ITS HISTORICAL

DEVELOPMENT

Introduction
Welcome to topic one. This topic is aimed at introducing you to the meaning of Mycology as a
subject and also give you the historical development on the study of this subject. You will also
learn why this subject is important for the advancement of humanity.

Learning Outcomes

By the end of this topic you should be able to:

i. Explain the meaning of Mycology

ii. Identify the major contributors to the development of this subject and explain their
contributions.
iii. State the importance of Mycology as a subject of Scientific study

Topic Content

1.1 Meaning and Importance of Mycology

Mycology is the study of fungi. The word mycology etymologically means the study of
mushrooms (Gr. Mykes = mushroom, logos = discourse/study). For a long time, man has had a lot
of interest in fungi, taking them as food substance and also acknowledging them as poisonous
substances. Edible forms of fungi were termed mushrooms while the poisonous forms were termed
the toadstools (derived from the German term todestuhl = death chair). During these early times,
mushrooms were taken as special food mainly for the wealthy and the rulers. Accidental poisoning
by mushrooms was also a common occurrence. The Roman emperor Claudius Caesar (AD 54)
was murdered by his wife who mixed his food with the fungus Amanita phalloides. This
necessitated a clear identification of the edible and the poisonous varieties of mushrooms.

Fungi are highly diverse and versatile organisms with ubiquitous distribution in the biosphere,
being found in both aquatic and moist terrestrial habitats as well as in air as spores. They are found
in moist soil and decomposing organic matter as saprophytes. They are also found growing in
some plants and animals as parasites where they mostly cause diseases, or as symbionts where
they are of ecological importance.

Man has always had lots of interactions with fungi in many spheres, as children through playing
with puffballs, mushrooms or toadstools that commonly grow during the rainy seasons, while
housewives always experience their effects through food and fruits spoilage in the kitchen, as well
as in the use of yeast in baking of bread or making “mandazis”, preparation of fermented food such
as vegetables in which fungal organisms play a role. Farmers on the other hand experience fungal
effects through the many plant diseases of fungal origin such as rusts, smuts, blights and mildews,
stem and root rots that affect fruits, vegetables, the crops in farms as well as the spoilage of stored
cereals. Other people may have experienced fungal organisms through the diseases that are
associated with them such as histoplasmosis, coccidiomycosis, aspergillosis, ringworms, athlete’s
foot or fungal allergies. Also noted is the production of antibiotics from fungal organisms such as
Penicillin, griseofulvin and antiamoebin, cheese and other milk products made through fungal
activities such as the Camberbert and Roqueforti cheese, soya sauce. A number of drugs are also
manufactured through fungal activities such as ergotamine and cortisone. Fungi are also used
industrially in the manufacture of many products such as organic acids, enzymes, fats, steroids,
anti-carcinogenic compounds and feed supplements.

Fungi play very important role in recycling of nutrients through decomposing of plant and animal
bodies and liberating various elements such as Sulphur, nitrogen, phosphorus, potassium, iron,
magnesium, calcium and zinc. Without fungal and some bacterial activities such elements would
be locked in these bodies and not be available for reuse for autotrophs to build up new organic
matter. Their multiple and complex enzymes enable them to attack organic substrates of different
kinds thereby releasing the simple elements back to the soil and thus enhancing soil fertility.
Why study mycology/Importance of Fungi?

Fungi are important to man in a number of ways:

 Important food source

 Important role in industries, such as pharmaceutical, food, enzymes,
 Disease agents
 Provision of important chemical compounds used in agriculture and industries
 Environment cleaning
 Offer tools for various scientific investigations and discoveries
 Ecological role in plants growth

1.2 Development of the science of Mycology

From its earlier times where the focus of fungi study was only on the large macroscopic fungi, the
study has undergone tremendous development especially with the discovery of the microscope.
Earlier studies mainly focussed on collection, description and illustration of macroscopic fungi.
These studies mainly concentrated in Europe (Greece, Italy, and the Netherlands, among other
countries). Some of the people credited with this work include C. E. Clusius who in 1601
described many mushrooms through artistic illustrations, van Sterbeck who in 1675 published a
book on the identification of edible fungi to aid in differentiating them from the poisonous forms.

A major step in the study of fungi was made with the development of microscopy by
Leeuwenhoek (1632 – 1723) which made it possible to observe microscopic organisms, fungi
included. The first illustration of microfungi was made by Robert Hooke (1635 – 1703) who drew
the sporangia of Mucor and the teliospores of the rose-rust Phragmidium mucronatum. P. A.
Micheli (1679 – 1737) made much use of the microscope through which he extensively studied
many fungi and their reproductive structures. In 1729, Micheli published his work which made
him to be reputed as the Founder and Father of Mycology. He proved experimentally that fungi
originate from their spores.

Another major contributor to the study of mycology is Carl Linnaeus who though did not directly
appreciate the importance of fungi as a group, his idea of binomial nomenclature greatly
contributed to the orderly naming of the fungi. Prior to this, fungal nomenclature was very
cumbersome with very long descriptive names. Taking up on this, a number of scientists focussed
on fungal nomenclature with the most notable being E. M. Fries (1794 – 1878) who published a
book Systema Mycologicum (1821) among other publications focussed on the systematics of
fungi. Fries came to be known as the Linnaeus of Mycology due to his tremendous contribution
to the taxonomy of fungi. The publication Systema Mycollogicum is still being regarded highly in
the systematics of mycology.

In the 19th century, through microscopic study of many fungi such as rusts, smuts, various
pycnidial fungi, great improvement on fungal identification was achieved. The idea of some fungi
existing in more than one morphological form at different environmental conditions or stages in
their life cycle, termed polymorphism, was established by L. R. Tulasne (1815 – 1885), a French
Botanist and Mycologist, who is also referred to as the “reconstructor of mycology”. L. R.
Tulasne and his brother Charles Tulasne (1816–1884) showed relationships between the conidial
stages and the sexual fruiting bodies (Perfect stage) of some fungal organisms. This was made
possible through the study of fungal life cycles that showed different morphological forms of an
individual fungus. The creation of the group of fungi that lack sexual stages, called “Fungi
Imperfecti” or the Deuteromycetes, was done by Fuckel (Karl Wilhelm G. L. Fuckel) a
German botanist who worked largely on fungi, in 1869. He also comprehensively studied the life
history of Myxomycetes.

Another major contributor to mycological studies was De Bary (Heinrich Anton De Bary, 1831
– 1888) who is also referred to as the founder of modern mycology. He made various studies on
the life histories of various groups of fungi, mechanism of parasitism, symbiosis and saprophytism,
the nature of lichens, among other biological characteristics. He discovered the phenomemnon of
heteroecism (need to exist in more than one host to complete life cycle) in rusts, especially in
Puccinia graminis tritici. He gave accounts of the development and sex in several fungi. He
wrote a book in 1866 titled the “Morphologie und Biologie der Pilze Flecten, und Myxomyceten”,
whose English translation was published in 1887 entitled “Comparative Morphology and Biology
of the Fungi, Mycetozoa and Bacteria”. De Bary is also called the father of Plant Pathology as
it was he who proved that the fungus Phytophthora infestans was the cause of the potato blight
disease.
Further contribution on fungal taxonomy was made by an Italian mycologist, P. A. Saccardo
(1845 – 1920). He brought together the scattered knowledge on systematic mycology under one
massive 25 volume document called the Sylloge Fungorum Omnium hucusque cognitorum in
1882. This work, commonly only referred to as the Sylloge, is still very much relevant in the study
of fungi. He classified the Fungi Imperfecti (Deuteromycetes) based on spore groups which is
extremely popular and being referred to by many mycologists up to now.

Other major contributors include: Robert Koch (1843 – 1910) a German Microbiologist who came
up with the theory of plant diseases, most of which are caused by fungal organisms. In the 19 th
century, lots of development on physiology, ecology and genetics of fungi was made through
intensive studies by various mycologists and other scientists using fungi as tools for their research.

The modern application of fungal organisms in many industrial processes is attributed to these
studies. Some of the major developments attributed to these organisms include,

1. use of fungi in commercial production of enzymes:

2. developments in fungicidal research and production of materials
3. fungal transformations in soils
4. role of fungi in food manufacture
5. role of fungi in food spoilage
6. role of fungi in rotting of textile fibres
7. fungi and antibiotics production
8. control of fungal and bacterial diseases in plants, animals and also human
9. production of novel therapeutic compounds to control some diseases such as cancer
10. study of genetic processes through the use of fungal organisms
11. use of fungal organisms in the production of hormones

Lots of studies still continue presently, especially on fungal physiology; basis of respiration,
growth factors, nutrition, enzymes, intermediary products of metabolism, etc.
Topic Summary

In this topic, you have learned the meaning and importance of the subject Mycology. The
development of this subject from being used to study mushrooms and toadstools for their
identification, to the understanding of fungal morphology and reproduction process to their
physiological processes. The contributions made by a number of scientists such as Leeuwenhoek
and microscopy, Robert Hooke and P. A. Micheli and their observation of fungal microscopic
structures, Linnaeus and naming of organisms which was built on by Fries and Tulasne brothers
in improving fungal nomenclature. Other scientist, P. A. Saccardo made major contribution to
Fungal taxonomy through his publication of many volumes of his work. The contribution of Robert
Koch from his work on plant diseases caused by fungal pathogens is also very significant in the
mycological studies. From these studies and many other modern studies, use of fungal organisms
in solving human challenges has been greatly promoted in 20th and 21st century where many novel
products, some therapeutic have been product through biotechnological manipulation of some
fungal organisms.

Glossary

Mycology - the scientific study of fungal organisms.

Heteroecisms – having more that one host in a life cycle

Further Reading

Mandatory Reading
An Introduction to Mycology. By R. S. Mehrotra and K. R. Aneja. New Age International
Publishers 1996 (Reprint 2005).
Optional Reading

Topic Activity

Collect different types of organisms that you think are fungi and state all the visible features that
make you qualify them as Fungi. How different are these organisms’ from higher plants that you
can see around you within your home or University compound.
Assignment [this assignment requires submission]

Write a short essay, describing six personalities who made significant contribution to the study
of Mycology, describing the contributions they made and how these contributions impacted on
mycological studies.
 TOPIC TWO: FUNGI AND THEIR CHARACTERISTICS

2.1 Introduction

In this Topic, you will study the characteristics associated with the fungal organisms, their habitats
and the adaptations they have to enable them survive in these habitats. Fungal morphological habit
from the plasmodial forms associated with the Myxomycota, the unicellular/yeast forms to the
more advanced filamentous mycelia forms will be studied. Nature of the hyphae, septate or
aseptate forms, nuclear oragnisation in funal cells such as uninucleate, multinucleate,
homokaryotic and heterokaryotic cells will be discussed. Further, the fungal tissues termed the
Plectenchyma and its forms will be explained. Associations in fungi including parasitism,
mutualism and lichen organisations will be explained.

Learning Outcomes

By the end of this topic you should be able to:

1. Describe the major characteristics of Fungal organisms.

2. Differentiate fungi from other four Kingdoms of Life.
3. Describe the different forms of fungal thallus.
4. Describe the cellular and hayphal organization in fungal organisms.
2.1.1 What are Fungi

Fungi are a diverse group of organisms not easy to be given a single definition. They bear various
characteristics associated with nutrition mode, thallus organisation, cell wall structure, nuclear
organisation, life cycle, sexuality, their habitat and distribution.

2.2: Characteristics of Fungi

1. All are eukaryotic

Possess membrane-bound nuclei (containing chromosomes) and a range of membrane-bound

cytoplasmic organelles (e.g. mitochondria, vacuoles, endoplasmic reticulum).

2. Most are morphologically filamentous

Composed of individual microscopic filaments called hyphae, which exhibit apical growth and
which branch to form a network of filaments called a mycelium. Some are however unicellular
e.g. yeasts.

3. Fungal cells are surrounded by a rigid wall which is composed primarily of chitin and glucans,
although the walls of some species contain cellulose.

4. Many reproduce both sexually and asexually both of which often result in the production of
spores.

5. Their nuclei are typically haploid and hyphal compartments are often multinucleate. However
some members of the Oomycota and some yeasts possess diploid nuclei.

6. All are achlorophyllous, lacking chlorophyll pigments and are incapable of photosynthesis.

7. All are chemoheterotrophic (chemo-organotrophic). They utilise pre-existing organic sources of

carbon in their environment and the energy from chemical reactions to synthesise organic
compounds they require for growth and energy.

8. Possess characteristic range of storage compounds e.g. glycogen, sugar alcohols and lipids.

9. May be free-living or may form associations with other organisms i.e. parasitic or mutualistic
(symbiotic).

2.2.1 Difference between fungi and other organisms

a. Fungi contain membrane-bound nucleus (like plants and animals but unlike bacteria), so
they are eukaryotes (Greek Eu = true; karyon = nucleus).
b. They contain cell wall made of chitin (unlike plant cells that have cellulosic walls and
animal cells, which have no cell wall).
c. They have no chlorophyll (unlike plants and algae which possess the pigment), so they are
not photoautotrophic, but they obtain energy from beakdown of organic matter from
their environment, hence they are heterotrophs.
 d. They do not ingest food, but secrete enzymes, which digest food outside of their bodies,
and then absorb obtained nutrients (absorptive heterotrophy), unlike animals which carry
out ingestive heterotrophy.

2.3 Thallus organisation

 Filamentous: Hyphae that forms mycelium

 Unicellular (yeasts)
 Plasmodium
 Pseudoplasmodium

The fungal body is mostly made up of undifferentiated filamentous structure called the hyphae.
Unicellular forms are rare, mainly represented by the yeast (Saccharomyces). The hypha is a
microscopic tubular filament. The mass of hyphae is called the mycelium (pl. mycelia). The
hyphae are usually branched (rarely unbranched), with the branches ramifying on or inside the
subs tratum, to form a three dimensional network.

Mycelial (filamentous):

Lateral branches arise behind the apex in acropetal order which each of these daughter brancjes
behaving as the parent hyphae and forms secondary and tertiary branches behind their growth tips.
The mycelium extends over or through whatever substrate the fungus is using as a source of food.

Hyphae may be septate or aseptate. The aseptate hyphae lack cross walls, termed septa, between
the cells hence have nuclei scattered in the cytoplasm making them multinucleate, a condition
termed coenocytic.
Fig. Aseptate (Coenocytic) and Septate hyphae

However, even in the aseptate hyphae, septa are normally formed to cut off old, empty portions of
the hyphae in the hinder regions and to delimit the sex organs. Such septa are formed due to
changes in the local concentration of cytoplasm and are called adventitious septa. The other
ordinary type of septa normally formed in association with the nuclear division, are termed the
primary septa. The primary septa produce the septate hyphae which result into cells that have
roughly comparable sizes. The adventitious septa are usually complete while the primary septa
have a pore in the centre allowing cytoplasmic continuity between the cells of the hyphae.

Other than the unicellular forms, other thalli observed in fungi include the plasmodium which is a
multinucleate mass of protoplasm without a cell wall. Such a thallus is found in the Myxomycetes
and the Plasmodiophoromycetes. The other thallus type is the amoeboid plasmodium, which is
formed by aggregation of unicellular, uninucleate, naked cells. The aggregated cells look like
plasmodium and is called the pseudoplasmodium. Such thallus is found in two groups of
organisms, the Acrasiales and the Labyrinthuales, which are placed alongside the Myxomycetes
and Plasmodiophoromycetes in the sub-division Myxomycota.

2.3.1 Fungal cell structure

Fungi have eucaryotic cell structure. They have double membrane-bound cell organelles like
nucleus and mitochondria, tubular endoplasmic reticulum (ER), the golgi bodies and the
ribosomes. Their ribosomes lie free in the cytoplasm and are not attached to the ER as is common
in most other eucaryotic cells of other organisms. Another structure found in the fungi cells are
the lomasomes which appear as particles or vesicles formed between the cell wall and the
plasmamembrane. Their function is however not known.

Figure: Fungal Cell Ultrastructure (from StuddyAdda.com)

Fungal tissues are called the plectenchyma. These tissues are formed by branching, interweaving
and coiling of the hyphae. Fungal tissues are relatively simple and limited in variety. They differ
mainly in the compactness of the organization of the hyphae and the degree of division into short
cells. Two types of plectenchyma are identified: The pseudoparenchyma and the prosenchyma.

The prosenchyma is a type of tissue which has loosely woven hyphae.

The Pseudoparenchyma is a type of tissue that has closely packed hyphae which show some
resemblance to the parenchyma of higher plants.
Figure: Types of Fungal tissues (Plectenchyma) (Pseudoparenchyma and Prosenchyma)

Plectenchymatous tissues in higher fungi are formed in somatic structures such as mycelial
strands, rhizomorphs, pseudorhizas, sclerotia, pseudosclerotia and stromata. These structures
are anatomically divisible into two regions: an outer hard rind called the cortex, which is made of
pigmented pseudoparenchyma and the inner soft, hyaline, prosenchymatous core, called the
medulla.

2.3.2 Cell wall structure

Fungal cell walls contain 80 – 90% carbohydrates; the remainder being proteins and lipids. The
fungal cell wall contains chitin (which is also the constituent of the exoskeleton of insects) and
not cellulose, the universal cell wall material of plants. Cellulose however occurs in the cell walls
of the Oomycetes and Hyphochytridiomycetes. The chitin wall is accompanied by glucose
polymers other than cellulose and mannans. The composition of cell wall is variable and is
modified by age and environmental factors.

Chitin is a linear polymer of N-acetyl β-glucosamine units linked by β 1, 4 glycosidic bonds. The
chitin may be overlain or intermeshed with other glucans, like callose, which consists of β-1, 3-
linked glucose residues.

2.3.3 Fungal nuclear status

Nuclei of fungi show some distinct characteristics that differ from other eukaryotic organisms.

 Their nuclei are generally very small and not easily visible, almost below the limit of
resolution of the light microscope.
 During mitosis, the nuclear membrane does not disintegrate and instead the nucleus divides
by constriction of this membrane in the centre.
 This type of nuclear division is termed karyochorisis
 In Karyochorisis, nuclear membrane remain intact as the chromosomes separate and move
to the opposite poles
 Karyochorisis is also termed as a closed mitosis
 First the invagination of inner nuclear membrane occurs that divides nucleoplasm into two
parts. Then subsequent invagination in outer nuclear membrane results in formation of
daughter nuclei.

Meiotic division in fungi is however similar to that of higher plants and animals

In most fungi, the nuclei are haploid throughout the life cycle with the diploid stage only found in
the zygote nucleus which does not multiply. Prolonged diploid phase is however found in some
fungi like the Oomycetes and yeasts.

In some higher fungi like the Ascomycetes and the Basidiomycotes, the nuclei of opposite mating
types may be found in one cell of the hyphae giving a condition termed Dikaryon.

In some cases, a cell in the hyphae may contain more than one nuclei which are genetically
dissimilar. The occurrence of such genetically dissimilar nuclei in the same cell is termed
Heterokaryosis. This phenomenon is common in many fungal groups and plays a major role in
bringing about variability and sexuality in fungi. Heterokaryosis is brought about by three main
ways:

 Mutation in the nuclei

 Anastomosis: which refers to the fusion of hyphae most of which are intra-specific
 Inclusion of dissimilar nuclei in some spores.

2.4 Nutrition in Fungi

Fungi show heterotrophic mode of nutrition, depending on the external supply of food. They are
chemoheterotrophic (chemo-organotrophic) - synthesising organic compounds they need for
growth and energy from pre-existing organic sources in their environment, using the energy from
chemical reactions. Having a rigid wall, fungi obtain their nutrients through absorption of small
molecules (e.g. simple sugars, amino acids) in solution across the fungal wall and plasma
membrane.

Larger, more complex molecules (e.g. polymers such as polysaccharides and proteins) are first
broken down into smaller molecules, which can then be absorbed. This degradation takes place
outside the fungal cell or hypha by the extracellular enzymes. Water is essential for the diffusion
of extracellular enzymes and nutrients across the fungal wall and plasma membranes, hence
actively growing fungi are usually restricted to moist environments.

Figure: Extracellular digestion and absorptive nutrition in fungi.

Three types of heterotrophy are shown by fungi namely:

Saprophytism

Parasitism

Mutualism

2.4.1 Saprophytism

Mode of heterotrophy in which fungi get their nourishment from the dead organic matter in their
surroundings. Such fungi are termed as saprophytes or saprobes. Saprophytes play a crucial role
in nature by decomposing dead organisms and releasing their nutrients for reuse by the living.

2.4.2 Mutualism/Symbiosis is a type of heterotrophy in which organisms live in a mutually

beneficial symbiotic relationship with one another, often involving energy transfer. These types of
relationship are also termed associations in fungi. Fungi show two types of mutualism, one with
algae and the other with roots higher plants, forming structures called Lichens and mycorrhizae.

Lichens

Lichens are structures made up of two different organisms, a fungus and either a cyanobacterium
or green algal cells.

In lichens, the fungal partner form distinct layers of hyphae that trap the photosynthetic algal cells
in the middle layer. The photosynthetic algal cells provide organic matter through photosynthesis
for fungal partner in the lichen.
The fungal partner prevented the algal cells from desiccation by trapping moisture. Recent
evidence indicates that the photosynthetic cells may grow faster when separated from the fungus.
Perhaps the fungus is parasitizing the photosynthetic cells.

Lichens derive most of their water and minerals from rainwater and air. This allows them to survive
on bare rock, tree trunks, inhospitable places.

Lichens are so efficient at absorbing nutrients from the air that they can be used to monitor air
quality because some kinds do not survive in polluted air.

They play an important ecological role: breaking down rocks and starting the process of soil
formation.

Mycorrhizae

A mycorrhiza is a mutualistic relationship between a fungus and a plant root.

The fungus functions like a root by growing into the soil and absorbing nutrients for the plant. The
plant provides the fungus with products of photosynthesis (sugar).

Many plants do not do well or do not grow at all without the fungi. Approximately ninety percent
of all plants develop mycorrhizae. Two types of mycorrhiza are identified: ecto and endo-or
arbuscular mycorhizzae

Ectomycorrhizal Fungi

Ectomycorrhizal fungi form a dense network of hyphae around plant roots. The hyphae may
penetrate the root, but they do not penetrate the root cells.

Ectomycorrhizal fungi have been described as the "dominant nutrient-gathering organs in most
temperate forest ecosystems" because nearly every tree in temperate and northern forests form
these associations with fungi.

Arbuscular Mycorrhizal Fungi

The hyphae of arbuscular mycorrhizal fungi (endomycorrhizal fungi) penetrate the plant root cells.
The portion of hyphae within the plant cell forms a highly-branched structure called an arbuscule,
which aids in the transfer of nutrients between the two species.

They extend into the soil and absorb water and minerals for the plant. The plant provides the fungus
with sugar.

Arbuscular mycorrhizal fungi are common in grasslands and tropical ecosystems. They are found
in eighty percent of all terrestrial plants.

Mycorhizae benefits

The plant benefits by more-efficient mineral (especially phosphorus) uptake while the fungus
benefits by the sugars translocated to the root by the plant.

2.4.3 Parasitism

Some fungi are parasitic, living in or on another organism (called a host) from which they obtain
their nutrients, causing serious damage to these hosts, both plants and animals. Parasitism is the
type of relationship in which one organism, the parasite benefits from the relationship, while the
host gets no benefit but supplies the parasite with nourishment while the parasite may also
produce harmful chemical compounds to the host. Parasitic fungi usually have specialized tissues
called haustoria that penetrate the host's body for food absorption.

Topic Summary

This topic has explained to us the major Fungal characteristics that include their eukaryotic nature,
achlorophyllous, heterotrophic, extracellular digestion followed by nutrients absorption that
require moisture, thallus organization that ranges from plasmodium, yeast and mycellial structure
and also tissue organization termed the plectenchyma. The nature of the cell wall that has chitin
and cellulosic materials was also described and the unique nuclear division without nuclear
membrane degredation termed Karyochorisis were all explained. Finally the different modes of
heterotrophy starting from saprophytism, parasitism and mutualism were explained with the two
common examples of mutualism where the structures Lichens and Mycorrhizae are formed were
also discussed.

Glossary

Eucaryotic cell – cell that bears membrane bound cellular organelles such as nucleus,
mitochondria, plastids,

Achlorophyllous – cell lacking the chlorophyll pigment.

Heterokaryotic – a condition of the cell bearing two nuclei which are genetically dissmilar

Plectenchyma – the term used to refer to the fungal tissues.

Prosenchyma – type of fungal tissue with loosely fused hyphae that make them soft to feel.

Mycorrhizae – root structure formed through the symbiotic growth of some fungi and the roots
of trees

Vessicular Arbuscular Mycorrhizae (VAM) – Intracellular mycorrhizae that grow and make
the cytoplasm swell into a bulged structure.

Anastomosis – coming of two or more hyphae together that fuse to form one single hyphal
thread.
References: