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Unit - 1

Origin of Life and Viruses

CONTENTS Learning Outcomes: At the end of


the unit learners should be able to :
1. Origin of life : Five
kingdom 1. Understand the origin of life from
classification. non-living matter of primitive
2. Discovery of earth.
microorganisms. 2. Know the basic characteristics of
3. Viruses -Shape and life forms in five kingdom
symmetry, TMV and classification of Whittaker.
Gemini virus; 3. Acquire the knowledge on
Multiplication of chronological events that led to
TMV; Prions and the discovery of microbes.
Viroids.
4. Understand and explain the living
4. Plant viral diseases, and non-living characteristics of
transmission and viruses.
control.
5. Develop critical knowledge on
5. Viruses- vaccine plant disease symptoms due to
production, bio- viral pathogens.
pesticides and
cloning vectors. 6. Realize the value of viruses in
medicine, agriculture and modern
biological fields.
Origin of life: Concept of primary Abiogenesis:
Chapter - 1
Five Kingdom classification.
1.1 Origin of Life:
Learning Outcomes: At the end of the chapter you should be
able to :
1. Develop a critical understanding on abiogenesis theory
with supporting experimental results.
2. Explain the merits and demerits of R.H. Whittaker’s five
kingdom classification.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 1


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




 

        

        
      
        
   
  




Fig. 1 Fig.1.1 Geographical regions on earth showing evidences on the origin of the earliest
life forms
https://upload.wikimedia.org/wikipedia/commons/a/aa/
Champagne_vent_white_smokers.jpg
https://en.wikipedia.org/wiki/Abiogenesis#/media/File:Stromatolites.jpg

2 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Introduction:
(= body). Life signifies a state of animation; it is a condition that
distinguishes biological organisms from non-living matter,
concerning capacity for growth, reproduction, and metabolism
preceding death. The theory of biogenesis states that ‘life can arise
from pre-existing life’.

Primary abiogenesis : This concept refers to the origin of life


from non-life on the earth, and it is contradictory to biogenesis.
The earliest life forms on the earth were much simple which led to
the origin of highly complex forms of life through evolution. A. I.
Oparin (1923) and J. B. S. Haldane (1928) independently
hypothesized the abiogenesis concept. This is also called chemical
origin of life or naturalistic theory or Oparin-Haldane theory.
Oparin and Haldane opined that life might have originated 3.5
billion years ago, from some non-living organic compounds in the
oceans of primitive earth through a series of chemical reactions.
They stated that the initial atmosphere on earth was of reducing
type, which was hydrogen rich and oxygen poor (Fig.1.1.1).
According to them:
i. In the primitive atmosphere of primitive earth, molecules
tend to donate electrons; energy favouring chemical
reactions may be due to high temperature, lightning and
solar radiation.
ii. Simple inorganic molecules could have reacted to form
organic compounds such as amino acids and nucleotides and
are gathered in the oceans as a ‘primordial soup’.
iii. The process of transformation of the non-living chemicals
in to living matter extended over a billion years.
iv. Abiogenesis is not possible under present conditions,
because presently the earth has an oxidizing atmosphere.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 3



  
        
  
 

        

   
   
 
      

  

        
   

Fig.1.1.2 Apparatus used by Miller and Urey to create conditions on the primitive earth.
Source: Miller-Urey_experiment-en.svg; GFDL, CC-BY-SA license

4 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Lederberg considered 3 stages i.e., chemogeny, biogeny, and
cognogeny in origin of life through abiogenesis. About 4 billion years
ago, the earth’s atmosphere had ammonia, methane and water
vapours; there was no free oxygen. Various simple and organic
molecules were formed from those gases.This is called prebiotic
evolution. During chemogeny simple organic compounds like
hydrocarbons, carbohydrates, fatty acids and glycerol, and amino
acids were formed. They underwent condensation, polymerization,
and chance chemical reactions to form complex organic molecules
and colloids. Those colloidal molecules aggregated into coacervates,
proteinoid microspheres, protocells, and liposomes depending on
their composition. In the next stage nucleic acids (naked genes) and
nucleoproteins were formed. At the early stage of biogeny, the RNA
containing nucleoproteins changed into bubble-like structures,
protocells (eobionts or protobionts). During the biogeny, microbes
with varied modes of nutrition are evolved. Cognogeny evolution
of mechanism of perception, expression and communication formed
eukaryotes. Evolution of eukaryotes might have occurred by
endosymbiosis (Lynn Margulis, 1990) or by invagination of the cell
membrane of primitive prokaryotic cells.

Activity: Discuss and debate with your peer groups on different


theories about origin of life by collecting material from standard
reference books/web resources. Collect photos of the
hydrothermal vents, volcanoes etc., with the conditions resemble
the pre-biotic earth.

Miller and Urey experiment: S. Miller and H. Urey (1953)


conducted an experiment to test the Oparin-Haldane hypothesis.
They devised a simple apparatus containing water and reduced gas
mixture, created a prebiotic environment on the planet earth to
demonstrate abiogenesis. The apparatus consisting of two glass

APSCHE : SEMESTER-I BOTANY TEXT BOOK 5


      

         
  
        

       
 



    
U

         


 



  

 



6 APSCHE : SEMESTER-I BOTANY TEXT BOOK


flasks (a larger, 5L and a smaller, 500 ml) connected together by
two glass tubes forming a loop (Fig.1.1.2). The big flask has a pair
of electrodes; one of the connecting glass tubes provided with
condenser and stopcock.

Miller and Urey added methane (CH4 3


), and
) in a sterile and fully sealed larger flask (represents
2

atmosphere); those chemicals believed to be the major components


of the early earth’s atmosphere. Whereas the smaller f lask
(represents ocean) has half-full of water (H2O). The water in the

vapour was allowed to enter the larger flask. The flask with heated
water represents water on the earth’s surface; while the recycled
water vapour represents water that evaporates from lakes and seas.
Continuous electrical sparks (800°C) were fired between the
electrodes to simulate thunder lightning (believed to be very
frequent on the early earth) in the water vapour and gaseous
mixture. Consequently, simulated atmosphere was cooled; so that
the water condensed and trickled (represents precipitation) back
into smaller flask via a U-shaped tube at the bottom of the apparatus
in a continuous cycle.

Fig. 1.1.3 Schematic diagram showing


the 5-kindoms in Whittaker’s
classification
Source: https://classification-of-
organisms.fandom.com/wiki/
Kingdom?file=Classification.jpg

APSCHE : SEMESTER-I BOTANY TEXT BOOK 7


After a day, the solution collected at the trap had turned to
pink colour and after 7 days of continuous operation, the solution

mercuric chloride was added to prevent microbial contamination.


The reaction was stopped by adding barium hydroxide and sulfuric
and evaporated to remove impurities. Miller identified, five amino
acids i.e., glycine,α β β-

chromatography. In addition to amino acids, simple organic


compounds like hydroxy acids, aliphatic acids, sugars and urea were
also identified from the cooled water. Thus, this experiment clearly
demonstrated that some building blocks of life can be produce in a
cell free environment or without living-being.
Five Kingdom Classification:

Introduction: Robert Harding Whittaker was an American plant


ecologist. He proposed a broad taxonomic classification of living
organisms in 1969. Whittaker system of classification is simple and
conceptually coherent. His approach of classification was functional
kingdoms, primarily ecological and only secondarily taxonomic
groupings. It reflects the three major directions of evolution and
ecological functions i.e., producers, consumers, and decomposers.
It is based on two fundamental characteristics of living organisms
i.e., mode of nutrition and cellular organization. The main criteria
for his classification include cell structure, body organization, mode
of nutrition, reproduction and phylogenetic relationships.
Accordingly, all the living-beings were grouped into five - kingdoms
viz., Monera, Protista, Fungi, Plantae and Animalia.

Kingdom Monera: The organisms in Monera are prokaryotic and


unicellular and cosmopolitan in distribution. Some organisms live

8 APSCHE : SEMESTER-I BOTANY TEXT BOOK


      

       
    
     


     
      
  

      
       





     
       
   


APSCHE : SEMESTER-I BOTANY TEXT BOOK 9


in extreme ecological conditions. Membrane bound cell organelles
including nucleus are absent in these organisms. The cell walls are
composed of pseudomurein or peptidoglycans or mycolic acid.
Cell walls are absent in mycoplasmas. The cell membrane contains
proteins and lipids. The cells may be flagellated or non-flagellated;
may be motile or non-motile. Except the cyanobacteria and a few
true bacteria, all others are heterotrophs; may be saprophytes,
parasites or symbionts. Reproduction is mostly by cell division.
Archaebacteria, eubacteria, cyanobacteria, mycoplasmas and
actinomycetes etc., are included in this kingdom.

Kingdom Protista: Unicellular and eukaryotic organisms are


placed under Protista. The members of this kingdom are mostly
aquatic and ubiquitous. The cell contains a well-defined nucleus and
membrane bound cell organelles. The cell wall is composed of silica
or cellulose plates; in euglenoids pellicle made of protein is present.
The protozoans and slime moulds lack cell walls and have plasma
membranes only; but spores of slime moulds show cell walls. The
plasma membrane is composed of proteins and lipids. The cells may
be motile with the help of flagella or cilia or pseudopodia; or may
be non-motile. The protists have varied types of nutrition, may be
autotrophic (photosynthetic) or heterotrophic. Reproduction is
mostly binary fission rarely sexual. Chrysophytes, dinoflagellates,
euglenoids, slime moulds, protozoans etc., are included in this
kingdom.

Kingdom Fungi: This kingdom comprises of eukaryotic organisms


with unicellular or multicellular body. The fungi are world-wide in
occurrence, live mostly in warm and humid places on the earth.
The cell contains conspicuous nucleus and membrane bound cell
organelles. The cell walls are made up of chitin and polysaccharides.

10 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Plasma membranes are made up of proteins and lipids. The
organisms are generally non-motile; but spores and gametes are
flagellated and motile. This kingdom is unique, consists of
heterotrophic organisms only; may be saprophytes or parasites or
symbionts. Reproduction occurs by vegetative, asexual and sexual
methods. This kingdom includes Phycomycetes (e.g., Rhizopus),
Ascomycetes (e.g., Yeasts), Basidiomycetes (e.g., Puccinia) and
Deuteromycetes (e.g., Colletotrichum).

Kingdom Plantae: All eukaryotic, multicellular and chlorophyll


containing plants are included in this kingdom. Organisms of this
group live in different aquatic and terrestrial ecosystems of the
world. The cells are obviously nucleated and contain different cell
organelles including chloroplasts. The cell walls are made of
cellulose. Almost all the plants are autotrophic (photosynthetic);
but insectivorous (e.g., Nepenthes) and hemi-parasitic (e.g., Cuscuta)
plants are partial heterotrophs. The plants are sedentary, but spores
or gametes of some plants are motile, in some case carried away by
abiotic factors or biotic organisms. Growth is usually indefinite by
means of apical cells or meristems. Reproduction is by means of
vegetative, asexual and sexual methods. Algae (except unicellular
forms), Br yophytes, Pteridophytes, Gymnosperms and
Angiosperms are included in this kingdom.

Kingdom Animalia: Eukaryotic, multicellular and heterotrophic


animals are kept in this kingdom. These organisms are inhabitants
of many aquatic and terrestrial ecosystems of the earth. The cells
are enucleated and possess all cell organelles except chloroplasts.
The cell wall is lacking and the cells are bounded by plasma
membranes. Nutrition is holozoic and depend on plants for their
nutrition, either directly or indirectly. The ingested food is digested
in an internal cavity, consequently stored as glycogen or fat. Most

APSCHE : SEMESTER-I BOTANY TEXT BOOK 11


of the animals are capable of locomotion. The growth is definite
and attains definite shape and size after a certain period of growth.
Evolutionarily advanced animals have an elaborate sensory and
neuromotor mechanism. Sexual reproduction by copulation is
predominant.

Objections to Five Kingdom classification: The inclusion of


unicellular algae (plants) and protozoa (animals) in the kingdom
Protista is not justifiable. In this system of classification, a
distinction between unicellular and multicellular algae is not
possible. Kingdoms Monera and Protista have diverse organisms,
keep them together is not justifiable. For example, those kingdoms
consist of organisms with and without cell walls; photosynthetic
and non-photosynthetic organisms; cellular or filamentous
organisms. Virus has not been included in any of the 5 kingdoms.
Archaebacteria differ from other bacteria in structure, composition
and physiology. Mycoplasma are quite different form bacteria where
they have been placed along with prokar yotes. Symbiotic
associations are not considered in this classification system. For
example, lichens are organisms which are formed by the symbiotic
association between fungi and algae.

Activity: Conduct a group discussion among your peers on various


classification systems of life forms i.e., 2-kingdom to 6-kingdom,
discuss and debate on their merits and demerits. Collect photos on
various life forms in 5-kingdoms of Whittaker and make an album
to acquire an in-depth knowledge.

12 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Discovery of microorganisms, Pasteur
Chapter -2
experiments, germ theory of diseases.

Learning outcomes : At the end of the chapter you should be able to :


1. Realize the importance of microscope in exploring microbial
world.
2. Gain knowledge on the contributions made by various
microbiologists.

1.2.1. Discovery of Microorganisms:


Introduction : Microbe is a French word, which derives from two
Greek words, Mikros (=small) and bios (=life). The existence of
microorganisms was predicted many centuries ago; the Jain
scriptures from 6 th century B.C. mentioned that ‘nigodas’ (=
microbes), pervade in entire universe and not visible to human eye.
Period Chronological events in discovery of microbial world
127 B.C Marcus Terentius Varro, a Roman scholar states that some
minute creatures enter the human body through the mouth
and nose, and cause serious diseases.
1025 A.D Avicenna, a Persian physician hypothesised the presence of
microbes in his book, “The Canon of Medicine”.
1546 A.D GirolamoFracastoro, an Italian physician states that
invisible, infectious and transferable entities cause the
epidemic diseases.
1590 A.D Zacharias Janssen was the first to invent a microscope (from
the ancient Greek, micro-small and scope-to look).
1625 A.D Galileo Galilei () crafted a compound microscope
1625 A.D Giovanni Faber coined the term microscope (from the
ancient Greek, micro-small and scope-to look).
1931 A.D E.Ruska and M.Knoll from Germany, invented electron
microscopev
Galileo Galilei’s compound microscope is called as ‘little eye’
and submitted to the Academia deiLincei. The science of
microbiology reached new horizons after the advent of electron
microscope by E.Ruska and M.Knoll from Germany in the year 1931.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 13



        
      

      

       
       
   


       
   
   
       

         

  

  
       
 
 

14 APSCHE : SEMESTER-I BOTANY TEXT BOOK
The credit for discovering microorganisms in the 17th century goes
to Robert Hooke and Antonie van Leeuwenhoek. They used simple
microscopes that magnified objects from about 25-fold to 250-fold.

Robert Hooke contributions : Robert Hooke was an English


polymath had the habit of examine various material (including
sponges, wood, seaweed, leaf surfaces, hair, peacock feathers, wings
of flies, eggs of silkworms, mites, a flea and a louse) of biological
origin, with the aid of a hand-crafted, leather and gold-tooled
microscope. His observations were recorded in the form an
illustrated book, Micrographia. This book contained detailed
account of 60 reflections. Leeches in vinegar and bluish mould upon
a mouldy piece of leather were two of his microscopic observations
presented at the Royal Society meeting in 1663. Hooke observed a
micro fungus from a ‘small white spot of a hairy mould’ on a book
cover made up of red sheep skin and He called the organism as
‘microscopical mushroom’ (Fig.1.2.1). Hooke thought the ‘knobs’
might be ‘seed cases’, and attempted to release and observe the
‘seeds’ (sporangiospores), but failed because they were too small to
be seen at the magnification he used. The microscopic view of the
mould published by him in Micrographia is now realized as of Mucor
with sporangia in different stages of development.

Antonie van Leeuwenhoek Contributions : Leeuwenhoek was


a Dutch business man regarded as the Father of Microbiology owing
to establishment of microbiology as a scientific discipline. He
designed and made 25 single-lens microscopes and explored the
microbial life from various material in the nature and reported to
the Royal Society, London. Leeuwenhoek (1677), gave the first
detailed description of protists in his famous paper ‘letter on the
protozoa’. In 1676, he scrutinized some ‘pepper water’ left idle for
3 weeks, identified bacteria. Both protozoans and bacteria were

APSCHE : SEMESTER-I BOTANY TEXT BOOK 15





   




Fig. 1.2.1 Microscope used by Robert Hooke and and his drawing from Micrographia.

A. Microscope used by Robert Hooke


Source: https://commons.wikimedia.org/ wiki/File:

RobertHooke,Micrographia,detail;_microscopeWellcome_M0005217.jpg
B. Microscopic view of a “hairy mould” colony described by Robert Hooke in 1665 (in

The reproductive structures (sporangia) are characteristic of the micro-fungus


Mucor. Sporangia in different stages are identified by the letters A, B, C, and D.
Hooke included a scale reference; the length of the bar under the diagram represents
1/32 of an inch.
Source: http://www.academia.dk/MedHist/Biblioteket/ Print/images/ scheme-
12.png

16 APSCHE : SEMESTER-I BOTANY TEXT BOOK


referred as ‘animalcules’ or ‘little eels’ by Leeuwenhoek (Fig.1.2.2).
In his 39th letter (1683), Leeuwenhoek described rods, cocci, and
spirochetes in scraping from his teeth. He computed the area,
volume, and number of microbes. He emphasized the role of microbe
in making wine, curd and cheese, and spoilage of food.

Fig.1.2.2 Microscope used by Leeuwenhoek and his drawings of micro-organisms.

Lazzro Spallanzani contributions : Italian scientist, Spallanzani


taken hay infusion in eight bottles and boiled all of them. Four of
the bottles were corked while other four made airtight.After a few
days he could observe growth of microbes only in corked bottles,
there were no microbes in the airtight bottles. Thus, he theorized
the contamination of corked bottles owing to the presence of
microbes in the air.
1.2.2. Louis Pasteur experiments : Pasteur was a French
microbiologist, well known for his explicit contributions in
Bacteriology; thus, honoured as the Father of Bacteriology. He
denies the theory of spontaneous generation by conducting several
experiments. He took a liquid broth in a flask, boiled the liquid

grew in it.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 17



   
 
   





         
    



 
   
 



       
     
        



18 APSCHE : SEMESTER-I BOTANY TEXT BOOK


experiments at high altitudes, he could observe growth of microbes
in a few flaks, suggesting less dust and occurrence of fewer microbes
at high altitudes. In another set of experiments, Pasteur used swan
neck flasks with added fermentable liquid (meat broth). Though
air was allowed into the flask through its swan neck, it remained
free of microbial growth for an extended period. It was presumed
that the neck prevented particles from falling into the broth. No
growth occurred in the broth unless the flask was tilted, allowing
the liquid to come into contact with the contaminated walls of the
neck. The broth was also contaminated when the swan neck was
broken. All these experiments clearly indicate that living organisms
which grew in such broths came from outside dust, rather than
spontaneously arising within the liquid or from pure air.

Activity: Prepare microbial culture media/broth, expose in open


air, set in a culture room for 1or 2 days observe various microbes
grown on such media using microscopes. Discuss with your
classmates on different microbe present in the surrounding
environment.

1.2.3. Germ theory of diseases :


Introduction : The existence of microbes and their role in
infectious diseases was not understood for many centuries. Till the
end of the 19th century, the miasma theory was the prime theory of
disease spread. It states that a venomous form of bad air (Greek
word miasma means pollution) deriving from decomposing organic
matter was responsible for infectious diseases. Against to this
concept many physicians like Avicenna (1025), Ibn al-Khatib (14th
century), GirolamoFracastoro (1546), Marcus von Plenciz (1762)

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stressed the role of microbes for infectious diseases, thereby formed
a basis for the germ theory of diseases (the pathogenic theory of
medicine).
The germ theory of diseases was taken a shape in the early
19th century; which had a scientific basis from the experimental
evidences by Louis Pasteur (1850) and Robert Koch (1880).
According this theory germs (microorganisms) or pathogens are
causing diseases. The word germ may refer to virus, prions, viroids,
bacteria, protists, fungi etc. Developments in Bacteriology and
Virology led to the recognition of the actual organisms that cause
many diseases, formed a strong basis for the advocacy of germ
theory of diseases.

Fig.1.2.3 Experimental Scheme of Louis Pasteur to demonstrate


the existence of micro-organisms.
Source: https://creativecommons.org/licenses/by-sa/4.0

20 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Louis Pasteur’s pasteurization experiment (c.f. 1.2.2 in this
chapter) illustrates the fact that particles in the air cause the spoilage
of liquid broth. His experiments gave significant evidences
supporting the idea of germ theory of disease. Antoine Bechamp
recognized that silk worm larvae suffering from pebrine (pepper
disease) were unable to spin silk thread. In 1870, Pasteur recognized
and isolated a microorganism (later identified as Nosema bombycis,
a microsporidian parasite) accountable for pebrine in silkworm
larvae. Pasteur technique for screening the silkworm eggs with
pepper disease saved the France’s silk industry; the same method is
still being used for control of silkworm diseases. Many people all
over the globe died due to diphtheria disease in 17th to 19th centuries.
Edwin Klebs (1883) identified the bacterium causing diphtheria,
Corynebacteriumdiphtheriae. Alexandre Yersin (1894) isolated a
bacillus bacterium (later named as Yersinia pestis, in honour the
discoverer) causing bubonic plague, a pandemic disease of humans.
Anthrax disease was more prevalent and caused death of live-
stock during 17th and 18th centuries. It is primarily a fatal skin disease
affecting live-stock, and also humans who have contact with infected
animals. The isolation and identification of anthrax pathogen,
Bacillus anthracis by Robert Koch (1877) was the first step towards
medical bacteriology; hence, he is honoured as the Father of Medical
Bacteriology. He worked out the disease cycle of anthrax bacterium.

This work of Koch completely ruled out the theory of


spontaneous generation and supported the germ theory of diseases.
He also revealed the disease cycle of tuberculosis caused by
Mycobacterium tuberculosis in 1882 and cholera caused by Vibrio
cholera in 1884 (Fig.1.2.4). Koch proposed 4 principles, popular as
Koch postulates, that explain the relationship between the pathogen
and disease.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 21


Source:https://commons.wikimedia.org/wiki/ Source:https://commons.wikimedia.org/wik
File:Cholera_bacteria_SEM.jpg i/File:Mycobacterium_tuberculosis.jpg

Fig. 1.2.4 Scanning Electron Micrographs of Vibrio cholera and Mycobacterim tuberculosis

Activity: Collect data on various diseases of plants and pathogens


by interacting with your teacher or a plant pathologist; similarly
interact with doctors to collect data on diseases of humans and
animals and their pathogens. Make a consolidated report with
preventive and control measures.

Koch postulates:
i. The microorganism must be found in plenty in all hosts
suffering from a disease, but should not in healthy
organisms.
ii. The microorganism isolated from a diseased organism and
grown in pure culture.
iii. The cultured microbe should cause disease when introduced
into a healthy organism.
iv. The microbe must be sub culture from the inoculated,
diseased experimental host, and
v. The microbe should be identical to the original specific
pathogenic agent.

22 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Implications of the germ theory of diseases : Consequent to
the advocacy of the germ theory of diseases, the emergence and
advancement of some branches occurred in biology and allied fields.
1. Preventive and control measures in both plant and animal
pathology, and also health management have their roots in
the germ theory.
2. Mining of disease resistant genes, breeding for disease
resistance in plants can be traced back to this theory.
3. The practices of disinfection, sterilization, personal hygiene,
and proper food preparation have their basis in germ theory.
4. Immunology and epidemiology emerged as explicit scientific
branches following this theory.
5. Advancement in vaccination, application of antibiotics to
prevent and control microbial diseases is also due to this
theory.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 23


Virus- shape and symmetry, structure and
Chapter -1.3
multiplication, Prions and virods

Learning outcomes : At the end of the chapter you should be


able to:
1. Explain the diversity in shape, structure and symmetry
of viruses.
2. Acquire knowledge on replication mechanism in
phytophages.
3. Understand the significance of prions and viroids in pre-
biotic world.

The word Virus (= venom) is derived from Latin. The branch of


microbiology that deals with viruses is called Virology. Martinus
Willem Beijerinck (1898) was the first to identify a new pathogen
of non-bacterial nature infecting tobacco plants, named it a virus.
He is regarded as the Father of Virology. He used the phrase,
‘Contagium vivum fluidum’ to describe the virus by thinking that
it was somewhat liquid in nature. Viruses are obligate intra
cellular parasites with a diameter of 20 to 300 nm. They are non-
cellular entities without independent metabolism and molecular
machinery for replication, often referred to as viral particles.
Viruses infect all life forms i.e., plants, animals, and
microorganisms. Viruses occur in all ecosystems of the earth. Of
the millions of types of viruses in the environment, about 9,000
species have been described in detail The virus has a nucleic acid
core (genome) surrounded by a protein coat (capsid), referred to
as nucleocapsid. The genome is either DNA or RNA, but not both.
In some viruses, an envelope made up of a bilayer of lipoprotein
and glycoprotein is present; they get lipoprotein layer from the
nuclear or plasma membrane of the infected host cell. Some
viruses comprise viral enzymes (lysozyme, reverse transcriptase)
that are essential for infection of a host cell and coded for within
the viral genome. The virus with all necessary components to
infect a host cell is called, a virion.

24 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig1.3.1 Shapes of Viruses

a) TMV b) Ebola virus c) Vaccinia virus

d) Rhabdovirus e) Corona virus f) Bacteriophage

g) Orf Vius h) Flexuous tailed phage i) Adenovirus

Source: https://commons.wikimedia.org/wiki/File:TMV_under_TEM.png
https://commons.wikimedia.org/wiki/File:Ebola_virus_(2).jpg
https://upload.wikimedia.org/wikipedia/en/f/fa/Poxvirus.jpg
https://upload.wikimedia.org/wikipedia/commons/5/56/Rhabdovirus.png
https://www.flickr.com/photos/niaid/49534865371/
https://commons.wikimedia.org/wiki/File:Bacteriophage.jpg
https://www.cdc.gov/mmwr/preview/mmwrhtml/mm5503a1.htm
https://manyworlds.space/2020/04/17/viruses-the-virosphere-and-astrovirology/
https://commons.wikimedia.org/wiki/File:Adenovirus_3D_schematic.png

1.3.1. Virus shape and symmetry


Virus shape : The virions are apparently of diverse shapes
(Fig1.3.1). They may be rod-shaped (TMV), filamentous (Ebola
virus), brick-shaped (Vaccinia virus), bullet- shaped (Rhabdovirus),
spherical (HIV, influenza, Herpes viruses etc.), tadpole-shaped
(Bacteriophages), elliptical and coiled (Orf virus), head and flexible

APSCHE : SEMESTER-I BOTANY TEXT BOOK 25




    
       

   
  

    

â

Genome in Viruses: Viruses are broadly classified as DNA Viurses
and RNA Viruses based on their genetic material (Baltimore, 1971).

Group-1: Viruses with dsDNA; mRNA is transcribed from DNA


(Small-pox virus).
Group-2: Viruses with ssDNA; mRNA is transcribed from DNA
(Human Papilloma Virus).
Group-3: Viruses with dsRNA; m RNA is transcribed from the
minus (–) strand of RNA (Rota viruses).
Group-4: Viruses with ssRNA (+sense); RNA itself acts as mRNA
(Corona viruses).
Group-5: Viruses with ssRNA (-sense); mRNA is transcribed
from RNA (Ebola virus).
Group-6: Enveloped viruses with two identical + strand ssRNAs;
DNA is transcribed from RNA (Retro viruses) and then
mRNA is synthesized on DNA (HIV).
Group-7: Viruses with dsDNA; RNA is synthesized on DNA and
back into DNA, it encodes for mRNA (Hepatitis-B
Virus).

26 APSCHE : SEMESTER-I BOTANY TEXT BOOK


tail (Flexuous tailed phage), Icosahedral with 12 spokes
(Adenovirus).
Symmetry of viruses : Viral symmetry is based on the
arrangement of capsomere units in the capsid. Basically, two types
of symmetry i.e., helical and icosahedral are recognized, respectively
corresponding to the spiral and spherical shapes of viruses (Fig.
1.3.2).

Fig 1.3.2 Symmetry of viruses


http://go.microsoft.com/fwlink/p/?LinkId=255141
https://basicmedicalkey.com/6-viruses-basic-concepts/

A. Helical (spiral) symmetry: Both capsomers and genetic


material together arranged in a spiral manner to form a rod like
structure. Viruses with this symmetry are mostly enveloped and all
of them possess RNA. Tobacco Mosaic Virus (TMV) with ssRNA
and 2,130 capsomers is a classic example for this symmetry. Viruses
causing measles, rabies, and influenza in human-beings are of helical
symmetry. Corona viruses are essentially of helical symmetry, but
appear in spherical shape due to the presence of envelope. Some
viruses having helical symmetry appear to be ‘open-ended’, while
in others the protein subunits seal the capsid at one or both ends.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 27



    
    
 
  

  
T2, T4 


      
       
       

x         




Fig 1.3.3 Complex symmetry of virus

28 APSCHE : SEMESTER-I BOTANY TEXT BOOK


B. Icosahedral (cubical) symmetry : An icosahedral is a polygon,
generally it consists of 12 vertices (corners), 20 facets (sides) and
30 edges. Each facet is an equilateral triangle. Icosahedral capsid is
the most stable, it is composed of pentagonal or hexagonal
capsomeres at the vertices. Human pathogenic viruses like
Adenovirus, Picornavirus, Papovavirus, herpes virus possess this
kind of symmetry. In such viruses, the protein subunits come
together to produce a geometric structure that is very close to being
spherical. Capsids in such viruses are always fully closed.This kind
of viruses are classified based on triangulation (T) number.
C. Complex symmetry: Some viruses display complex symmetry;
they are composed of a number of distinct capsomeres with discrete
shape and symmetry. Due to complexity of their capsid structure,
they do not have either icosahedral or helical symmetry. Pox virus
and some bacteriophages contain this symmetry. The T-even phages
(T2, T4 etc.,) have binal symmetry with an icosahedral head and a
helical tail (Fig.1.3.3).

Know more: The minimum number of capsomers required to


construct an icosahedron is 12, each composed of five identical
subunits forming the adjoining corners of the capsid
structure.The icosahedral structure displays manifold evenness,
with 2-fold axes running along each edge, 3-fold axes cantered
in each triangular face, and 5-fold axes at each corner. The
capsid in adenoviruses is highly distinguishable from that of
other viruses; it consists of 12 projecting fibres, one present at

APSCHE : SEMESTER-I BOTANY TEXT BOOK 29


  
  
   


  


Fig 1.3.4. Structure of TMV (1) ssRNA (2) Capsomer (Protomer) (3) Capsid
https://upload.wikimedia.org/wikipedia/commons/6/6d/
Tobacco_mosaic_virus_structure.png

Fig1.3.5. Gemini virus – Maize Streak Virus (MSV)

https://talk.ictvonline.org/ictv-reports/ictv_online_report/ssdna-viruses/w/geminiviridae

30 APSCHE : SEMESTER-I BOTANY TEXT BOOK


1.3.2 Structure of TMV and Gemini virus
Structure of Tobacco Mosaic Virus : Tobacco Mosaic Virus (TMV)
is a phytophage belongs to the genus Tobamovirus. TMV was the
first virus discovered by Dmitry Iwanowski (1887). W.M.Stanley
(1935) crystallized TMV, demonstrated that it remains active even
after crystallization. TMV infects members of Solanaceae family, in
particular tobacco. The infection causes mosaic like mottling and
discoloration on the leaves of host plants.
Rosalind Franklin and her associates (1953) have described
the structure of TMV. The virion is rod shaped with helical
symmetry. It is 3,000Å (300nm) in length and 180Å (180nm) in
diameter;and, its molecular weight is 40 x10 6 Daltons. It is a
nucleoprotein, consists of a centrally located Ribonucleic acid (RNA)
enveloped by a protein coat (capsid).
The capsid accounts for 94.4% of weight. It is composed of
2,130 grape like sub-units, capsomers that are arranged in 130
helical turns around a central hole of 40Å (4nm) radius. Therefore,
16.3 capsomers are present in every helical turn. Each capsomer is
a single polypeptide. It possesses 158 amino acids and has a
molecular weight of 17,500 Daltons. The amino acids in a capsomer
are assembled into four main alpha-helices, which are joined by a
prominent loop proximal to the axis of the virion.
The RNA (genetic material) is single stranded. The positive
sense RNA is un-segmented, helically coiled and accounts for 5.6%
of weight. The diameter of RNA helix is about 80 Å; the RNA
molecule lies about 50 Å inward from the outer-most surface of the
rod. This RNA consists of 6,600 nucleotides, its molecular weight
25,000 Daltons. There are 49 nucleotides in every helical turn and
3 nucleotides per a capsomere.The ssRNA is little more in length
(about 3300 Å) slightly protruding from one end of therod. The 3’

APSCHE : SEMESTER-I BOTANY TEXT BOOK 31


The family Geminiviridae consists of 9 genera and 360 species. Viruses
in the genera Becurtovirus, Capulavirus, Curtovirus, Eragrovirus,
Grablovirus, Mastrevirus,Topocuvirus and Turncurtovirus have a single
genomic component, whereas those in the genus Begomovirus have
either one or two components. Twinned virions contain a single copy of
circular ssDNA, ranging from 2.5 to 3.2 kb. Hence, for viruses with
bipartite genomes, two virions containing different genomic components
are required for infection, the total genome being about 5.2 kb. It is the
largest known family of ss DNA. These DNA viruses are transmitted by
insects such as leaf hoppers and white flies. Viruses of this family cause
Bright Yellow Mosaic, Yellow Mosaic, Yellow Mottle, leaf curling,
stunting, streaks, reduced yield in host plants.

  







 
       
 
    

   
    
    
 

32 APSCHE : SEMESTER-I BOTANY TEXT BOOK


terminus has a tRNA like structure and the 5’ terminus has a
methylated nucleotide cap. The 4 genes in the genome encode for
(a) Replicase with methyl transferase (b) RNA helicase (c) RNA
dependent RNA polymerase (movement protein) and (d) a capsid
protein (Fig.1.3.4).
Structure of Gemini Virus : Gemini viruses are non-enveloped
with icosahedral symmetry. They are classified under Class-II in
Baltimore (1971) classification. They belong to a family of plant
viruses, Geminiviridae. A virion consists of capsid of 30nm length
and 18-20nm diameter. The capsid is composed of 22 capsomers.
The name of the virus is derived from the geminate (twinned) capsid
i.e., capsid consists of 2 sub-units. The nucleic acid is circular single
stranded DNA. The genome may be non-segmented with 2500 to
3000 nucleotides. In some cases, the genome has 4800 to 5600
nucleotides long and consists of two segments. In such case, both
the segments must be transmitted to cause full systemic infection
in the host plants. The genome encoding genes diverge in both
directions from a virion strand origin of replication (i.e. geminivirus

and non-structural proteins (Fig.1.3.5).


1.3.3 Multiplication of TMV : TMV is an obligate intracellular
parasite that replicates using the molecular machinery of the host
cells; it multiplies within the infected host cells. The multiplication
of virus consists of 5 steps (a) Entry (b) Un-coating (c) Intracellular
development (d) Assembly and (e) Release
a. Entry: TMV cannot directly enter the host cells. The complete
virus penetrates and enters through damaged host cells.
b. Un-coating: After entry nucleic acid of the virus is released from
the capsid within the cytoplasm of host cells. The removal of viral
protein coat requires host enzymes.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 33


 


    
 
       
  
       

Fig 1.3.6 Schematic diagram of TMV Multiplication

34 APSCHE : SEMESTER-I BOTANY TEXT BOOK


c. Intracellular development: The viral RNA freed from capsid
enters into the nucleus (possibly into the nucleolus) of the host
cell. Replication of viral RNA occurs in 2 steps. The viral-RNA first
induces the formation of specific enzymes called ‘RNA polymerases.
A single stranded RNA (- sense) called as replicative RNA is
synthesized using the ss RNA (+ sense) of TMV as a template strand.
This replicative RNA strand is complementary to the viral genome
and serves as ‘template’ for producing new RNA single strands which
are the copies of the parental viral-RNA (+sense). The newly
synthesized viral-RNAs are released from the nucleus into the
cytoplasm. They serve as messenger-RNAs (mRNAs). Each mRNA
directs the synthesis of capsomers by utilizing the ribosomes and
t-RNA of the host cell.
d. Assembly: After the production of desired capsomeres. They
are organized around the new viral RNA resulting in the formation
of complete virus particle, the virion.

e. Release: The TMV does not cause lysis of the host cell like that
of virulent bacteriophages. The host cells remain alive after the
intracellular development and assembly of viral particles. The virus
moves from one cell to the other causing systemic infection. They
may infect healthy plants on transmission from an infected plant
by some means.
1.3.4 Prions and Viroids
Prions: The word prion is a short form for ‘proteinaceous infectious
particle’; it was coined by Stanley B. Prusiner (1982). Prions are
considered as sub-viral infectious agents; discovered in a search for
the agent causing a deadly disease, scrapie in sheep. They are
misfolded proteins with about 250 amino acids. Prions transmit
their misfolded shape to normal variants of the same protein. Prions

APSCHE : SEMESTER-I BOTANY TEXT BOOK 35




     
      
 
       


Fig. 1.3.7A schematic diagram showing replication cycle of Prions


Source: https://en.wikipedia.org/wiki/Prion#/media/File:Prion_Replication.png

Know more: Diener (1989) hypothesized that viroids may represent


“living relics” from the widely assumed, ancient, and non-cellular ‘RNA
world’, that existed before the evolution of DNA or proteins. Viroids
have evolutionary significance, representing the most plausible RNAs
capable of performing crucial steps in the evolution of life from
inanimate matter i.e., abiogenesis.

Activity : Make animations on multiplication of viruses, prions and


viroids using a computer software. Draw illustrative pictures of
replication in phytophages, bacteriophages, prions and viroids.

36 APSCHE : SEMESTER-I BOTANY TEXT BOOK


can cause several fatal and transmissible neurodegenerative

tribes of Papua New Guinea) and bovine spongiform encephalopathy


(BSE) in cattle (commonly known as “mad cow disease”), were shown
to be transmitted by prions. The protein that makes prion (PrP) is
naturally present in healthy humans and animals; but the PrP in
infectious material has a different structure. Proteases cannot
degrade prions but inactivate them by modifying the structure.
Phenols, detergents and urea can also inactivate prions. Prions are
resistant to nucleases, formaldehyde, UV radiation and heat (at
80°C).
Viroids: Viroids are acellular infectious pathogens. A viroid is a
short strand of circular and self-replicating single stranded RNA.
Vioroids are variously described as degenerated viruses, primitive
viruses, escaped introns or abnormal host RNAs. Theodor Otto
Diener (1971) was the first to recognize a virion causing potato
spindle tuber disease. It is now called Potato spindle tuber viroid,
abbreviated PSTVd. Later on Citrusexocotosis viroid (CEVd) was
identified. Viroid lack protein coat and so differ from virus. Some
viroids are ribozymes with catalytic properties. They display self-
cleavage and ligation of unit size genomes from larger self-
replicating intermediates. Replication of viroids occur in 3 steps
through an RNA based mechanism in the nucleus (Pospiviroidae) or
chloroplast (Avsunviroidae) of the host plant. The replication
involves a double stranded RNA. A viroid requires the host cell
enzyme, RNA polymerase-II to catalyse ‘rolling circle’ synthesis of
new RNA using the viroid as template. Viroid infections can be
transmitted by aphids, through cross contamination after
mechanical injuries to plants during horticultural or agricultural
practices, or by leaf contact between plants. All known viroids are

APSCHE : SEMESTER-I BOTANY TEXT BOOK 37


intracellular parasites of angiosperms, and most of them cause
econom ic losses due t o t ransm it t able diseases. Chrysanthemum
stunt, cucumber pale fruit, coconut cadang disease are some
important plant diseases due to viroids. Viroids induce symptoms
in plants by RNA silencing. The viroid siRNAs comprise sequences
capable of complementary base pairing with the plant’s own
messenger RNAs; thus, induce degradation or inhibit translation
to cause the typical viroid symptoms.

Fig. 1.3.8 Diseases caused by viroids Fig. 1.3.8. Diseases caused by viroids

(a) Potato spindle tuber disease (b) Citrus exocortis disease.

Fig. 1.3.9 A schematic diagram showing replication cycle of viroids.


Source: https://en.wikipedia.org/wiki/File:Rolling_circle_replication_of_RNA_viroids.jpg

38 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Symptoms ofplant diseases causedby the
Chapter- 1.4 Viruses. Transmission of plant viruses and
their control.

Learning outcomes : At the end of the chapter you should be


able to:
1. Identify the viral diseases based on the symptoms.
2. Explain the mode of viral disease transmission and control
measures.

1.4.1General account on symptoms of plant diseases caused


by the Virus : Plant viruses cause diseases with specific symptoms
on the host and cause severe harm and loss to the crops. Plant virus
is not host-specific, but cause different types of diseases on a
different host. Based on the symptoms of different host plants by
the same virus their nomenclature and disease name have been
classified. The symptoms vary according to the factors like age,
nutrition, and environment. Sometimes a combination of two or
more viruses can cause disease in a single plant. Based on the

Types of Plant viral diseases and their symptoms


1. Chlorosis: The green parts of the plants lose chlorophyll or green
color disappeared at some places leaving the yellow spots due to
the presence of plant viruses (Fig. 1.4.1

Fig 1.4.1. Chlorosis of leaves Fig 1.4.2. Necrosis of leaves

APSCHE : SEMESTER-I BOTANY TEXT BOOK 39




     



  
      

  


 
    


    
 

    
    

FACTS: The largest class of plant virus-transmitting vectors are
insects; other vectors include mites, nematodes and chytrid fungi.

Activity: Collect the plants attacked by viruses from your


surrounding location; and record the features of the diseases,
identify them based on the symptoms.

40 APSCHE : SEMESTER-I BOTANY TEXT BOOK


2. Mosaic:
and there in the green tissue appears like a ‘mosaic” pattern and
therefore, the disease is known as mosaic disease. The virus causing
most mosaic diseases will not allow to flower or affect the dormancy
of buds. It is mechanically transmitted and usually through aphid
vectors in nature (Fig.1.4.2)
3. Yellows:
chlorophyll completely from the host tissue is known as yellows
(Fig.1.4.3)
4. Vein clearing and vein banding: The characteristic symptom
is yellowing of the entire network of veins in the leaf blade. In severe
infections, the plant becomes highly stunted due to the turning of
younger leaves to yellow and becomes entirely chlorotic. while, in
vein banding, bands of lighter or darker in color appear along the
main veins of the plant leaves. The whole leaf is yellow while keeping
the veins green (Fig.1.4.4)
5. Necrosis: The phenomenon of the appearance of brownish
necrotic spots due to the death and drying of the tissues is known
as necrosis. These spots are also known as necrotic lesions as ring
spots. They are usually characterized by the appearance of chlorotic
or necrotic rings on the leaves, fruit, and stem. These ringspot-

symptoms usually produced on the host plants by the virus are


Bunchy top, galls, hypertrophy, atrophy, rolling (leaf roll of potato)
curling (leaf curl of potato), curling of leaves, stunting (corn stunt),
and dwarfing ( barley yellow dwarf) of plants.

Know more: The plants that react promptly and characteristically


to particular viruses are called indicator plants or differential hosts.
Only susceptible plants act as indicator plants in testing but not
virus-resistant plants.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 41


Fig 1.4.4yellow vein banding on
Fig 1.4.3 Bhendi Vein Clearing
leaves of sweet cherry plants

1.4.2 Transmission of Plant Viruses and their control


Transmission of plant viruses
Mechanical Transmission : In nature plant viruses are
mechanically transmitted from diseased to other plants by rubbing
leaves together injecting plant extract, by the action of animals,
etc. Viral particles are liberated in damaged cells during rubbing
and the cells are broken. Transmission through this mechanism
occurs in such plants which are closely planted. Similarly, viral
particles attached to the surface of the animal body are transmitted
when they rub their body first on infected plants and then on healthy
plants. Viral particles enter through the injuries made by the
animals. They remain adhered to the plant surface, epidermis or
hairs. Similarly, birds also transmit viruses by this method, for
example, TMV. Some viruses survive on human clothing and
agricultural tools such as cutting knives, sickles, etc. These tools
are frequently used in agriculture and horticulture. Hence their use
first on infected plants then healthy plants facilitates viral
transmission.
Vegetative and Graft Transmission :
grafting techniques such as stem grafting, tuber grafting in potatoes,

42 APSCHE : SEMESTER-I BOTANY TEXT BOOK


root grafting, etc. This is widely used commercially for the
propagation of plants. Generally, viral particles are present almost
in all parts of systematically infected plants. Viruses will certainly
be transmitted to the progeny, if any part of such mother plants is
used for vegetative propagation through rhizomes, bulbs, corns,
tubers, cuttings, etc. Therefore, one must not use the infected
vegetative part of vegetatively propagating plants such as dahlia,

Pollen Transmission :
of virus germinate and eventually facilitate the virus to infect the
ovules of plants when they fall on the stigma of female plants. Barley
stripe virus, tobacco ringspot virus, fruit ringspot virus, bean
common mosaic virus, Datura mosaic virus are few examples of

Seed transmission of viruses is very rare but


many viruses are transmitted through seed. However, a very low
level (0.1%) of seed transmission had epidemiologically been found
out. Some examples of seed transmitted viruses are tomato ringspot
virus, bean mosaic virus, cowpea mosaic virus, tobacco ringspot
virus, cucumber mosaic virus, mung, and urd bean mosaic viruses,
etc. Seed transmitted viruses are present in the endosperm, seed
coat or embryo. After seed germination, virus-infected seedlings
are produced.
Nematode Transmission :
the roots of plants. Such nematodes also act as a vector for some
viral pathogens. Vectors are the organism that assists in the
transmission of viruses. Examples of some plant-parasitic
Longidorus, Paratrichodorus, Trichodorus, and
Ipanema.
but the transmission does not involve viral replication inside the

APSCHE : SEMESTER-I BOTANY TEXT BOOK 43


vector. There is virus vector specificity for transmission by
nematodes. Nepoviruses (polyhedral) are transmitted
Xiphenema Longidorus, Tobraviruses
transmitted by paratrichodorus and Trichodorus.
Fungal Transmission : Soil-inhabiting fungi transmit several
Olpidium brassicae, Polymyxa graminis,
and
are obligate endoparasites of higher plants. Their zoospores infect
the roots of new hosts, introduce viruses, and produce virus-specific
symptoms. These fungi acquire virus from virus-infected plants
O.
transmits tobacco necrosis virus
S. subterranea
virus.
Insect Vector Transmission Vectors are highly mobile and play
an important role in the natural ecology of viruses. In many cases,
the relationship between virus and vector is an intimate biological
association, besides mere mechanical transfer. They have slender,
needle-like parts to which they pierce the cells and suck the cell sap
of host plants. The virus is transmitted to by plants when the

Aphids: Aphids are the most notorious and important groups of


plant vectors. They are found in large numbers during the spring
and winter seasons. They show a preference for feeding strain. Most
of the vectors belong to the sucking types contained within two
families, Aphidoidea (aphids) and Cicadelidae (leaf hoppers), while
a few each belong to the Aleyrodidae (white Alles), the Cercopidae
(spittle insects), and the Coccidae (mealy bugs). A few viruses are
transmitted by both chewing and sucking insects. It is of interest
that practically all vectors of the so-called yellows group of viruses

44 APSCHE : SEMESTER-I BOTANY TEXT BOOK


are leafhoppers while practically all vectors of the mosaic group are
aphids. Aphids play a more important role in virus transmission

over 50 viruses.
Control of plant viruses: Viruses attack bacteria, algae, fungi,
herbaceous plants and trees. The diseases caused by them may
damage roots, leaves, stems, fruits, flowers which in turn cause great
economic losses by the reduction in quality and yield of plant
products. Diseases are responsible for serious losses to the crops.
They are all more important in the plantation crops which are
propagated vegetatively. These diseases because of their economic
importance and loss have received due attention and virology has
made considerable progress. As the knowledge about plant viruses
is rapidly increasing, new techniques or processes have been devised
to replace the existing method of control. However, the best way to
overcome a virus disease is through quarantine, inspection, and
certification. The seed certification and the use of virus-free clones
of important fruit and vegetatively propagated crops have received
the wider application. The use of heat and chemotherapeutic
treatments has been extended to several virus diseases of fruit and
other cash crops such peach, strawberry raspberries, and sugarcane.
The other important methods of preventing disease spread and loss
may be categorized as follows: Eradication of infected plants and
susceptible weeds, their destruction, checking possibilities of spread
of disease. Ruthless eradication by rouging and burning of rogued
plants is one of the common methods of practical value. Removal
of susceptible weeds on which the viruses overwinter or over
summer is also useful. The major advances in controlling virus
diseases have been with vegetatively propagated clonal varieties.
The best method in this connection is to select to stock used for

APSCHE : SEMESTER-I BOTANY TEXT BOOK 45


propagations free from the disease. Many methods are in vogue to
the control of virus diseases in plants. It is a difficult problem to
control viral diseases once plants are infected. Prevention or
alleviation of the effects of viruses can be done to free them from
the virus. It involves the elimination of sources of virus from
infected plants, the use of cross-protection methods, control of
vectors, and breeding for resistance.
1. Genetic Host Resistance : Genetically modified plants with
superior resistance to some viruses were developed with the advent
of recent advances in virology and plant cell molecular biology.
Resistant types should be planted whenever they are available as
different cultivars and species show different degrees of resistance

brought down by numerous cultural practices. A reservoir of virus


can be done by removal of symptomatic plants or known alternative
weed or volunteer plants and by scouting. Wash hands frequently,
use cleaner or sanitized tools, and disposable overcoat while doing
cutting, grafting, or propagating seedlings vegetatively. Growing
susceptible plants in isolation, closely spaced plants tend to escape
infection, alterations of dates of sowing or changed planting dates,
growing plants in sterilized soil helps in checking soil-borne
infection, leaving fields fallow and crop rotation are some practices.
The loss caused by the plant virus
can be avoided by the practice of geographical isolation. Before
introduction, the isolation of newly received plant material into the
rest of a production system can also minimize the unintentional
introduction of pathogens. Many cultivars of fruit and ornamental
species have been done with the clean tissue as a source of
propagation for large-scale production of virus-free plants. This can

46 APSCHE : SEMESTER-I BOTANY TEXT BOOK


be done by heat therapy that makes free individual plants or cuttings
of the virus. In this, prolonged exposure of infected tissue to
relatively high temperatures, like 20 to 30 days at 380 C makes the
virus inactivated. Another successful way to eliminate viruses,
particularly in herbaceous plants is to use meristematic tip culture
techniques. This culture is to develop virus-free callus tissue that
can then be used to generate new virus-free clones of the original
plant. It is based on the fact that the virus is usually absent in the
actively growing shoot tip of an infected plant. This procedure has
been used to clear many herbaceous cultivars of viruses.

disease control are done by soil treatment or by spraying on stems


and leaves of growing plants. Antibiotics and bases in nucleic acid-
like substances have the function of suppressing the multiplication
of plant viruses. Casein, alginic acid, etc. are polymeric materials
derived from the living body and have the function of preventing
contagious of plant viruses to other plants. Emulsion, wettable
powder or solution having a concentration of the affective
component is sprayed on plants or irrigated into soils. Bavistin
reduces virus-induced symptoms but does not affect the virus. A
chemical carbendazim is present in fungicide. Chemical treatment
in combination with heat treatment or meristem tip culture may
have an advantage in a few instances.
5. Biological Control:
management of a community of organisms and harnessing disease
suppressive microorganisms to improve plant health. The plant, the
pathogen, the biocontrol agent (antagonist), the microbial
community in and around the plant, and the physical environment
show sustained interaction for disease suppression by the use of

APSCHE : SEMESTER-I BOTANY TEXT BOOK 47


6. Management of insect vector populations:
be eliminated by direct chemical control using suitable insecticides.
In the field, it is difficult to control vectors unless coordinated on a
regional basis whereas, highly effective in closed production systems
such as interior spaces or greenhouses. Natural or synthetic
insecticides are commonly used to control insects and are derived
from nicotine (Nicotiana), pyrethrum (Chrysanthemum), rotenone
(Derris), and neem. Synthetic pesticides are sprayed or applied in
soil drench or as carbofuran or phorate granules e.g., Malathion,
Roger, Dimecron, etc.
Sometimes infection
of a susceptible plant with a mild or attenuated strain of virus helps
in protecting plants against later infection by a more severe strain
of the same virus. Protection by mild strain viruses is also helpful
in protecting crops against severe strains of the virus. Mild strains
of tomato mosaic virus are particularly helpful in controlling
infection by severe strains in tomato plants. The naturally occurring
satellite in CMV strains has been used as a biological control agent
to protect tomato plants against disease induced by severe strains
of CMV.

Activity: collect the plants attacked by viruses from your


surrounding location and record the features of disease.
Based on these, identify what is the disease.

Know more: The plants that react promptly and


characteristically to particular viruses are called indicator
plants or differential hosts. Only susceptible plants act as
indicator plant in testing but not the virus resistant plants.

48 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Chapter - 1.5 Significance of viruses in vaccine production,
bio-pesticides and as cloning vectors

Learning outcomes : At the end of the chapter you will be able


to:
1. Understand various types of viral vaccines, biopesticides
and viral vectors.
2. Evaluate the significance of viruses in medicine,
agriculture and biotechnology fields.

A. Viruses in vaccine production:


Vaccines have been designed to enhance the body immune
system by recognizing and to block the disease causing viruses.Many
viruses have been investigated for the human welfare in terms of
health, agriculture, environment and other purposes.Virus ability
will be checked in terms of expression of foreign pathogens proteins
and induction of particular immunological responses against
antigens in vivo. Stimulation of potent humoral and cellular immune
responses and delivery of antigen encoding genes to enhance
antigen presentation is possible by an effective strategy through
viral vectors. During the 18th century Edward Jenner and his
associates for the first time succeeded over an epidemic of smallpox
virus by the term of vaccination.

The success of traditional vaccines has been proved in the


numerous infectious diseases’ prevention and eradication for many
years. Some diseases require a more complex immune response, such
as HIV and malaria for which viral vectors play an important role
for the development of a successful vaccine. Initially the approaches
used to develop vaccines were empirical and dependent upon the
neutralizing antibodies induction so as to generate protection.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 49


     
 

   



  


      

     

    
  
   
 mRNA 
(COVID-19) 

         
NPV
V

  
   

       
       

50 APSCHE : SEMESTER-I BOTANY TEXT BOOK


However, this approach may not be successful for some emerging
diseases due to the need to generate immune responses in a broad
spectrum. Thus, it includes besides mucosal immunity T-cell
responses, neutralizing antibodies, which show correlation with
protection.

Viral vaccines are developed from viruses and contain either


inactivated or attenuated viruses. Inactivated or killed viral vaccines
do not cause disease and contain viruses which have lost their ability
to replicate. Majority of vaccines in use at present belong to whole
virus vaccines that would be either live or killed virus. The most
common examples of viral vaccine are BCG (BacilleCalmette Guerin)
and MMR (mumps, measles and rubella). In fact, alternative
methods for producing vaccines have been possible because of recent
advances in molecular biology.

The possibilities of production of vaccines are here given below:


Live whole virus vaccines: Live virus vaccines are capable of
inducing a protective immune response and prepared from
attenuated strains that are almost or completely devoid of
pathogenicity. e.g., Smallpox, Chickenpox.
Killed whole virus vaccines: The preparation was simply
inactivated by the destruction of replication. But, the outer virion
coat has left intact. e.g., Rabies, Flu, Polio.
Subunit vaccines: It is purified or recombinant viral antigen. From
animal tissues non replicating vaccines have been derived from
crude preparations of virus to identify the sites of peptide. Highly
purified subunit vaccines can be produced from the major antigenic
sites of viral antigens. e.g., Hepatitis B, HPV.
Toxoid vaccines: A toxin product is used which is harmful made
from the germ that causes a disease. Immunity to the parts of the

APSCHE : SEMESTER-I BOTANY TEXT BOOK 51


        
   


    
* * * * * * * * *
germ created rather than the germ itself. Thus, the toxin is targeted
for immune response instead of the whole germ. e.g., Diptheria,
Tetanus.
Anti idiotype antibodies: In experimental animals these vaccines
do mimic as foreign antigens to induce immunity against viruses,
bacteria and protozoa. e.g., Anti-Id cancer vaccines
Viral vector vaccines: There is a use of vector with modified
version of a different virus to deliver protection. COVID-19
mRNA vaccines: mRNA vaccines do not contain a live virus and
make proteins in order to trigger an immune response. They have
several benefits including shorter manufacturing times and they
won’t cause disease in the person who got vaccinated. COVID-19

Know more: Scientists are concentrating to create new types


of vaccines: DNA vaccines are easy and inexpensive to provide
strong and long-term immunity. They are plasmids usually with
a strong viral promoter enable to drive the in vivo transcription
and translation of the gene or complementary DNA of interest.
Recombinant vector vaccines act like a natural infection. They
are especially good at acquiring immune system against germs,
as virus proteins have been expressed in several microorganisms.

Learn about the recommended immunization schedule for


children (aged 0-6 Yrs.)

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B. Viruses as Bio-pesticides :
The crops are being attacked by pests or insects and involved
directly with crop loss. Chemical insecticides came into the market
to encounter this problem. But, pest control by chemical agents has
been regarded as ecologically unacceptable. Several adverse effects
of chemical insecticides were reported and created the situations
to search for an alternative to reduce the adverse effect. They are
gradually replaced by biopesticides because of increased
environmental consciousness to reduce their use in the future.
Pesticides that are derived from plants, animals, microbes,
or any other biologically available source are defined as
Biopesticides. They show potential benefits like safety for humans,
other non-target organisms and preservation of biodiversity are
just the examples. Invertebrates are being infected by viruses of a
few families and Baculoviridae family is one such classic example of
this kind of insecticides. The family Baculoviridae consists of diverse
members and virus morphology is basis of virus classification. It is
divided into two genera, Nucleopolyhedrovirus (NPV) and
Granulovirus (GV). However, only a few arthropod species have
been infected by baculoviruses. There are viruses specific for a type
of insect which are similar to bacterial insecticides. Besides this,
there are other viruses that can control insects such as sawflies and
Lepidoptera. These include such as granulosis virus, entomopox
virus, etc.Many baculoviruses are isolated from insects of
Lepidoptera, Hymenoptera and Diptera orders. They possess DNA
genome which is double stranded having the length of 80 to 200
kbp.Their specificity is usually very narrow and are safe to people
and other living organisms.
Interference of host physiology and/or introduction of an
insect-specific toxin accelerates the killing action by baculoviruses.
A baculovirus that produces occlusion bodies was constructed by

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Chang et al. with the fusion of protein consisting of
recombinant. By this experiment it has proved that a potential
biopesticide is feasible and can be possible to combine the
advantages of the virus and the bacterial toxin.
C. Viruses as cloning vectors : The plasmids are not suitable to
clone larger pieces of DNA for genomic library preparation. These
larger inserts increase the plasmid size that make them unsuitable
for transformation. A linear DNA molecule is there in the phage
which will produce two fragments following a single break. The
chimeric phages are produced by joining of the fragments with
foreign DNA. They can be isolated after allowing bacterial infection
and collecting progeny particles after a lytic cycle. However, the
insertion capacity can be optimized by the modification of phage
DNA itself according to the purpose. Several phages are described
below which are used commonly in DNA cloning.
Bacteriophage : Bacteriophages are viruses which infect bacterial
cells. Phage ë and M13 Phage are utilized in gene cloning. The
maximum packaged volume into the phage can be 53 kb DNA. If
the vector DNA is too small, it cannot be packaged properly into
the phage. Advantages of Phage Vectors:
· For cloning large inserts, they are more efficient than
plasmids.
· Phage plaques screening is much easier than identification
of recombinant bacterial colonies.
Cosmids as Vectors : Cosmid vectors are prepared by insertion of
the phage cos sequence into a small plasmid vector. The cos sites of
plasmid make the DNA to get packed in lambda particle. They allow
the packing of DNA in phage particle in vitro, permitting selection
as well as purification. They perpetuate in bacteria like plasmids,
and do not carry the genes for lytic development. They show high
efficiency to produce a complete genomic library of 106 -107 clones

54 APSCHE : SEMESTER-I BOTANY TEXT BOOK


from a mere 1pg of DNA insert. However, the disadvantage is their
inability to accept more than 40-50kbp of DNA.
Phagemids as Vectors: Like cosmid, artificially by combining the
features of phages with plasmids, Phagemid vectors are prepared.
In general, the phagemids show following characteristics: Multiple
cloning site, Inducible lac promoter, Origin of replication which is
derived from both phage and plasmid.
Vectors for Cloning Larger DNA Fragments : Vectors for cloning
large DNA fragments are often required, as human and other
mammalian genomes are very large. These vectors show various
cloning capacity. A number of such vectors have been developed that
are very useful for characterization and expression of large genes
or gene complexes.
Expression vectors in Mammalian cells: Usually, several vectors
derived from mammalian viruses have been used for the expression
of cloned genes in mammalian cells. These are as follows:
i. DNA viruses like papoviruses, e.g., SV 40 and polyoma and
adenoviruses cannot be used for expression of many large
genes because of their very low capacity for foreign DNA.
ii. Retroviruses are single stranded RNA and cannot replicate
without being integrated. It can infect large variety of cell
type and can clone large genes. But there must be high
concentration as these viruses require replicating cells for
expression or for fully differentiated cells.
iii. The nonpathogenic Adenoassociated virus overcomes from
the limitations associated with retrovirus.
iv. Herpes virus is very useful for expression of large genes as it
is non-integrating and of large size.

The pathogenicity genes must be deleted to use the viruses


as vectors. At the same time the replication associated genes were

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retained along with high copy number. A selection marker for the
vector system also needed, for example, the neomycin resistance
gene which provides resistance against the cytotoxic compound
G418. Such expression vector can be used to express cloned proteins
in cultured mammalian cells. Generally, the level of expression by
these vector systems is credited for importance of those proteins
which can be correctly folded or processed only in human cells.
Co-curricular Activities :
1. Prepare a project report of vaccines so far developed for
serious contagious diseases that cause threat to human
society
2. Enumeratethe viruses which are deadliest to human
population
3. Collect and prepare various types of viral biopesticides
avaialable for plant diseases.
4. Prepare a project report on viral diseases in commercially
important millet and other crops growing in your locality.
Self Analysis
1. Distinguish the concepts of abiogenesis and biogenesis.
2. Draw conclusions by studying various concepts and experimental
evidences on origin of life on the earth.
3. Make a brief note on diseases in plants due to obligate and
facultative microbial pathogens.
4. Illustrate the living and non-living characteristics of viruses.
5. Analyse the prebiotic acellular world by studying the features of
prions and viroids.
6. Develop a concept map on evolutionary paths among acellular
particles and cellular organisms.
7. Distinguish the symptoms caused by different viral pathogens in
plants.
8. Discuss the transmission of viruses from plant to plant in nature.
9. Enumerate the different control measures for plant viral diseases.

56 APSCHE : SEMESTER-I BOTANY TEXT BOOK


For advanced learners – Additional material about the topics

Several different types of potential vaccines for COVID-19 are in


development Inactivated or weakened virus vaccines, which use a form of
inactivated or weakened virus that won’t cause disease, but still generates an
immune response.
Protein-based vaccines use harmless fragments of proteins or protein shells
that mimic COVID-19 virus to safely generate an immune response.
Viral vector vaccines use a safe virus to generate an immune response by not
causing disease but serves as a platform to produce coronavirus proteins.
Latest and modern approach pioneering RNA and DNA vaccines, that uses
genetically engineered RNA or DNA to generate a protein that to prompt an
immune response safely.

Viruses affecting insect behavior

The behaviour of insects affected by viruses have been described for many
decades. Recent studies have shown that plant virus and insect interactions are
old and complex. Upon virus infection induction and release of volatiles from
the plant happened to attract vectors to infect plants. Once a vector is feeding
on the plant, the virus may induce anti feeding compounds to encourage the
vector to move off to a new plant. It also seems to be virus is able to manipulate
the insect to prefer uninfected plants. Viruses in a negative way can also affect
herbivorous insects. Whiteflies feeding on plants infected with Tomato spotted
wilt virus show slower growth and reduced fertility. So, it is important to know
about the complex ecology of viruses.

Viruses for Gene Therapy

Pathogen derived resistance


1. Protein based protection 2.Nucleic acid-based protection
In 1985, Sanford and Johnston developed the concept of parasite or
pathogen derived resistance. The goal of constructing genetically engineered plants
resistant to virus infection is to express a portion of the viral genome, either with or
without expression of an encoded protein that will interfere with some particular
aspect of the multiplication cycle. Successful strategies based on PDR include coat
protein mediated resistance, expression of a coding region embedded in the replicas,
the 54 kD protein of TMV, use of antisense RNAs that are the complement of the
plus or minus, sense template of the virus or use of sat RNAs that can presumably
overwhelm the viral RNA replicas and thereby suppress specific events required for
infection. Interference in the replication cycle, PDR might modulate the disease
symptoms and result in only a localized infection.

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Interactive links Online resources
https:// https://www.youtube.com/
www.pbslearningmedia.org/ watch?v=xyhZcEY5PCQ
subjects/science/life-science/ https://www.youtube.com/
evolution/origin-of-life-on-earth/ watch?v=0cwvj0XBKlE
https://www.livescience.com/ https://www.youtube.com/
53272-what-is-a-virus.html watch?v=0cwvj0XBKlE
https:// https://www.youtube.com/
courses.lumenlearning.com/ watch?v=CeVk9yC0_vk
microbiology/chapter/viroids- https://www.youtube.com/
virusoids-and-prions/ watch?v=2S7hlBmxO6s
https://www.biointeractive.org/ https://www.youtube.com/
classroom-resources/deep-history- watch?v=VQy0tJ1yu9w
life-earth https://www.youtube.com/
https://www.pbs.org/wgbh/nova/ watch?v=dvKJWcRec-4
origins/life.html https://www.youtube.com/
https://www.ncbi.nlm.nih.gov/ watch?v=zfHno163zRo
pmc/articles/PMC3718341/https:/ https://www.youtube.com/
/www.youtube.com/ watch?v=0h5Jd7sgQWY
watch?v=fR5434QR9AA https://www.youtube.com/
https://www.polyplus- watch?v=WZH380uimgc
transfection.com/blog/article/viral- https://www.youtube.com/
vectors-for-gene-therapies/ watch?v=p2iflV7EbeU
https://www.hhs.gov/ https://www.ibiology.org/plant-
immunization/basics/types/ biology/plant-virus/
index.html https://www.youtube.com/
https://www.esgct.eu/ watch?v=IDSjiN3We-k
publicpatients/gene-and-cell- https://www.ncbi.nlm.nih.gov/pmc/
therapy-glossary/gene-therapy- articles/PMC5127017/
101/vectors-101.aspx https://pubmed.ncbi.nlm.nih.gov/
http://www.biocomes.eu/biological- 9804053/
control/ https://entnemdept.ufl.edu/
https://www.livescience.com/ capinera/eny5236/pest1/content/
56598-deadliest-viruses-on- 03/3_plant_diseases.pdfhttps://
earth.html www.ncbi.nlm.nih.gov/pmc/articles/
PMC3490135/
https://www.googleadservices.com/
pagead/aclk?sa

58 APSCHE : SEMESTER-I BOTANY TEXT BOOK


https://vaccine-safety-training.org/ c. S.Miller and H.Urey
live-attenuated-vaccines.html d. Charles Darwin and
Hugo deVries
References 3. Actinomycetes and
Harold J. Morowitz (1992) dinoflagellates respectively
Beginnings of Cellular Life. Yale belong to following
University Press, USA kingdoms
Smith, J. M. and E.Szathmary (1997) a. Monera and Protista
The Major Transitions in Evolution, b. Protista and Monera
Oxford University Press, Oxford. c. Protista and Plantae
Dennis H. Bamford and M. d. Monera and Fungi
Zukerman (2021) Encyclopaedia of 4. The following kingdoms are
Virology, Elsevier Publ., Amsterdam, respectively consist of
Netherlands predominant
Roger Hull (2002) Matthews’ Plant photoautotrophs and
Virology, Elsevier Publ., Amsterdam, exclusive heterotrophs with
Netherlands varied nutrition habits.
Handbook of Plant Virology by a. Fungi and Plantae
Jawaid Khan, Jeanne Dijkstra b. Protista and Fungi
A Text Book of Virology: V. N. c. Plantae and Fungi
TiwariA Text Book of Microbiology: d. Plantae and Animalia
R.C. Dubey and D.K. Maheswari 5. Ingestion of food and
https:// holozoic nutrition are
bsppjournals.onlinelibrary.wiley.com/ characteristic to organisms
doi/10.1111/mpp.12241 in the following kingdom.
a. Plantae
Self-Assessment b. Protista
1. The proposer of biogenesis c. Animalia
theory d. Fungi
a. Rudolf Virchow 6. The following are
b. T.H.Huxley respectively regarded as
c. Aristotle Father of Microbiology and
d. J.B.S. Haldane Father of Bacteriology.
2. The concept of abiogenesis a. Robert Hooke and Louis
was conceived by Pasteur
a.Rudolf Virchow and b.Robert Hooke and
JB.S.Haldane Antonie van
b. A.I.Oparin and J.B.S. Leeuwenhoek
Haldane

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c. Louis Pasteur and 12. Structure of TMV was
Antonie van elucidated by
Leeuwenhoek a. Dmitry Iwanowski
d. Antonie van b. W.M.Stanley
Leeuwenhoek and c. Rosalind Franklin
Louis Pasteur d. Stanley B. Prusiner
7. An electron microscope was 13. The genetic material in TMV
made by is
a.Zacharias Janssen a. ssRNA with + sense
b.Galileo Galilei b. ssRNA with – sense
c.E.Ruska and M.Knoll c. dsRNA
d.Giovanni Faber d.dsDNA
8. The pathogen causing 14. The following is regarded as
plague is the Father of Virology.
a.Corynebacteriumdiphtheriae a. Martinus Willem
b.Nosemabombycis Beijerinck
c. Bacillus anthracis b. Dmitry Iwanowski
d. Yersinia pestis c. Theodor Otto Diener
9. Disease cycles of d. W.M.Stanley
tuberculosis and cholera 15. Scrapie in sheep is caused by
were worked out by a. Viroid
a. Antonie van b. Prion
Leeuwenhoek c. Virus
b. Louis Pasteur d. Virusoid
c. Robert Koch 16. Identify the self-replicating
d. Edward Jenner proteins and RNA respectively
10. Shape and symmetry of from the following
TMV is a. Prions and Viroids
a. Rod shape and helical b. Viroids and Prions
b.Rod shape and c. Virusoids and Prions
icosahedral d. Virus and Prion
c. Rod shape and binal 17. Choose the correct statement
d. Rod shape and complex related to viral vector vaccine?
11. The genetic material in a. inactivated or weakened
Gemini virus virus that doesn’t cause
a. ssRNA disease
b. ssDNA b. harmless fragments of
c. dsRNA proteins
d. dsDNA

60 APSCHE : SEMESTER-I BOTANY TEXT BOOK


c. safe virus that cannot Glossary :
cause disease Acellular: A state reflecting lack of
d. genetically engineered intact cell.
RNA to generate a Attenuated vaccine: Viable
protein pathogen created by reducing the
18. The vector formed from virulence
combining the features of Bacillus thuringiensis: Gram-
phages with plasmids. positive, soil-dwelling bacterium used
a. Plasmid as biological pesticide.
b. Cosmid Bacteriophage :
c. Phagmid infects and replicates
d. SV 40
19. Baculoviruses are isolated Baculoviridae: Group of virus infect
from following insect orders insectsbiopesticides.
a. Lepidoptera Biogenesis: A concept on origin of
b. Hymenoptera life from similar life for m by
c. Diptera reproduction.
d. All the above The production of a living
20. In expression of many large organism from other living organism
genes DNA viruses cannot be of same species.
used Biopesticides:pesticides derived
a. very low capacity for from natural materials like animals,
foreign DNA plants, bacteria, viruses and certain
b. Replication
Capsid: Protein coat that encloses the
c. Protein expression nucleic acid of a virus.
d. Correctly folded and Capsomere: A protein subunit of the
processed
21. The vaccine type used for Chemogeny: A theory of chemical
Diptheria, Tetanus is evolution predicting origin of life
a. Taxoid from inorganic matter.
Cognogeny: The diversification of
b. Killed
life forms from simple to complex
c. Live state during evolution.
d. DNA Cosmid:

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Cyanophage: Any virus that infects Protocell: A macromolecular
and replicates within Blue Green structure that is theoretically made
Algae (Cyanobacteria).
Dalton: A unit of weight
frequently used to describe the size Retrovirus: virus with RNA as its
of a virus or viral particle.
Genome: The complete set of concept on origin of life from non-life
genetic information in an organism. (inanimate matter).
Mycovirus: Any virus that infects Reverse transcriptase: An enzyme
and replicates in a fungus. catalyzing the transcription of DNA
A unit of length from RNA.
used to measure the size of a virus. Reverse transcription: Synthesis
of DNA from a RNA template.
Peplomere: A lipoprotein subunit in An RNA
the peplos of a virus. molecule suppresses the expression
Peplos: A lipoprotein coat in some of a gene by dsRNA, through
vir uses that der ives from cell translational or transcr iptional
membrane of the host. repression.
Phagemid: cloning vector with Toxoid vaccines: A vaccine response
bacter iophage and plasmid against weak and specific bacterial
toxins.
Phytophage: Plant vir us, Virion: An independent particle of a
virus.
Viroid: A naked ssRNA causing
Prebiotic: Chemical and physical infectious diseases in plants.
processes existed before the Virusoids:
emergence of life. dependent
Prion: A self-replicating protein that
causes fatal neurological disease in encapsidation.
humans or animals. Zoophage: Animal vir us,

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Special Groups of Bacteria Unit - 2
and Eubacteria

CONTENTS Learning Outcomes: At the end of


this unit learners should be able to:
1. Archaebacteria,
Actinomycetes and, 1. Develop a critical
Cyanobacteria. understanding on
2. Cell structure and characteristics of special
nutrition of bacteria and eubacteria.
Eubacteria. 2. Explain the distribution,
3. Reproduction- structure, nutrition and
Asexual and reproduction of special
bacterial bacteria and eubacteria.
recombination .
4. Economic 3. Realize the ecological and
importance of economic significance of
Bacteria special bacteria and
5. A general account eubacteria.
on symptoms of
plant diseases 4. Explain the harmful effects
caused by Bacteria; of special
Citrus canker.

Chapter -2.1

2.1.1 Archaebacteria:

Learning Outcomes: At the end of the chapter you should be able


to:
1. Understand the characteristics of special types of bacteria and
discuss the same.
2. Compare and contrast the unique features of three groups of
special bacteria in this chapter.
3. Enumerate their relationships with other prokaryotes and
eukaryotes.

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
       
    





       
       
       
  
  
 
   ds DNA    

 
  


Fig.2.1.1.a Halobacterium salinarum Fig.2.1.1. b Sulfolobus

64 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Introduction: The word Archaea (singular archaeon) has derived
from a Greek word, Archaios (= ancient). The archaebacteria are the
earliest group of prokaryotic organisms, previously classified under
bacteria. It comprises of unicellular forms with variations in
morphology and physiology; distinct from both bacteria and
eukaryotes. Carl Woese (1990) placed these organisms in a separate
domain, archaea. Thus, archaebacteria are now termed as archaea.
This module gives a brief account on archaea, their relationship with
other two domains of life, bacteria and eukaryota.

Habitat: The first discovered archaea were extremophiles, living


in hot springs, acidic, alkaline and hyper saline habitats. During
recent years, archaea are being reported from a variety of ecosystems
on the earth; representing 20% of microbial cells in the oceans. Some
archaea are thermophiles (e.g., Methanopyrus), survive at very high
temperatures (above 100°C); observed from geysers, black smokers
and oil wells. Halophilicarchaea (e.g.Halobacterium) thrive at about
20-25% salinities. Acidophilic archaea (e.g., Picrophilus) grow even
at pH 0, equivalent to 1.2 molar sulphuric acid. Natronomonas is a

3.5 M sodium chloride. Archaea are also found in cold oceanic


environments of polar seas. Though archaea constitute about 40%
of microbial mass on the earth, not even a single archaeon is isolated
and studied in pure culture.

Morphology of cells: Archaea are tiny unicellular forms, usually


0.1 to 15 µm long; they may form filaments (200 µm) or aggregates.
The shape of archaeal cells, it may be spherical, rod-shaped, slender
hair like, plate like, lobed, triangular or square shaped, or
pleomorphic. Archaea may be non-flagellated or uni-flagellated or
multi-flagellated; flagella if present, attached directly into the outer

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Fig. 2.1.1.c. Thermoplasma Fig. 2.1.1.d. Desulfurococcus mobilis

     


    
      
     

       
Taq
  



Phospholipids in Bacteria and
Phospholipids in Archaea
Eukaryotes
Glycerol – ether lipids bound by ether Glycerol – ester lipids bound by ester
bonds. bonds.
Backbone is sn-glycerol-3-phosphate.
Archaeal lipid tails have long isoprenoid The lipid tails have straight chains
chains with multiple side branches, without side branches or rings.
sometimes cyclopropane or cyclohexane
rings.
In some archaea, the lipid bilayer is The membranes always consist of lipid
replaced by a monolayer. bilayer.

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membrane of the cell. In multi-flagellated forms, usually all flagella
are attached to one side of the cell (Fig.2.1.1 a,b,c & d).

Archaea have no internal membranes. The chromosome has


a single loop of dsDNA with smaller genome (0.5 to 5.8 Mbp) as

pairs) and the smallest (4,90,885 bp) genomes are reported in


Methanosarcina acetivorans and Nanoarchaeum equitans respectively.
The genome contains only 537 protein-encoding genes.

Cell structure: Gram staining indicates that archaea are of both


positive and negative types; but structurally similar to Gram-positive
bacteria. The cell consists of a single plasma membrane and cell wall,
without periplasmic space. But in Ignicoccus, a large periplasm having

Thermoplasma Ferroplasma) have a cell


wall.The surface-layer proteins (glycoproteins) are assembled to

arrangement of protein molecules enclosing the cell like a chain mail.


It provides both chemical and physical protection, and can

Archaeal cell walls do not contain peptidoglycans (as do bacteria) or


cellulose (as do plant cells). Members of Methanobacteriales order
contain pseudopeptidoglycan in their cell walls; which resembles
eubacterial peptidoglycan in morphology, function, and physical
structure. Pseudopeptidoglycan lacks D-amino acids and muramic
acid; the latter is replaced by N-Acetyltalosaminuronic acid having
â (1-3) glycosidic bonds. Therefore,archaea are resistant to attack
by lysozyme and â-lactam antibiotics like penicillin. Methanosarcina
and Halococcus contain complex cell wall polysaccharides instead of
pseudomurein, that are similar to chondroitin sulphate of animal
connective tissue.

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Gram-negative archaea possess a layer of protein and
glycoprotein around a single or double layered cell membrane. The
chemical composition of cell wall may vary in different forms. For
example, Methanolobus and Sulfolobus contains glycoprotein in their
cell wall, while Methanomicrobium and Desulfurococcus contain
protein cell walls.
Archaeal cell membranes have distinct lipids, indicating their
distant relationship with both bacteria and eukaryotes. The
phospholipids in cell membranes of archaea are unusual in four ways,
as compared to phospholipids in bacteria and eukaryotes.

Archaea are genetically dissimilar to bacteria and eukaryotes; about


15% of the proteins encoded by any one archaeal genome are unique
to the domain. The function of these unique genes is not known.

are also found in archaea. The transfer RNA (tRNA) in archaebacteria


lack ribothymine in the common arm (TØC arm); it is replaced by
pseudouridine. The DNA replication in archaea is similar to eukaryotes,
except for the primase that synthesize a short RNA primer from the
free 3’OH group. Archaea and eukar yotes have multiple RNA
polymerases and multiple polypeptides. The RNA polymerases of

in archaea more closely resembles eukar yotic than bacterial


transcription. The ribosomes in archaea structurally similar to bacterial
ribosomes; but resembles eukar yotes in their sensitivity to
met
to initiate the translation
process as in the case of eukaryotes. Archaea use a modified form

domain Archaea, specifically in the subdivision Euryarchaeota. It


involves unique cofactors and co-enzymes like methanofuran (MP),
methanopterin (MP), coenzyme M (CoM), coenzyme F420, and

eukaryotes.

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Nutritional type Source of energy Source of carbon Example
Phototrophs Sun light Organic compounds Halobacterium
Lithotrophs Inorganic Organic compounds / Methanobacteria
compounds Carbon fixation
Organotrophs Organic Organic compounds / Sulfolobus
compounds Carbon fixation

Nutrition: The archaea resemble eubacteria in having nitrogen


fixation, denitrification, chemolithotrophy, and hyperthermophilic
growth. During those metabolic processes, electrons are transferred
from one compound to another, energy is released consequently; it

reported in any archaea. The following nutritional types are


recognized in archaea:

Reproduction: Binary fission is the usual asexual reproduction in


archaea. During this process, the DNA replicates, daughter DNAs
are pulled apart with cell enlargement. Subsequently, the cell wall
pinches off in the centre of the cell, to form two daughter cells. In
some cases, multiple fission may form many daughter cells from
one mother cell. In some archaea, sexual reproduction occurs by
conjugation, transformation and transduction as reported in
bacteria. It is reported that, the cytoplasmic bridges between cells
ofHaloferaxvolcanii transfer DNA from one cell to another in either
direction. Aggregation of cells and DNA repair was reported in
Sulfolobusacidocaldarius, subsequent on exposure to DNA-damaging
UV irradiation; which is attributed for homologous recombination.
Archaea are the target for a number of distinct DNA viruses, which
may be accountable for transduction.

Classification of Archaebacteria: Archaea areclassified into 3


major groups based on their phylogenetic relationships.

a. Crenarchaeota: Archaea of this group are tolerant to extreme


heat (as high as 110 °C) and also survive in very acidic conditions.

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Crenarchaeota
and around deep-sea vents, where water has been superheated by
magma beneath the Earth’s surface.

b. Euryarchaeota: This type of archaea lives in hypersaline


conditions and produce methane. They also live in deep sea
sediments, produce pockets of methane beneath the ocean floor.
Euryarchaeota
respiration using carbon as the electron acceptor. Carbon cycle
becomes inoperative without this group of archaea.
c. Korarchaeota: This type of archaea is rare, found in
hydrothermal environments and are the least-understood. They
share some genes in common with both Crenarchaeota and
Euryarchaeota; also possess some unique genes. Thus, thought to
be the oldest lineage of archaebacteria; perhaps the oldest enduring
organisms on the earth.
Economic importance:
extremophiles; play a vital role in various environments including
human body. Some important uses of archaea are:
a. Archaea survive

conditions. For example, the Taq polymerase obtained from


Thermus aquaticus is used in PCR at high
temperatures.
b. Methanogens are used in the treatment of sewage water and

c. Archaea are found in intestine, rectum, and vagina of


humans and domestic animals. They have various functions
which include the maintenance of the microbial flora.
d. Sulfolobus acidocaldarius

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e. Archaea are used in bioremediation of polluted sites.
f. Some archaea finding application in extraction of silver and
other metals from electronic waste.

2.1.2 Actinomycetes:
Introduction: Actinomycetes are an extensive group of
heterotrophic prokaryotes that form hyphae at some stage of their
growth; thus, denoted as filamentous prokaryotes. The word
Actinomycetes is a derivative from Greek ‘aktis’ (= a ray) and ‘mukes’
(= fungus). This group of prokaryotes has been known for over a
hundred years. Previously, considered an exotic group of organisms
with affinities to both bacteria and fungi; but classified as aerobic
(some are facultative anaerobes and a few are obligate anaerobes),
spore-forming and gram-positive bacteria. The application of new
taxonomic techniques, has led to improvements in the classification
and identification of actinomycete genera and species.
Actinomycetes are universal with varied structure, produce distinct
secondary metabolites and enzymes; thus, they are recognized as
actinobacteria, a separate group from bacteria.
Habitat: The actinomycetes are cosmopolitan microbes, widely
spread in almost all the natural ecosystems on the earth. They are
mostly soil dwellers, but have been found in a diverse range of
aquatic ecosystems, and sediments obtained from deep sea.
Actinomycetes are reported in soil of Antarctica, and even from
desert soils. It is evident that actinomycete population more surface
layer of soil and gradually decreases with increase in depth; but,
individual actinomycete strains are present in all soil layers.
Morphology: Actinomycetes are generally possess rod-like single
cells (Fig. 2.1.2) at early stage of development. Later on, many
actinomycetes produce a very delicate and profusely branched
mycelium; it may embed into the soil or solid medium, if grown in

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 
      
      
      
       


  
      
     
B
   


      
    


  
  

         


72 APSCHE : SEMESTER-I BOTANY TEXT BOOK


After a period of growth, hyphae of different types develop from
the substratum and grow aerial, constituting secondary or aerial
mycelium. The delicate mycelia often grow in all directions from a
central point, resembling rays of the sun or of a star; thus, called as
‘ray fungi’. The aerial mycelium displays various colours i.e., white,
yellow, violet, red, blue, green, or grey; and excrete pigments into
the medium. Usually, the secondary mycelium is slightly wider than
the primary mycelium. The aerial hyphae possess an extra cell wall
layer (sheath). The hyphal tip endures septation within this sheath
and gives out a chain of conidia. Conidial cell contains a deeply
staining plump, which is an oval or rod-shaped nuclear body.
The hyphae in mycelium are generally non-septate usually

certain special conditions. The mycelium in some species may break


apart to form rod- or coccoid-shaped forms. Many genera form the

Fig. 2.1.2. a. Streptomyces griseus Fig. 2.1.2.b. Actinoplanes

Fig. 2.1.2. c. Sporangium of Frankia Fig. 2.1.2. d. Mycobacterium tuberculosis

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sporangia (or spore cases) and spores on aerial hyphae, on the colony

Cell structure: The cell of an actinomycete has an envelope


comprising a plasma membrane surrounded by a cell wall. The cell
wall of actinomycetes is a rigid structure. It maintains shape of the
cell and prevents from bursting due to high osmotic pressure. It
does not contain chitin and cellulose (as do fungi). The chemical
composition of cell wall is similar to that of Gram-positive bacteria.
The cell wall is composed of a variety of complex compounds i.e.,
peptidoglycans, teichoic and teichuronic acid and polysaccharides.
The peptidoglycan consists of glycan (polysaccharides) chains;
having alternating N-acetyl-d-glucosamine (NAG), N-acetyl-d-
muramic acid (NAM) and diaminopimelic acid (DAP) molecules.
which are unique in prokaryotic cell walls. Four cell wall types are
identified in actinomycetes on the basis of three features of the
peptidoglycan composition and structure. They are (i)
diaminopimelic acid isomer on tetrapeptide side chain position 3,
(e.g., Streptomyces) (ii) the presence of glycine in interpeptide
bridges (e.g., Monospora) and (iii) sugar content of peptidoglycan
(e.g., Frankia). In cell walls of Nocardia and Saccharomonospora,
arabinose and glucose are present, but glycine is absent in their
peptide bridges; thus, considered as fourth type (Fig 2.1.3).
The protoplasm in young cells of the hyphae is
undifferentiated; but in older parts of the mycelium cells, consists
of definite granules, vacuoles and nucleoid. Each cell has a free,
naked and circular ds DNA without associated proteins, in direct
contact with cytoplasm. Actinomycetes are unique in having G+C
content above 55%. For example, the G+C content in species of
Micrococcus and Streptomyces genera is reported to be 64-78%.
Membrane bound cell organelles are absent, 70S ribosomes are
present in the cytoplasm.

74 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig.2.1.3 Mycobacterial cell wall (1) Outer lipids, (2) mycolic acid, (3)
Polysaccharides arabinogalactan), (4) peptidoglycan, (5) plasma membrane, (6)
Lipoarabinomannan Phosphatidylinositol mannoside, (8) Cell wall skeleton
https://en.wikipedia.org/wiki/Mycobacterium#/media/File:Mycobacterial_cell_wall_diagram.svg

Nutrition: The actinomycetes are popular as saprophytic soil


microflora; they are associated with rotting organic material. They
are abundant in forest soil owing to the rich organic matter
availability. The vegetative forms of the actinomycetes are quite
hardy and adapt themselves to the changing soil conditions. Some
actinomycetes are pathogens. The Actinomycetes utilize a large
number of carbohydrates as energy source; or as sole source of
carbon, if available in the media. Most of the Actinomycetes are

able to utilize nitrates and ammonia as sources of nitrogen. Majority

containing cobalt salts; and many are able to synthesize rather


complex organic molecules with antibiotic properties.
Reproduction: Most species of actinomycetes reproduce by conidia
that develop in chains from the aerial hyphae. The

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filamentsbearingconidia are often spirally twisted. The complete
aerial hypha or sometimes its upper part produces conidia.The
conidial chains may be straight, wavy or coiled . The conidial wall
may be smooth, warty, spiny, or hairy.The conidia persist in the dry
state for many years. The conidia appear as a fine powdery coat on
the surface of culture medium. The conidia when fall on the ground
and germinate under favourable conditions; they produce one to
three or even four tiny germ tubes representing myceloid condition.
The primary mycelium in some species breaks up into small
fragments, arthrospores; which seems to be bacterial cells and might
easily be misidentified as bacteria.
Classification: Actinomycetes are traditionally classified as
bacteria. In the Bergey’s Manual of Determinative Bacteriology, they
are included in several sections of volume four. All actinomycetes
are classified under the order Actinomycetales; and, are divided in
to four families i.e., Streptomycetaceae, Actinomycetaceae,
Actinoplanaceae, and Mycobacteriaceae.
Economic importance: Most of the actinomycetes are finding both
ecological and economic benefits; a few are pathogenic showing
harmful effects.
A. Beneficial effects:
a. Actinomycetesare mostly inhabitants of soil; saprophytes
degrade the organic matter in the soil and play a critical role
in the mineralization process. Streptomyces spp. convert
lignocellulose to humus thereby increases the soil fertility.
This group of microbes colonize the rhizosphere of plants
inhibit the plant pathogens.
b. Frankia sp. are nitrogen fixers, live as symbionts in the root
nodules in Casuarina spp.
c. Actinomycetes are potential producers of antimicrobial
substances.About 2/3 of the known antibiotics are from

76 APSCHE : SEMESTER-I BOTANY TEXT BOOK


actinomycetes; Streptomyces species produce antibiotics such
as streptomycin, neomycin, erythromycin etc.,
d. Cholesterol oxidase enzyme from Nocardia useful to resolve
cholesterol in blood serum.
e. Streptonigrin and retrostatin synthesised by Streptomyces
spp. inhibit reverse transcriptase.
f. Bacteriolytic or mycolytic enzymes from actinomycetes are
used in clarification of beer and wine; also serve as nontoxic
food preservatives.
g. Mesophilic and thermophilicactinomycetes are being used
in degradation of agricultural and urban wastes to make
compost.
B. Harmful effects:
1. Actinomycetes are the most common cause of infection in
dental procedures and oral Abscesses.
2. In rare cases, these bacteria (Actinomycesisraelii) can

3. Mycobacterium tuberculosis causes tuberculosis in human


beings and many mammals.

Petrichor: The word petrichor derives from two Greek words,


Petros (=rock) and ichor (=the fluid that flows in the veins of the
gods in Greek mythology). Australian researchers, Isabel Bear and
Dick Thomas (1964) coined this term to replace ‘argillaceous
odour’. Petrichor refers to the earthy scent produced when rain
falls on dry soil. This phenomenon is due to geosmin, a meta-
bolic by-product of actinobacteria mixed with oils produced by
plants during dry periods

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

 
  
        

 
 


 

Fig. 2.1.4.a. Chroococcus Fig. 2.1.4.b. Aphanocapsa

Activity: Collect the


Actinomycetes and Cyanobacteria
from your college campus or your
native place and obser ve the
structural features under
microscope.

Fig. 2.1.4.c. Oscillatoria

78 APSCHE : SEMESTER-I BOTANY TEXT BOOK


2.1.3 Cyanobacteria:

Introduction: Cyanobacteria are a vast group of photoautotrophic


prokaryotes distributed globally. The term ‘cyano’ derived from
=blue). Previously these organisms were
named Blue-Green Algae (BGA). Owing to prokaryotic and Gram-
negative staining, they are separated from algae; and are termed

photosynthetic prokaryotes; evolutionarily adapted to low oxygen


environment. Because of these distinct features they are separated
from eubacteria and treated as a special group.

Habitat : Cyanobacteria can be found in almost every terrestrial

plants (in Anthoceros and Cycas

Morphology of the thallus :

(Oscillatoria) habit.Thus, display notable variability in their


morphology. Filaments in cyanobacteria are referred as trichomes.

(Fig. 2.1.4
Cell structure : The cell structure in cyanobacteria (Fig.2.1.5)
resembles eubacteria; it consists of an envelope, cell wall, plasma
membrane, cytoplasm, and cytoplasmic inclusions. In general, most
of the cyanobacteria possess an “envelope” outside of the cell wall;
which is called a sheath, glycocalyx, or capsule. Many sheaths show

APSCHE : SEMESTER-I BOTANY TEXT BOOK 79


     
dsDNA  S
  
       
 
    




       

* * * * * * * *
a microfibrillar substructure, predominantly composed of
polysaccharides. But in some forms,>20% of the weight may consist
of polypeptides. The two-layered cell wall is Gram-negative type; it
gives shape, mechanical strength, and protection to the cell. The
outer layer of the cell wall is composed of lipopolysaccharides and
proteins with a â-barrel-shaped membrane domain. The inner cell
Oscillatoria
princeps); it consists of muramic acid, glucosamine, alanine, glutamic
acid, and 2,3-diaminopimelic acid. The cell wall has incomplete pore
pits from the cell interior, through which the cytoplasmic membrane
comes in contact with the outer layer of the cell wall; they allow the
passage of mucilage secreted by the cell. The cell wall has many
fimbriae (fili) as external appendages; flagella are not found.
The plasma membrane contains two types of lipids,
phosphoglycerolipids, and galactolipids. The typical α-helix based
membrane domain architecture is seen in proteins of the plasma
membrane.
80 APSCHE : SEMESTER-I BOTANY TEXT BOOK
The cytoplasm present inner to the plasma membrane is
divisible into the peripheral, coloured chromoplasm and the central,

photosynthetic pigments namely Chlorophyll-a, c-phycocyanin,


allophycocyanin, c-phycoerythrin, and carotenoids. The outer region
of the cell is traversed by an intracellular membrane system,
photosynthetic lamellae (thylakoids); they may be often arranged
in two or more parallel stacks, or disposed irregularly. The pigments
are dispersed freely in thylakoids. The characteristic blue-green
colour of cyanobacteria is due to copious c -phycocyanin.
Centroplasm is considered as an incipient nucleus, that lacks 2-
layered membrane and nucleolus. This region consists of a double
stranded, naked and circular DNA and RNAs; the DNA appears like
a microfibril has no histones. Some cyanobacteria contain one to
many plasmids. Membrane bound cell organelles are absent, 70S
ribosomes are present. The cytoplasmic inclusions like oil droplets,
carboxysomes (polyhedral bodies), gas vacuoles, cyanophyce-
anstarch ,volutin granules and glycogen granules are present
peripheral part of the cytoplasm.

Fig.2.1.5 Cell structure of a Cyanobacterium


https://en.wikipedia.org/wiki/Cyanobacteria#/media/File:Cyanobacterium-inline.svg

APSCHE : SEMESTER-I BOTANY TEXT BOOK 81


Activity: Learn about the composition and preparation of culture
media for Actinomycetes and Cyanobacteria. Isolate the
Actinomycetes and Cyanobacteria from your college campus or your
native place and culture them using specific media.

Specialized cells in filamentous forms : The filamentous


cyanobacteria have specialized cells like heterocysts, akinetes and
hormagonia (Fig. 2.1.6), to perform certain specified functions.

Fig.2.1.6 Heterocyst and Akinete in Anabaena


https://www.proteinspotlight.org/back_issues/023/

a. Heterocysts: Thick-walledcells meant for nitrogen fixation in


filamentous forms, unique tocyanobacteria. These are most
common in members of Nostocaceae, Oscillatoriaceae, Rivulariaceae
and Scytonemataceae. Usually, vegetative cells in trichomes enlarge
in size to form heterocysts under nitrogen deprived conditions in
the surrounding environment. But they differ from vegetative cells
in having less granular cytoplasm, additional wall layers and lack of
functional Photosystem-II. Based on the presence or absence of
these special cells, cyanobacteria can be classified as heterocystous
and non-heterocystous forms. Heterocysts may be present in the
middle of the trichome (intercalary) or at terminal position; they
may be formed as single cells or in chains. Heterocyst is connected

82 APSCHE : SEMESTER-I BOTANY TEXT BOOK


with juxtaposed vegetative cell by means of microplasmodesmata,
through a canal like polar body. Thylakoids are reticulate in
heterocysts with a few ribosomes.
b. Akinetes: These are thick-walled resting cells formed under
unfavourable conditions (nutrient deficiency and/or light
limitation), may vary in shape and size. They consist of bi-layered
wall, cyanophycean granules, glycogen, lipids and carotenoid; devoid
of polyphosphates. Normally, akinetes are present adjacent (Nostoc,
Gloeotrichia) or distant from heterocysts. These cells tolerate drying,
freezing, and long-term storage in anoxic sediments, and apparently
do not require a resting period before germination.
c. Hormogonia: A short chain of cells (5-15) with gliding motility
or planktonic with floatationof gas vacuoles is referred as a
hormagonium. The cell diameter in horamagonium is less than
those of vegetative trichomes of the respective forms. They may be
formed in different ways i.e., (i) fragmentation of trichome into
pieces (e.g., Oscillatoria), (ii) fragmentation followed by rounding
of the end cells (e.g., Nostoc), (iii) delimitation of cells into intercalary
groups (e.g., Gloeotrichia), (iv) formation of biconvex daed cells in
the middle of the filament (v) break up of weak plasmodesmata
between heterocyst and the adjacent vegetative cells. The cells in
this motile (migrating) filaments undergo cell divisions only after a
specified period od dispersal.
d. Termini: The terminal cell in some filamentous cyanobacteria
may differentiated into various shapes, it may be conical, long,
tapered, or hooked. In some forms, the terminal part of the trichome
consisting of several subterminal cells may be tapered. An extended
terminal taper of cells (multicellular hairs without thylakoids and
pigments is seen inCalothrix. The function of termini is not yet
realized.

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Movements : Almost all the cyanobacteria have creping or gliding
movement. They move in forward or backward direction, or spirally
or like a pendulum. These movements are aided by plasmodesmata
or by the contraction and expansion of the cell walls; in some cases,
the mucilage sheaths may also responsible for movements. Both
phototaxis and photophobic movements are observed in
cyanobacteria.
Nutrition : Cyanophytes differ clearly from most other
prokaryotes, they have a significantly limited range of their
nutrition. Most of them are obligate photoautotrophs. They cannot
grow in darkness even in the presence of organic substrates, except
some strains of Nostoc, Tolypothrix and Chlorogloea. Most of the
cyanobacteria utilize molecular nitrogen from the atmosphere and
fix it as inorganic forms nitrogen; this property makes them truly
autotrophic. They fix the nitrogen in free-living or symbiotic state;
as such the fixed nitrogen may be assimilated by themselves as well
as by other organisms. Heterocystous cyanobacteria such as Nostoc,
Anabaena, Aulosira, Cylindrospermum, Mastigocladus, and Tolypothrix
are important diazotrophs. Non-heterocystous forms are also
capable of nitrogen fixation during dark and anoxic conditions.
Reproduction : The cyanophytes have vegetative and asexual
methods of reproduction, do not have sexual reproduction. The
vegetative reproduction may be by cell division (e.g., Synechococcus),
fragmentation (e.g., Oscillatoria), hormogonia formation (e.g.,
Nostoc), hormospores (e.g., Westiella lanosa), planococci and
palmelloid stage. Cyanobacteria produce different asexual spores
like akinetes (e.g., Anabaena), endospores (e.g., Dermocarpa),
exospores (e.g., Chamaesiphon) and nannocyte (e.g., Microcystis).
Classification : Fritsch (1945) included cyanobacteria in algae,
under the class Myxophyceae. He classified various genera of

84 APSCHE : SEMESTER-I BOTANY TEXT BOOK


cyanobacteria into 5 orders namely, Chrococcales,
Charmosiphonales, Nostocales, Pleurocapsales, and Stigonematales.
Economic importance : Cyanobacteria have both beneficial and
harmful effects, apart from their ecological role.
A. Beneficial effects:
a. The cyanobacteria are primary producers in almost all the
ecosystems in the world, thereby key organisms for food chains
and food webs.
b. Many cyanobacteria form a thick crest on wet soils, helpful in
reclamation of land.
c. Being potential nitrogen fixing organisms, they are used as
biofertilizers. For instance, Azolla containing symbiotic Anabaena
azollae is grown in rice fields.
d. Spirulina spp. are rich in protein (60-70%) have been gained
recognition as Single Cell Protein (SCP). Central Food
Technological Research Institute (CFTRI) recommended
Spirulina for heart and obese patients.
B. Harmful effects:
a. Some members of Cyanophyceae cause damage of building
plasters, stones etc.

The water of reservoirs contaminated by Cyanobacteria,


develop a foul odour. Drinking of such unhygienic water may cause
gastric troubles in human beings.
Siderophores: Many Cyanobacteria excrete low-molecular-weight
(400–1000 kDA) compounds, siderophores having an extremely high
affinity for iron. They are meant to cope up with the iron-limited
environment. Siderophores form a strong hexadentate octahedral
complex with ferric iron (Fe3+). This complex transports inside the
cell, Fe3+

APSCHE : SEMESTER-I BOTANY TEXT BOOK 85


Chapter -2.2 Cell structure and Nutrition of Eubacteria

2.2.1. Cell structure of Eubacteria :


Introduction : The typical cell structure of a eubacterium (true
bacterium) can be revealed by the electron microphotography (Fig.
2.2.1). The eubacterial cell is prokaryotic in nature comprises of
six main components namely capsule, cell wall, cell membrane,
extracellular structures, cytoplasm and genetic material.
Capsule: Capsuleis the outermost protective layer present outside
the cell wall. It is 0.2 µm in thickness (e.g., Pneumococcus). But in
some bacteria, it is very thin (Haemophilus influenza). The capsule
is mucilaginous with 98% water and 2% of complex polysaccharides
(e.g., Enterobacter). In some bacteria, capsule is composed of
polypeptides (e.g., anthrax bacillus) or hyaluronic acid (e.g.,
Streptococcus). Capsule is a characteristic feature of gram +ve
bacteria.
In Gram +ve or soil bacteria an amorphous, loose and
undefined viscous layer present outside the cell wall called slime
layer (Leuconostoc). It is a highly charged hygroscopic layer, acts as
a barrier between cell and its surroundings. It protects the cell
against the desiccation, physical force and stress; from attack of
bacteriophages, enzymes and imparts virulence.In some bacteria,
an extracellular covering, glycocalyx is present. Glycocalyx is an
extracellular protective layer formed by the glycoproteins and
glycolipids of the bacterium (Streptococcus).
Cell wall : Bacteria possess a rigid cell wall immediately outside
the cell membrane. Cell wall is absent in Mycoplasma. The cell wall
is made up of peptidoglycans; in which peptidyl part is N-
Acetylmuramic acid (NAM) and glycan is N-Acetylglucosamine
(NAG). NAM and NAG are residues in present equal proportions

86 APSCHE : SEMESTER-I BOTANY TEXT BOOK


and arranged alternatively. Cell wall is porous and allows the passage
of substrate molecules of about 2 nm size. The cell wall provides
structural integrity to the cell. It determines the size, shape and
protects the cell from osmolytic damage. Based on the Gram staining
technique, two types of cell walls identified in bacteria; thus, bacteria
are categorized as Gram-positive and Gram-negative types.

Cell wall in Gram +ve Bacteria Cell wall in Gram-ve Bacteria


Thicker than the Gram -ve bacteria Thinner than the Gram +ve bacteria
Peptidoglycans (murein) are about 95% Peptidoglycans are 5-10%
Possess less amount of lipids Possess more lipids.
Take up crystal violet dye and Do not take up crystal violet and are
are stained purple. stained pink.
Ribitolteichoic acids and Do not contain teichoic acid.
glycerol teichoic acids are present.
E.g., Staphylococcus, Bacillus E.g., Salmonella, Escherichia

Know more: The matrix polysaccharides, teichoic acids are unique


to the cell walls of Gram +ve bacteria. Lipoteichoic acid, a major
component of Gram +ve cell walls is antigenic in function.

Cell membrane : Cell membrane is present beneath the cell wall.


It is the boundary layer of the protoplast made of phospholipids
and proteins. It is a selectively permeable membrane, It possesses
enzymes meant for electron transport and generation of proton
motive force. Phototrophic and nitrifying bacteria have membrane
infolds called mesosomes (chondroids). They are of two types i.e.,
short peripheral mesosomes and long central mesosomes. The
peripheral mesosomes possess respiratory enzymes and considered
as equivalent to mitochondria of eukaryotes. The central
mesosomes help in the replication of nucleoid and cell division.

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
  
  
           

        

       
    
  

 
        
      


Fig.2.2.1 Cell structure of a Eubacterium


https://upload.wikimedia.org/wikipedia/commons/6/61/Figure_04_02_01.jpg

88 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Gram-negative bacteria contain a second plasma membrane
superficial to their thin peptidoglycan layer called S-layer. It
comprises of phospholipids, proteins and lipopolysaccharide (LPS)
mixture.Matrix present in between inner cytoplasmic membrane
and S-layer (outermembrane) is called as periplasmic space or
periplasm.
The lipopolysaccharides in the outer cell membrane are
unique to Gram-negative bacteria and impart specific antigenic
propertie to the cells. Porins in this membrane may allow the entry
of many ions, sugars and amino acids into periplasm through passive
transport.

Know more: Sterols are absent in cell membranes of bacteria.


However, in some bacteria hopanoids are reported, they are
structurally similar to sterols and likely to serve same function.
In addition to typical saturated and un-saturated fatty acids,
bacterial cell membranes have a wide variety of fatty acids. They
include fatty acids with additional methyl, hydroxy or even cyclic
groups. The relative proportion of these fatty acids may change
to maintain optimum fluidity of the membrane (e.g., following a
temperature change in the medium).

Extra-cellular structures : Bacterial cells have flagella and


fimbriae as extracellular sructures.

a. Flagella: Flagella are the whip likeextensions of cell membrane


emerged through the cell wall is called Flagellum. Exceptcocci, all
bacteria possess flagella. Flagella are responsible for motility of
bacteria.. The flagellum in bacteria composed of thousands of
protein sub-units, flagellins. it consists a whip-like filament
connected to a motor complex by a hook-like structure. Based on
the arrangement of flagella bacteria are classified as monotrichous,
lophotrichous, amphitrichous and peritrichous types.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 89



     
 
  
   
      
s 
     
  
         
DNA
 

    

        
    
      
       
     

Know more :

energy and precursors for growth.

90 APSCHE : SEMESTER-I BOTANY TEXT BOOK


b. Fimbriae or pili: Fimbriae (singular fimbria) are extremely thin,
short, hallow, filamentous and non-flagellar appendages; they are
made up of protein subunits, pilins. Fimbriae are mostly observed
in Gram-negative bacilli (e.g., Salmonella typhi) and also cocci (e.g.,
Neisseria gonorrhoeae. Fimbriae help in attachment of bacteria to a
surface or to host cells. In a few bacteria like Myxococcus, fimbriae
may facilitate motility and assembly into multicellular structures.
Pili are structurally similar to fimbriae, but are longer and present
in low numbers per a cell. Pili are antigenic in nature, classified as 3
types namely, common pili, sex pili (F or fertility pili) and col I
(colicin) pili. Common pili help in attachment of bacteria to host
cells. The sex pili are meant for transfer of genetic material during
conjugation. The col I pili produce colicin or hemolycin, that causes
lysis of red blood cells of host to release haemoglobin.

Cytoplasm: The cytoplasm present inner to the cell membrane. It


does not contain any membrane bound cell organelles like
mitochondria, Golgi complex, ER etc. However, spindle shaped,
membrane bound gas vesicles meant for buoyancy are seen in
planktonic bacteria. Ribosomes, some cell inclusions, and nucleoid
are the apparent structures in cytoplasm of bacteria. In bacteria,
70S types of ribosomes are present as chains, polysomes. Each

cellular inclusions include non-living material like glycogen, lipid


droplets, crystals, volutin granules (poly-metaphosphate), poly-β-
hydroxybutyrate (PHB) and pigments. Sulphur granules are common
2

cytoplasm of hydrogen oxidizing (Knall gas bacteria) and nitrogen


oxidizing bacteria (Nitrosobacteria and Nitrobacteria) proteinaceous

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structures, carboxysomes are present as microcompartments. They
contain the enzymes for carbon fixation, especially RuBisCo and
carbonic anhydrase. Membrane associated vesicles, chromatophores
are present in photosynthetic bacteria (e.g., Rhodospirillum rubrum),
and they contain bacterial chlorophylls and carotenoids.

Genetic material: The genetic material in bacteria appear as


nucleoid and plasmids.

Living magnets: Magnetotactic bacteria (Magnetospirillum


magneticum, Magnetobacterium bavaricum) contain micro
compartments called magnetosomes. They demonstrate
magnetotaxis i.e., sense and align themselves along a magnetic
field just like a compass needle. Magnetosomes are composed of
the mineral magnetite or greigite, and are surrounded by a lipid
bilayer membrane.

a. Nucleoid: It appears as a low electron dense central area as


compared to rest of the cytoplasm in bacteria. It is devoid of nuclear
membrane, nucleolus and nuclear sap, often called as genophore or
bacterial chromosome. The genophore is a double stranded circular
DNA (rarely linear as in Borrelia burgdorferi) without histone
proteins; thus, present as a naked structure, in direct contact with
cytoplasm. In Escherechia coli, the genophore has 4,600 base pairs.

b. Plasmids: In most of the bacteria a number of small circular


DNA (plasmids) are located outside the nucleoid region, in
cytoplasm. Plasmids can be easily gained or lost by a bacterium and
can be transferred between bacteria as a form of horizontal gene
transfer. Therefore, plasmids are called as extrachromosomal DNA.
Plasmids are autonomously replicating units contain about
100genes. They confer antibiotic resistance, pathogenicity,
degradative property, fertility, secretion of enzymes etc.,

92 APSCHE : SEMESTER-I BOTANY TEXT BOOK


2.2.2. Nutrition of Eubacteria:

precursors for growth.

Introduction: Nutrition refers to a process by which living beings


obtain and utilize the inorganic and/or organic material for their
survival, growth and reproduction. Like all living organisms,
bacteria require certain nutrients like carbon, nitrogen, sulphur,
phosphorous, oxygen etc., for their biological processes. Bacteria
obtain carbon and nitrogen from various inorganic and organic
nutrients from the environment. Bacteria can be categorized in to
two major groups based on their nutrition rather energy sources,
they are autotrophs and heterotrophs.
I. Autotrophs: Certain bacteria synthesize food for their growth,
by obtaining the carbon via atmospheric CO2 and hydrogen from
H2, H2S and NH3. The autotrophic bacteria can be classified into
two groups based on the source of energy i.e., photoautotrophs and
chemoautotrophs.
a. Photoautotrophs: Photoautotrophic bacteria contain pigments.
Purple sulphur bacteria (e.g., Chromatium glycolicum) contain
bacterial Chlorophyll a, bacterial Chl.b, and carotenoids. Green
sulphur bacteria (e.g., Chlorobium vibrioforme) contain chrlorobium
chlorophyll. They do not utilize water, perform non-oxygenic
photosynthesis and hence do not release O2. Derive energy from
light. They synthesize their food by obtaining hydrogen from
sulphites and thiosulphates and CO2 from the atmosphere.

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b.Chemoautotrophs:Chemoautotrophic bacteria
(chemolithotrophs) are non-photosynthetic and cannot trap light.
This type of bacteria obtain energy by the oxidation of inorganic
molecules in their environment. Chemolithotrophs oxidize
hydrogen sulphide, elemental sulphur, ferrous iron, molecular
hydrogen and ammonia. The energy released in those exergonic
reactions is utilized for synthesis of all necessary organic compounds
using CO2 as carbon source. These bacteria help in recycling of many
inorganic minerals.
Category Example Inorganic Product Energy
energy Source released
Sulphur Beggiatoa Hydrogen sulphide Sulphur 50 K.Cal.
bacteria
Iron bacteria Ferrobacillus Ferrous carbonate Ferric hydroxide 81 K.Cal.
Nitrifying Nitrosomonas Ammonia Nitrites 78.5 K.Cal.
bacteria
Hydrogen Hydrogenomonas Hydrogen Water 56 K.Cal.
bacteria

II. Heterotrophs : Majority of bacteria are heterotrophic, derive


their energy from organic compounds. Such bacteria are
cosmopolitan, reported from ever y ecosystem containing
decomposing organic matter. Some of them live as
photoheterotrophs; many of them are parasites or symbionts in
plants and animals.

a. Photoheterotrophs: This type of bacteria use light for energy


and organic compounds as carbon source. They can utilize
carbohydrates, fatty acids, alcohols etc., in their environment to get
the carbon. Examples: Purple non-sulphur bacteria (Rhodobacter,
Rhodospirillum) green non-sulphur bacteria (Chlorof lexia,
Thermomicrobia) and heliobacteria ( Heliorestis).

94 APSCHE : SEMESTER-I BOTANY TEXT BOOK


b. Chemoheterotrophs: Chemoheterotrophic bacteria cannot
synthesize organic material for their metabolism. Some bacteria of
this group are saprophytes, while others depend on living plants
and animals as parasites or symbionts.

i. Saprophytes: Saprophytic bacteria live on dead remains of


other living beings, or decaying organic matter. This group
of bacteria (Bacillus spp., Vibrio japonicus) are mostly soil
dwellers, regarded as the natural scavengers. They are the
key players in mineralization and also recycling of minerals;
degrade complex polysaccharides like lignin, cellulose,
hemicellulose in wood and other plant material. Saprophytic
bacteria produce extracellular enzymes to breakdown the
complex organic compounds into simple molecules; in due
course obtain energy and absorb digested material in soluble
form. Saprophytes degrade carbohydrates and proteins
respectively by fermentation and putrefaction. Lactic acid
bacteria convert the sugar in milk to lactic acid through
glucose, by the fermentation process. The putrefying
bacteria digest the proteins into peptones, polypeptides,
peptides, and finally amino acids.

ii. Parasites: Parasitic bacteria obtain their nutrients from


living hosts. Most of them are pathogenic causing harmful
effects to the host plants (e.g., Xanthomonas, Erwenia,
pseudomonas) or animals (e.g., Bacillus anthracis) or human-
beings (e.g., Diplococcus pneumoniae).

iii. Symbionts: Some bacteria live symbiotically in plants (e.g.,


Rhizobium) and human beings (e.g., Escherichia coli). Both
the host and the bacterium get benefit out of this

APSCHE : SEMESTER-I BOTANY TEXT BOOK 95


Purity of water in the Ganga River: Despite various biological
pollutants, the water in river Ganges is pure. It was previously
ascribed to bacteriophages. But now a days it is realized that
Bdellovibrio bacteriovorous living in Ganga water keep it pure by
attacking other bacteria. This is a Gram-negative and obligate
bacterium. It lives as parasite in periplasmic space of other Gram-
negative bacteria in the form of a structure, bdelloplast; that
modifies the cells of both parasite and host.

Activity: Take a sample of non-pathogenic bacteria, stain them


with Gram staining and report the type of bacteria based on
pigment retained.

Activity: Take the sample of Lactobacillus from curd and observe


the movements using hanging drop method.

96 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Reproduction in Bacteria - Asexual and bacterial
Chapter - 2.3
recombination

Learning outcomes : At the end of the chapter you should be


able to:
1. Understand the mode of reproduction of prokaryotic cell
2. Understand the impact of environment on structure of
asexual spores.
3. Gain the knowledge of different types DNA recombination
to develop news trains.

2.3. Reproduction in Bacteria :


Introduction: Bacterial reproduction is mainly depends on nutrient
availability in the surroundings. Based on the availability of
nutrients, bacteria show two types of reproduction i.e., Asexual
reproduction (Binary fission and Endospores) and Bacterial
recombination.
A. Asexual reproduction:
a. Binary Fission : During binary fission,a pairof daughter cells is
formed from the parent cell. Daughter cells are genetically identical
individuals. Binary fission, is simple and rapid process. The single,
circular DNA chromosome of bacteria occupies a specific location,
the nucleoid, within the cell without histone proteins. The bacterial
chromosome is attached to the plasma membrane at about the
midpoint of the cell. DNA undergoes bidirectional replication and
form two strands. The newly formed double strands moves away
from the cell wall attachment toward the opposite ends of the cell.
As the cell elongates, the growing membrane directs the transport
of the chromosomes. After the chromosomes have cleared at the
midpoint of the elongated cell, cytoplasmic separation begins. A
septum is formed between the nucleoids, extending gradually from
the periphery toward the center of the cell into two daughter cells
with distinct cell walls (Fig.2.3.1).

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
  

      


        
 
                          
     

 
{ ²³ y Ž
T (1946) ¿±+E ¹>wŸ HŽ¿£ T. <‘“ ý ËFF
q T>=H •s Á
F+F
F-

Fig.2.3.1 Stages of binary fission of E.Coli


1.The bacterium with tightly coiled DNA.
2: The DNA of the bacterium has replicated .
3: The DNA is pulled to the separate poles of the bacterium.
4: The new cell wall fully develops inwards resulting in the completesplit of the bacterium.
5: The new daughter cells have tightly coiled DNA, ribosomes, and plasmids.

98 APSCHE : SEMESTER-I BOTANY TEXT BOOK


b. Endospore : Endospore is special resistant, dormant structure
develop within vegetative bacterial cells of Gram+ve bacteria,
Bacillus and Clostridium (rods), Sporosarcina (cocci), and others.
Endospores are viable for around 100,000 years, because they are
highly resistant to environmental stresses such as heat, ultraviolet
radiation, gamma radiation, chemical disinfectants, and desiccation
and hence, they are highly pathogenic in nature. Endospores are of
great practical importance in food, industrial, and medical
microbiology.
Position of the endospore in the mother cell or sporangium
frequently differs among species. Spores are either centrally placed,
close to one end (sub-terminal), or definitely terminal and the
position of endospore is significant in the identification of bacteria.
Electron microscopic structure of endospore is complex (Fig. 2.3.2).

Source:
clip_image002-129.jpg (272×517)
(biologydiscussion.com

Fig2.3. 2. Structure of endospore and endospore formation CW:corewall, CX:cortex,


SC: sporecoat, EX:exosporium N; Nucleoidand CR: bacterial ribosome

APSCHE : SEMESTER-I BOTANY TEXT BOOK 99


F
F 
FF
HfrDNA
F
F
   F    
      
 Hfr  F    
FF
F



   S
DNA
S
  




Know more: Formation of endospore is suited for research on


the construction of complex biological structures because their
intricate entities of bacteria are developed in a very organized
fashion over a period of few hours.

100 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Structure of endospore: The spore is surrounded by three layers;
exosporium, protein coat and cortex. Exosporium is thin and
delicate outer layerfollowed by thick and multi layered protenaceous
and middle spore coat and innermost cortex made up of
peptidoglycan. Spore-coat is impermeable and resistance to
chemicals whereas the cortex occupy half of the spore volume. The
central part of endospore is called core or protoplast which is
metabolically inactive and has ribosomes, a nucleoid, DNA repair
enzymes and dipicolinic acid (DPA) complexed with calcium.
DPA weighs more than 15% of the spore’s dry weight. Between the
core and the cortex, the spore cell wall is present. The cortex
protect the spore from both heat and radiation damage by removing
water from the protoplast osmotically. Spore coat protects the spore
against digestive enzymes and strong chemicals such as hydrogen
peroxide. DNA is repaired during germination and out growth after
the core has become active once again. Sporogenesis or sporulation
(spore formation), normally commences when growth ceases due
to lack of nutrients. In Bacillus megaterium, duration of sporulation
is 10 hrs. The transformation of dormant spores into active
vegetative cells is a complex process like sporogenesis. It occurs in
three stages: (1) Activation, (2) Germination and (3) Outgrowth.

B. Genetic recombination in bacteria : Bacterial cells do not


possess sex chromosomes or sex organs like eukaryotic cells.
However Bacteria develop new strains by genetic recombination
either between two cells or in a single in three ways (a) Bacterial
conjugation, (b) DNA transformation and (c) DNA transduction.
(a). Bacterial conjugation : The transfer of genetic information
by direct cell to cell contact between two cells in the conjugation
(Lederberg and Edward L. Tatum,1946). Bernard Davis (1950)
evidenced that direct contact was required for the recombination
during conjugation, by U tube experiment.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 101


(i) F+ x F - mating : In this type of conjugation, the donor is (F+)
and the recipient is (F-) strains, and the gene is transferred from
donor to recipient in a nonreciprocal and polar method (Lederberg
and Tatum). Hence it is called F+X F- mating. In this conjugation
only F- strains frequently became F+. An extra chromosomal F factor
carrying the genes for pilus formation and plasmid transfer is
present in the F+ strain. During F+X F- Mating or conjugation, the
F factor replicates by the rolling-circle mechanism and form two
copies. One of the copies moves to the recipient cell (Fig. 2.3.3. a)
called the entering strand. The entering strand is copied to produce
double-stranded DNA. The donor and recipient cells are joined by
the sex pilus or F pilus and form a connecting bridge as a channel
for DNA transfer. The channel is either the hollow F pilus or a special
conjugation bridge formed between the two cells (E. coli).

(ii) Hfr conjugation : In this type of mating, the HFr cell is the
donor, and the F- cell is the recipient. In HFr cell the F factor is
present as an episome and it is integrated into the bacterial
chromosome. Hfr strain has a high frequency of recombination.
Chromosomal gene transfer is also more efficient than F+ cells.
Hence HFr strains transfer bacterial genes with great efficiency and
do not usually change recipient bacteria to donors. Integrated F
factor can direct the synthesis of pili. HFR strain replicates the DNA
by the rolling-circle model (Fig. 2.3.3.b). After the replication, the
chromosome moves through the pilus or conjugation bridge. Gene
transfer can be in either a clockwise or counter-clockwise direction
around the circular chromosome, depending on the orientation of
the integrated F factor. After the replicated donor chromosome
enters the recipient cell, it may be degraded or incorporated into
the F- genome by recombination.

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Fig.2.3.3. a. Bacterial conjugation- F+x F- mating

Fig. 2.3.3.(b) HFRXF- mating


https://en.wikipedia.org/wiki/Hfr_cell#/media/File:Hfr_Recombination.png

Know more: Davis Experiment:U tube consisting of two pieces


of curved glass tubing fused at the base to form a U shape with
a fritted glass filter between the halves with two different
auxotrophic strain of E. coli and discovered that when the two
auxotrophic strains were separated from each other by the fine
filter, gene transfer could not take place.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 103


b. DNA transformation: The uptake of a naked DNA molecule or
fragment by a bacterial cell from the medium and incorporate into
the recipient bacterial cell in a heritable form is called
Transformation (Fred Griffith, 1928). DNA fragments of a lysed
bacterial cell are released into the surrounding environment and
enter into the competent bacterial cell resulting in the
transformation of the competent cell into a new strain with gene
recombination. A bacterial cell that can be able to take up DNA and
be transformed is called the competent cell. If a fragment contacts
a competent cell, it can be bound to the cell and taken inside
(Fig.2.3.4). Mechanism of transformation has been intensively
studied in S.pneumonia (Fig.2.3.5). A competent cell binds a double-
stranded DNA fragment, one of the strands is hydrolyzed by an
envelope-associated exonuclease and the other strand moves
through the plasma membrane with the help of associated
membrane proteins. The single-stranded fragment can then align
with a homologous region of the genome of the competent cell and
be integrated.
c. DNA transduction : Transduction is the transfer of bacterial
genes between two bacterial strains by viruses. Bacterial viruses or
bacteriophages participate in this type of recombination. These
viruses have relatively simple structures in which virus genetic
material is enclosed within an outer protein coat. The coat protects
the genome and transmits it between host bacterial cells. After
infecting the host cell, a bacteriophage (phage for short) often takes
control and forces the host to make many copies of the virus.
Eventually, the host bacterium bursts or lyses and releases new
phages with bacterial genes. The virus-containing these bacterial
genes then inject into another bacterium thus completing the
transfer. There are two different kinds of transduction i.e., (i)
Generalized and (ii) Specialized.

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Fig.2. 3.4 Mechanism of bacterial transformation

Fig. 2.3.5 DNA transformation with a fragment and a plasmid

i) Generalized transduction: It occurs during the lytic cycle of


virulent and temperate phages and transfers any part of the
bacterial genome. Virus particle is simply a carrier of genetic
information from the original bacterium to another cell. The virus
injects the DNA into another bacterial cell but does not initiate a
lytic cycle (Fig. 2.3.6). This phage is known as a generalized
transducing particle or phage. Once the DNA has been injected,
it will be incorporated into the recipient cell’s chromosome as
transferred genes. Generalized transduction was discovered by
Lederberg and Zinder.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 105


Fig 2.3.6. a. Generalized transduction through lytic cycle

ii) Specialized transduction : Specialized transduction is made


possible in the lysogenic life cycle of a virus. The best-studied
example of specialized transduction is the lambda phage. The
lambda genome inserts into the host chromosome at specific
locations known as attachment or att sites. The att sites of phage
and bacterial are similar and can form a complex with each other.
Consequently, specialized transducing lambda phages carry these
bacterial genes and this part of the virus is called a prophage. When
the cell wall of the host shows improper excision prophage is

106 APSCHE : SEMESTER-I BOTANY TEXT BOOK


induced to leave the host chromosome. As a result phage genome
contains portions of the bacterial chromosome that will transfer to
a new bacterial cell (Fig. 2.3.6).

Fig 2.3.6. b. Specialized transduction through lysogenic cycle

APSCHE : SEMESTER-I BOTANY TEXT BOOK 107


Chapter-2.4 Economic Importance of Bacteria

Learning outcomes: At the end of the chapter you should be


able to:
1. Understand and explain the uses of bacteria in agriculture
and industry.
2. Discuss the ill-effects of pathogenic bacteria.

Bacteria are ubiquitous with greater diversity in the mode


of nutrition. The characteristic features of bacteria in terms of cell
wall structure, metabolic regulation, and genetic material exchange
pattern accomplished its presence both in the beneficiary and
harmful activities. This chapter deals with beneficiary activities of
bacteria with a brief emphasis on agriculture and industrial
application.
A) Beneficial Activities of Bacteria: Bacteria are absolutely
essential to the presence of life. They play a major role in agriculture
and industrial production apart from food beverages vitamins
vaccines etc.
a) Role in Agriculture:
i)Decay and decomposition: Saprophytic soil bacteria are
generally saprophytic and play an important role in the
decomposition of organic matter. These bacteria help to decompose
dead and decaying organic matter by removing the harmful dead
matter as scavengers by ammonification and supply the decomposed
matter to soil by the process of mineralization. The dead bodies
and wastes of organisms (both plants and animals) are decomposed
by the activities of the soil saprophytic bacteria (Bacillus mycoides,
Bacillus ramosus etc.,).
Variety of elements of minerals such as carbon, oxygen,
hydrogen, sulfur, and phosphorus which are present in dead and

108 APSCHE : SEMESTER-I BOTANY TEXT BOOK


decaying organic matter are reduced to simple compounds such as
carbon monoxide, water, nitrates, sulphates and phosphates. Some
of these go back to the soil and the rest to the air. From the soil,
they can be absorbed as plant food.
ii) Soil fertility: Chemolithotrophic soil bacteria commonly
Nitrogen bacteria play an important role in maintaining soil fertility.
Nitrogen bacteria act as “Nature’s farmers. Nitrogen-fixing bacteria,
Ammonifying bacteria, and nitrifying bacteria enhance soil fertility
whereas denitrifying bacteria play a reverse role and deplete the
nitrogen fertility of the soil. Green plants generally absorb nitrogen
s nitrates and to some extent as ammonia from the soil by a series
of steps.
iii) Nitrogen-fixing Bacteria: Nitrogen-fixing bacteria are able
to make use of the atmospheric nitrogen and change it into
nitrogenous compounds This process of nitrogen transformation
is called nitrogen fixation. These bacteria are of two types i.e., free-
living nitrogen fixers and symbiotic nitrogen fixers. Azotobacter,
Beijerinckia (aerobic forms) and Clostridium (anaerobic) live free in
the soil. They take gaseous nitrogen from the air present between
the soil particles. The nitrogen combines with other elements
forming organic nitrogenous compounds. These compounds are
assimilated by the bacteria (Fig.2.4.1). Rhizobium leguminosarum
(syn.Bacillus radicicola) is a symbiotic nitrogen-fixing bacterium that
lives in the roots of Leguminosae members such as Bean, Medicago,
and others in the form of nodules. Besides the legumes, the nodules
are found on the roots of Alnus glutinosa, Casuarina, species of
Coriaria, and a few others (non-leguminous plants). The presence
of bacteria in the roots causes the formation of little nodules). In
Pavetta indica, the bacterial nodules are formed on the leaves.
Neither Rhizobium nor the legume root alone can fix nitrogen !

APSCHE : SEMESTER-I BOTANY TEXT BOOK 109




    


     
   

       
 
     

            

   


  

   





 
Know more: Principle of decomposition and mineralization of
saprophytic bacteria is extensively applied in sewage disposal
system of cities and urban septic tanks

110 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Know more: Rhizobium lives in the soil in the form of form of coccus.
The legume roots secrete substances which attract bacteria on to
their surface. The bacteria, in turn, secrete a growth hormone which
causes the root hairs to curl. The cocci enter the curled root hairs.
They grow in the root hair in the form of a continuous thread-like
mass which finally reaches the root cortex. The cortical cells become
filled with a dense mass of bacteroids and stimulate abnormal
growth. The cortical cells around the infection divide and redivide
and grow to form a nodule. A nodule comprises a central mass of
cells full of bacteroids. Around this zone of infection are a few cell
layers thick of bacterial free cortical zone. At the apex of the nodule
is the meristematic region and a vascular strand at its base Within
the host cells the cocci feed on the carbohydrates and other foods
and undergo a change in their form to V, T or Y shaped. The core of
the nodule is red owing to the formation of red respiratory pigment,
leghaemoglobin (Fig. 2.4.2).

Ammonifying Bacteria: The saprophytic soil bacteria break down


the proteins and other nitrogen-containing waste of the plant and
animal origin to amino acids by a process called Ammonification.
The amino acids are then converted into ammonia by a group of
bacteria called the ammonifyingbacteria (Bacillus ramosus,
B.mycoides). Common cereals can make use of ammonium
compounds as a source of nitrogen. The majority of plants, however,
cannot absorb ammonium compounds as a source of nitrogen
Nitrifying Bacteria: Ammonia in the soil is oxidized by
chemosynthetic autotrophic soil bacteria called nitrifying bacteria
(Nitrosomonas and Nitrobacter). They form nitrates and nitrites from
ammonium compounds in two steps:
i. Nitrosomonas oxidizes ammonium carbonate to nitrous acid
liberating energy.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 111


Fig 2.4.1 Nitrogen Fixing Bacteria :
a) Free Nitrogen fixers and b) Symbiotic Nitrogen fixers.
Source: https://www.uou.ac.in/lecturenotes/science/MSCBOT



       
 
 
 


Fig.2.4.2 Nitrogen fixing


bacteria:
A. Symbiotic bacteria in root
nodules of legume
B. Cross section of nodule and
C. Infected cortical cells with
bacteroids of Rhizobium sp.

Source: https://www.uou.ac.in/
lecturenotes/science/MSCBOT
http://go.microsoft.com/fwlink/p/
?LinkId=255141

112 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fact: Neither the ammonifying nor the nitrifying bacteria add to
the total quantity of combined nitrogen in the soil. The ammonifying
bacteria convert amino acids into ammonia. The process is called
ammonification.The nitrifying bacteria convert nitrogen from the
unavailable form of ammonium salts to the available nitrates. This
process converting unavailable ammonium salts into available
nitrates is called nitrification.

ii. The nitrous acid then combines with bases in the soil-forming
potassium nitrite. Nitrobacter oxidizes nitrites to nitrates again
liberating energy.
b. Role of Bacteria in industries: Bacteria are being exploited
by researchers for various industrial processes.
i) Dairy industry: The butter and cheese industries depend on
Lactic Acid Bacteria. The souring and curding of milk by lactic acid
bacteria is the common example of application in everyday life.
Streptococcus lactis, S. thermophilic, Lactococcus lactis, Lactobacillus
plantarum, Lactobacillus casei, Lactobacillus acidophilus, Lactobacillus
helveticus, Lactobacillus bulgaricus, etc., are used to produce butter,
cheese, curd, etc.
ii) Vinegar Industry: Vinegar (acetic acid) is oxidation product
of alcohol produced by acetic acid bacteria.These are Gram-negative
bacteria belong to the Acetobacteriaceae. In oxidative fermentation,
vinegar is produced as a by-product.
e.g., Acetobacter aceti, A.cerevisiae, A.malorum, A.oeni, A.pasteurianus,
A.pomorum, Gluconacetobacter entanii, G.liquefaciens, G.oxydans,
Komagataeibacter europaeus, K. hansenii, K. intermedius, K.
medellinensis, K. oboediens, K.xylinus.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 113


iii) Curing of tea, tobacco and indigo: Bacillus megatherium and
Micrococcus candisans are used in the curing and fermentation of
tea and tobacco leaves for commercial purposes.
iv) Leather, linen and sponges: Leather making, production of
linen and preparing sponges also use bacteria.The tough fibres,
which are left behind, are separated. These fibres are spun and
woven into linen cloth, ropes, etc.E.g., Achromobacter parvulus,
Aerobacter cloacae, Aerobacter aerogenes, Bacillus brevis, Bacillus cereus,
Bacillus megaterium, Bacillus sphaericus, Bacillus subtilis, Clostridium
butylicum, Clostridium beijerinckii, Clostridium saprogenes, Clostridium
perenne etc.

v) Coffee and cocoa: Coffee and cocoa beans are white in colour
and quite bitter in taste. The bacteria digest the bitter coverings of
seeds and give the characteristic colour, flavour and aroma.

Know more: Process of curding takes place in two steps. In the


first step, the lactose sugar of milk is fermented into glucose by
anenzyme lactose secreted by the lactic acid bacteria. In the
second step, there is transformation of glucose into lactic acid.
The latter sours the milk and coagulates the milk proteins
(casein) forming curd and whey.

114 APSCHE : SEMESTER-I BOTANY TEXT BOOK


A general account on symptoms of plant diseases
Chapter -2.5
caused by Bacteria; Citrus canker

Learning outcomes: At the end of the chapter you should be


able to:
1. Identify the symptoms of bacterial diseases in plants.
2. Explain the disease cycle of Citrus canker.

2.5.1 General account on symptoms of plant diseases caused


by Bacteria : Diseases in plants, animals and humans are caused
by heterotrophic bacteria showing parasitism. Such diseases are
called bacterial diseases. Plant diseases caused by bacteria were first
revealed by T.J. Burrill (1878). Phyto-pathogenic bacteria are
generally rod-like, non-spore forming and flagellate and either
aerobic (live in the presence of oxygen) or facultative anaerobes
(grow with or without oxygen). There are about 200 species of
phyto-pathogenic bacteria are reported and almost all of them are
parasites within the host plant or on the surface in plant debris. In
absence host, these bacteria exist in the soil as saprophytes. Parasitic
bacteria show dessimination by several ways. Some bacteria have
the ability to move short distances in water by flagella. Most of the
bacteria are disseminated by passive agents such as air, insects, water
and soil. Some parasitic bacteria are without flagella and these
bacteria gain entry into the host through wounds, stomata,
lenticels, hydathodes or through the thin epidermis.

A) Types of Bacterial Diseases : Bacterial symptoms are similar


to the symptoms in fungal plant disease. Based on symptoms,
bacterial diseases are broadly classified into Bacterial spots, blights,
wilts, scabs, cankers and soft rots of root.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 115


a. Bacterial spots: The most common symptom of bacterial disease
is leaf spots. Spots appear on leaves, blossoms, fruits and stems. A
bacterial spot which appear and advance rapidly it is a blight
(Fig.2.5.1.a).Spots on leaves have a rotten or fishy odour, watery
and are initially confined between the leaf veins and will appear
angular,in dicots.Bacterial ooze from the spot is the diagnostic
symptom for bacterial infections. Sometimes a chlorotic halo will
surround the bacterial lesion of an infected leaf. Spots may coalesce
causing large areas of necrotic tissue. Bacterial spots will appear as
streaks or stripes on monocotyledonous plants. Bacterial leaf spots
and blights are caused by the genera Pseudomonas and Xanthomonas.

b. Cankers: Canker symptoms appear on trunks, stems, twigs and


branches. The most conspicuous symptom of a bacterial canker
disease is the development of cankers and gum exudation
(gummosis). Cankers can be slightly sunken, dark brown and much
longer than broad. The cortical tissue of the canker is orange brown
to dark brown in colour (Fig.2.5.1.b). Gum is produced in most of
the cankers especially on branches and twigs. Cankers that do not
produce gum may have a sour odor, soft, sunken and moist. Cankers
that girdle trunks and branches can cause dieback of the portion of
the tree distal to the canker. Pseudomonas and Xanthomonas cause
canker disease of stone fruit and pome fruit trees and citrus.

c. Bacterial Galls: Gall tissue is composed of disorganized,


randomly proliferating cells that multiply in the intercellular
(between the cells) spaces in the vicinity of the wound. A wound in
the host is required for the pathogen to gain entry into the host
tissue (Fig.2.5.1.C). Entry of pathogen induce rapid and continuous
cell division (hyperplasia and hypertrophy) of the plant tissue and
form a gall.. Gall damage can be benign to deadly. Bacterial galls can
be produced by the genus Agrobacterium.

116 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig 2.5.1 Different types of Bacterial diseases.
A. Leaf spot, B. Canker, C. Crown Gall,
D. Vascular wilt, E. Soft rot, F. Bacterial Scabies.

Know more: Crown Gall of Agrobacterium tumefaciens most


often found in commercial nurseries. The colour of galls (tumors)
caused by that bacterium can be orange-brown. Pseudomonas sp.
bacterial infection causes gummosis.

Certain species of Arthrobacter, Pseudomonas, Rhizobacter and


Rhodococcus. Agrobacterium tumefaciens, A. rubi and A. vitis cause
galls, which are referred as crown gall, crown knot, root knot and
root gall. Species of these bacteria are thought to be present in most
agriculture soils. Crown gall first appears as small, whitish, soft
round over growths typically on the plants crown or at the main
root.

d.Bacterial Vascular Wilts: Vascular wilts caused by bacteria


primarily affect vegetables, field crops, ornamentals and some

APSCHE : SEMESTER-I BOTANY TEXT BOOK 117


tropical plants. The causal pathogen enters, multiplies and moves
through the xylem vessels of the host plant and interferes with the
translocation of nutrients and water by producing gum
(Fig.2.5.1.d). The symptoms of bacterial wilt disease include wilting
and death of the aboveground parts of the plant. In some cases
bacterial ooze seeps out through stomata or cracks onto the surface
of infected leaves. This ooze does not occur until the infected plant
tissue is dead. The pathogen will often destroy parts of the cell wall
of the xylem vessels resulting in pockets of bacteria, gums and
cellular debris.

e.Bacterial Soft Rots: Primarily the bacteria that cause soft rots
in living plant tissue are Erwinia sp., Pseudomonas sp., Bacillus sp.
and Clostridium sp. (Fig.2.5.1.e). Soft rots attack a large number
of hosts and are best known for causing disease in fleshy plant
structures both above and below ground. These bacteria are almost
always present where susceptible plants under stress are in the field
or in storage. Soft rot pathogens enter the host through wounds.
After entering the host tissue these bacteria produce enzymes that
break down the middle lamella causing separation of the cells at
the site of the infection. The cells die and disintegrate. Rotting tissue
becomes watery and soft and bacteria will form a slimy foul smelling
ooze. eg., Bacterial Soft Rot of melon (Erwinia carotovora), Bacterial
Wilt of Tomato (Ralstonia solanacearum)

f. Bacterial scabs: Bacterial scabs primarily infect underground


parts of plants such as potatoes. Common scab of potato is caused
by Streptomyces. Scabies causes localized scabby lesions on the outer
surface of the tuber followed by corky tissue will form below and
around the lesion. Rot pathogens can gain entrance into the host
tissue through these lesions and further degrade the host
(Fig.2.5.1.f).

118 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Know more : General classification some of the phytopathogenic
prokaryotes:

Kingdom: ProcaryotaeBacteria have Rhizobium-the cause of nitrogen-


cell membrane and cell wall and no fixing root nodules in legumes.
nuclearmembrane. Family: still unnamed
Division: Bacteria- Gram-positive Genus: Xylella- xylem-inhabiting
Class: Proteabacteria- Mostly single bacteria- leaf scorch and dieback
celled bacteria. disease on trees and vines.
Family: Enterobacteriaceae Candidatus liberobacter-Phloem
Genus: Erwinia - fire blight of pear inhabiting- Citrus greening disease.
and apple, Stewart’s wilt in corn, and Unnamed- laticifer-in habiting-
soft rot of fleshy vegetables. Bunchy top disease of papaya.
Pantoea- wilt of corn. Division: Firmicutes - Gram-
Serratia, S. marcescens,-a phloem- positive bacteria.
in habiting bacterium causing yellow Class: Firmibacteria– Mostly single
vine disease of cucurbits. celled bacteria.
Sphingomonas- brown spot of yellow Genus: Bacillus-causing rot of
Spanish melon fruit tubers, seeds, and seedlings and white
Family: Pseudomonadaceae stripe of wheat.
Genus: Acidovorax - leaf spots in Clostridium, causing rot of stored
corn, orchids and water melon. tubers and leaves and wetwood of elm
Pseudomonas -numerous leaf spots, and poplar.
blights, vascular wilts, soft Class: Thallobacteria– Branching
rots,cankers, and galls. bacteria.
Ralstonia -wilts of solanaceous crops. Genus: Arthrobacter- bacter ial
Rhizobacter - Bacterial gall of carrots. blight of holly, thought to be the cause
Rhizomonas-the corky root rot of of douglas- fir bacterial gall.
lettuce. Clavibacter- bacterial wilts in alfalfa,
Xanthomonas-numerous leaf spots, potato, and tomato.
fruit spots, blights of annual and Curtobacterium- causing wilt in
Perennial, plants, vascular wilts and beans and other plants.
citrus canker. Leifsonia- ratoon stunting of
Xylophilus- Bacterial necrosis and sugarcane.
canker of grapevines. Rhodococcus- fasciation of sweet pea.
Family: Rhizobiaceae Streptomyces-common potato scab
Genus: Agrobacterium, - crown gall
disease

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2.5.2. Citrus canker :
Introduction : Citrus Canker is a bacterial disease of worldwide
distribution in all citrus growing countries. It is a serious menace
and the disease causes serious damage in India, China, Japan and
Java. The pathogen incites severe canker disease on stems, leaves
and fruits of citrus species.Canker is not species specific, and it
attacks most of the species/varieties of citrus. The acid lime plants,
sweet orange and grape fruit are most susceptible species.
Symptoms of Citrus Canker Disease: Crest-like disease lesions
or scabby spots and small cankers (open wounds or dead tissue
surrounded by living tissue) appear on all over ground parts of the
plant such as leaves, young branches and fruits (Fig.2.5.2). The
trees are, however, not commonly killed. The lesions on the foliage,
at first, appear on the lower surface as small round raised spots.
These are translucent and of yellowish brown colour. Later the spots
turn white or greyish and finally rupture. The older lesions are corky
and brown, sometimes purplish. The necrotic brownish canker
regions are surrounded by a yellowish brown to green raised margin
and distinct watery yellow halo region. The yellow halo region is
free from the pathogen. The cankerous lesions contain the pathogen
in millions. The lesions on the twigs are usually irregular in form.
The lesions on the fruit are similar to those on the leaves but lack
the yellow halo (Fig.2.5.3).
FACT: Maririe observed that the halo regions are formed due to
the response of the host tissue to a diffusible metabolite of the
pathogen. Padmanabhan et al. (1975) reported accumulation of
malic acid in the halo region due to increased respiration in this
region.

Causal Organism: The causal organism is the bacterial pathogen


Xanthomonas citri, now called X. campestrispv. citri (Hasse)

120 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig 2.5.2 Citrus canker disease Fig 2.5.3 Canker lesions of leaf (A) and
a) fruit b) leaf c) twig Fruit (B) appear similar without halo

Dowson. It consists of a short, motile rod (1.5-2.0 x 0.5- 0.75 µ)


furnished with a single polar flagellum (monotrichous). It lacks
endospore formation. It is a gram negative, aerobic form
surrounded by a mucilaginous capsule. It forms chains.The climate
factors which favour the disease are the mild temperature and wet
weather. The most suitable range of temperature appears to be 20ºC
to 30ºC.
Disease Cycle: Infection takes place through the stomata and
wounds. The disease is not soil borne. The pathogen perennates in
the old lesions on the twigs still attached to the host plant.From
there it is carried by driving rains and by insects to new localities.
Man functions as the chief agent of dissemination by planting
infected nursery stock in new localities (Fig.2.5.4).
Control Measures of Citrus Canker Disease: In order to prevent
economic loss or to reduce to a low level Citrus canker can be
controlled in two ways A. Physical and Mechanical Measure and B.
Chemical Control.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 121


Fig 2.5.4 Life cycle of citrus canker caused by Xanthomonas citri.
Source: https://cutt.ly/NWfRvbw

A. Physical and Mechanical Control:


1. Eradication:The disease is controlled by the eradication of
diseased trees by removing the trees with advanced infection and
burning them.
2. Pruning: The infected trees may be cured by removing the
diseased leaves and branches with pruning scissors and then spraying
the trees with 1% Bordeaux mixture at regular intervals.
3. The use of disease free nursery stock for planting is the best
method of controlling the disease. Citrus nurseries should be raised
in places away from the regions of heavy and protracted rainfall.
”khatti”
4. Burning of the fallen infected leaves and twigs.
B. Chemical Control:
1. Bordeaux mixture and lime sulphur: Spraying with Bordeaux
mixture and lime sulphur is a useful measure to protect the fruit. It
should be done during the first three months after the beginning of
fruit formation. Spraying should commence before the onset of rains
and repeated during the rainy season.

122 APSCHE : SEMESTER-I BOTANY TEXT BOOK


2. Antibiotics: Antibiotic sprays are useful in controlling the
disease. Streptomycin sulphate and Phonomycin have been found
to be effective antibiotics (Rangaswamy, 1957).
3. Neem cake: Spraying with neem-cake is effective in controlling
citrus canker (Vaheeddudin 1959).

Know more: Common bacterial diseases of vegetable crops with some


typical symptoms.
 (brassicas)

 Pseudomonas spp.
 Rhizomonas suberifaciens
 (cucurbits);

 Pseudomonas corrugata
(tomatoes);
 Xanthomonas campestris pv.
phaseoli
 (beans)

 Erwinia carotovora pv. atroseptica

Know more: “Bacterial cellulose (BC) produced by some


microorganisms has been widely accepted as a multifunctional
nano-biomaterial. It is composed of linear glucan molecules
attached with hydrogen bonds, which appears similar to plant
cellulose. However, when compared with other conventional
natural or synthesized counterparts, BC performs better in areas
such as biomedicine, functional devices, water treatment,Nano
fillers, etc. for its distinct superior chemical purity, crystallinity,
biocompatibility, and ultrafine network architecture. When BC
is incorporated in a material or used as a scaffold, novel features
result that are related to BC’s intrinsic characteristics (Yang
Huang et al 2014.)”

APSCHE : SEMESTER-I BOTANY TEXT BOOK 123


Self- Analysis
1. Compare the general characteristics of three groups of special
bacteria studied by you.
2. Tabulate the unique features of each group of special bacteria
in your syllabus.
3. Compare and contrast the characteristics of special bacteria
with that of eubacteria.
4. Study the significance of actinomycetes in production of
antibiotics.
5. Evaluate the features of Archaea in relation with bacteria and
eukaryotes.
6. Justify the significance of taxonomic status of Cyanobacteria
in comparison with bacteria and eukaryotic algae.
7. Differentiate the cellular changes occur in Bacteria during
binary fission endospore formation.
8. Compare the genetical differences between HFr strain and F+
strain.
9. Compare and contrast the bacterial plant disease symptoms
with viral and fungal plant disease symptoms.

Self-Assessment
1. Taq polymerase used for 2. Transcription
PCR is obtained from mechanism
a. Cyanobacterium 3. Methionyl-tRNAmet
b. Archaea 4. Sensitivity to anisomycin
c. Eubacterium a. 1, 2 & 3
d. Actinobacterium b. 1, 3 & 4
2. Methanogens belong to c. 1, 2 & 4
following group d. 2, 3 & 4
a. Mycoplasma 4. The habitat of Actinomycetes
b. Chlamydia israelii is
c. Eubacterium a. Soil
d. Archaea b. Water
3. The following features are c. Skin
common for archaea and d. Oral cavity
eukaryotes. 5. The following prokaryotes
1. Multiple RNA have highest G+C content in
polymerases and the biome.
polypeptides

124 APSCHE : SEMESTER-I BOTANY TEXT BOOK


a. Cyanobacterium a. Conjunction
b. Archaea b. Transformation
c. Eubacterium c. Mutation
d. Actinomycetes d. Plasmids
6. The following cyanobacterium is 12. Bacteria that are
a source of Single Cell Protein. responsible for
a. Anabaena fermentation of dairy milk
b. Nostoc are
c. Spirulina a. Azatobacter
d. Oscillatoria b. Rhizobium
7. Siderophores have affinity with c. Lactobacillus
following metal ion d. Hay bacillus
a. Nickel 13. Role of bacteria in carbon
b. Iron cycle is
c. Zinc a. Photosynthesis
d. Copper b. Chemosynthesis
8. Identify the Gram +ve bacteria c. Breakdown of
from the following organic
a. Staphylococcus, Bacillus compounds
b. d. Assimilation of
c. Staphylococcus,Escherichia nitrogen
d. Salmonella,Bacillus compound
9. Bacterial chlorophylls are present 14. Mesosomes are the part of
in a. Plasma membrane
a. Rhodospirillum rubrum b. ER
b. Neisseria gonorrhoeae c. Lysosomes
c. Salmonella typhi d. Golgi
d. Streptococcus 15. A bacterium containing
pneumoniae prophage is called as
10. Identify the chemoautotrophic a. Lytic
bacterium in the following b. Lysogen
a. Chlorobium c. Lytogen
b. Beggiatoa d. None of these.
c. Heliorestis 16. Viral genome can become
d. Bacillus integrated into the
11. Lederberg and Tatum (1946) bacterial genomes are
described the phenomena of known as

APSCHE : SEMESTER-I BOTANY TEXT BOOK 125


a. Prophases microbiology-manageable/
b. b. Temperate phage tr10761.trhttps://www.youtube.com/
c. Bacteriophage watch?v=8SsF_ivLejY
d. Metaphage https://www.xylem-analytics.asia/
17. In the industrial production of parametersdetail.php?Blue-Green-
streptomycin, the secondary Algae-BGA-13
metabolite or byproducts https://microbiologysociety.org/why-
a. Vitamin – B12 microbiology-matters/what-is-
b. Vitamin – C microbiology/bacteria.htmlhttps://
c. Vitamin – B6 www.carolina.com/teacher-resources/
d. Ethanol Interactive/carolina-labsheets-
18. Conjugation was reported for introduction-to-prokaryotes-archaea/
the first time in following tr30045.trhttps://www.carolina.com/
bacterium
teacher-resources/Interactive/
a. Escherichia coli
b. Salmonella Typhi halobacteria-making-microbiology-
c. Bacillus anthracis manageable/tr10761.trhttps://
d. Streptococcus www.youtube.com/
pneumoniae watch?v=8SsF_ivLejYhttps://
19. Bacterial transformation
www.xylem-analytics.asia/
occurs by
parametersdetail.php?Blue-Green-
a. Physical contact of bacteria
b. By bacteriophages Algae-BGA-13https://
c. A bacterium takes genes microbiologysociety.org/why-
from surrounding microbiology-matters/what-is-
medium microbiology/bacteria.htmlhttps://
d. None of the above
www.youtube.com/
20. Crown gall is due to
a. Agrobacterium tumefaciens watch?v=4DYgGA9jdlEhttps://
b. Agrobacterium rhizogenes www.youtube.com/
c. Bacillus thuringiensis watch?v=OOFJyw0EYBU
d. Pseudomonas fluorescens Online sources:
https://ucmp.berkeley.edu/archaea/
Interactive Links: archaea.html
https://www.carolina.com/teacher- https://www.youtube.com/
resources/Interactive/carolina- watch?v=VGcT1-XaWgk
labsheets-introduction-to-prokaryotes- https://www.frontiersin.org/
archaea/tr30045.trhttps:// journals/microbiology/sections/
www.carolina.com/teacher-resources/ biology-of-archaea
Interactive/halobacteria-making-

126 APSCHE : SEMESTER-I BOTANY TEXT BOOK


https:// https://en.wikipedia.org/wiki/Bacteria
www.microbiologyresearch.org/ https://www.youtube.com/
content/journal/micro/10.1099/ watch?v=TDoGrbpJJ14https://
mic.0.000944?TRACK=RSS www.biologyexams4u.com/2012/12/
https:// csir-ugc-net-jrf-life-
www.microbiologyresearch.org/ sciences.htmlhttps://
content/journal/mgen/10.1099/ www.mcqbiology.com/2012/10/mcq-
mgen.0.000498?crawler=true on-microbiology-bacteria.htmlhttps://
https://hcb-1.itrcweb.org/ en.wikipedia.org/wiki/Bacteriahttp://
introduction/ go.microsoft.com/fwlink/p/
https://en.wikipedia.org/wiki/ ?LinkId=255141
Bacteria Reference Books:
https://www.youtube.com/ 1. Bhattachar jee, R.N., (2017)
watch?v=TDoGrbpJJ14 Introduction to Microbiolog y and
https://sciencing.com/nutritional- Microbial Diversity, Kalyani Publishers,
types-bacteria-2515.html New Delhi.
https://sciencing.com/types- 2. Dubey, R.C. &D.K.Maheswari (2013)
heterotrophic-bacteria- A Text Book of Microbiology,S.Chand&
6884639.html Company Ltd., New Delhi .
https://ucmp.berkeley.edu/archaea/
3. lczar Jr., M.J., E.C.N. Chan
archaea.html
&N.R.Krieg (2001)Microbiology, Tata
https://www.youtube.com/
McGrawHill Co, New Delhi .
watch?v=VGcT1-XaWgk
https://www.frontiersin.org/ 4. Presscott, L. Harley, J. and Klein, D.
journals/microbiology/sections/ (2005)Microbiology, 6th edition, Tata
biology-of-archaea
McGraw –Hill Co. New Delhi.
https://
www.microbiologyresearch.org/ 5. Fogg, G.E., W.D.P.Stewart, P.Fay and
content/journal/micro/10.1099/ A.E.Walsby (1973) The Blue Green
mic.0.000944?TRACK=RSS Algae, Elsevier Inc., Amsterdam,
https:// Netherlands
www.microbiologyresearch.org/
content/journal/mgen/10.1099/
and S. T. Williams (1984) The Biology
mgen.0.000498?crawler=true
of Actinomycetes, Academic Press,
https://hcb-1.itrcweb.org/
introduction/ Cambridge, U.S.A.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 127


Unit - 3
FUNGI & LICHENS

CONTENTS LEARNING OUTCOMES: At the end of


the unit the learners should be able to :
1. General
characteristics of  Understand the general characteristics
fungi and of fungi
Ainsworth  Get an in-depth knowledge on diversity
classification. of fungi based on Ainsworth
classification.
2. a) Rhizopus
(Zygomycota),  Acquire knowledge on structure ,
b) Puccinia reproduction and life cycles of two
(Basidiomycota), fungi.

3. Economic uses of  Explain the economic significance of


fungi. fungi.

4. Plant diseases  Identify the symptoms of plant


caused by Fungi; disease of due to fungal pathogens.
Blast of Rice.  Discuss the characteristics, ecological
5. Lichens and economic importance of lichens

The Fungi is one of the large and diverse group in the plant kingdom.
The branch of Botany deals with fungi is called mycology. They resemble algae
in many respects; hence, included in the division, Thallophyta and . In
the five-kingdom system of classification, the fungi are grouped in a
distinct kingdom within the eukaryotes. It includes diversity ranging
from unicellular yeasts and moulds to large multicellular mushrooms.
There are about 50,000 to 1,00,000 known species of fungi in the world.
According to Ainsworth (1961), there are 4,300 and 50,000 species of
fungi, at present a total of 50,000 to 1,00,000 known species of fungi
reported in the world.

128 APSCHE : SEMESTER-I BOTANY TEXT BOOK


General characteristics of fungi and Ainsworth
Chapter -3.1
classification (upto classes)

Learning outcomes: At the end of the chapter you should be able


to :
1. Discuss and debate the general characters of fungi in
compression with algae.
2. Explain the characteristic features of fungi included in
different taxa of Ainsworth classification.

3.1.1 General characters of fungi : Fungi are eukaryotic,


achlorophyllous, unicellular or multicellular organisms, which
reproduce by asexual and sexual spores. The plant body is simple,
and consists of network of branched filaments called the hyphae.
The entangled mass of hyphae is the mycelium. Hypha exhibit
apicalgrowth. Plant body organization show variation from
unicellular (e.g., yeasts), coenocytic (Mucor) and multicellular
(Penicillium, Mushroom). Fungi are eukar yotic and possess
membrane bound nuclei (containing chromosomes) and a range of
membrane bound cytoplasmic organelles (e.g., mitochondria,
vacuoles, endoplasmic reticulum). Protoplasm of a hypha or cell is
surrounded by a rigid wall composed of chitin and glucans. In
some species cell wall contain cellulose. Their nuclei are typically
haploid and hyphal compartments are often multinucleate. Some
yeast possesses diploid nuclei. Mode of nutrition is
chemoheterotrophic. Fungi are saprophytic, parasitic or symbiotic
in nature. Reserve food is trehalose, glycogen, sugar alcohols and
lipids. Reproduce vegetatively, asexually and sexually and often
produce fruit bodies (Ascomycetes and Basidomycetis).

Somatic or Vegetative structure of plant body : Based on


vegetative and reproductive nature of the thallus, fungi are classified
into two types.

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
   
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
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
              
  

       



     
  
  


The mode of nutrition in fungi: Many of the fungi are saprobes, and
bring about the decay of organic materials. When they act as parasites,
they attack living protoplasm and cause diseases of plants, animals and
human beings. They also utilise pre-existing organic sources of carbon in
their environment and the energy from chemical reactions to synthesize
the organic compounds for their growth and energy. By this, fungi get food
materials converting complex compounds into simple molecules.

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a. Holocarpic thallus: In holocarpic forms therefore, somatic
and reproductive phases do not exist together at a time.

b. Eucarpic thallus: The thallus is differentiated into


vegetative part for the absorption of nutrients, and a
reproductive part for reproductive structure. Such thalli are
called as eucarpic. e.g., Pythium aphanidermatum.

In Chytridiaceae and Rhizidiaceae the unicellular thallus is


evolved into a rhizomycelium which bears one reproductive
structure (eucarpic monocentric) while in Cladochytriaceae the
rhizoidal mycelium bears several reproductive units (eucarpic
polycentric). The slime moulds (Myxomycetes) and yeasts are also
unicellular forms. Based on cell organization, thallus of Fungi shows
two kinds of somatic plant body.

1. Non mycelial fungi which includes unicellular forms

2. Mycelial fungi with filamentous forms.

Mycelial thallus: Unicellular thallus:Lower fungi have a very simple


and primitive type of thallus. The thallus is unicellular and the entire
thallus gets transformed into a single reproductive structures.
Holocarpic monocentric) e.g., Olpidium or into several reproductive
units (holocarpic polycentric) e.g., Synchytrium. In Synchytrium, the
mycelium is completely absent whereas in yeasts, the mycelium is
not well developed. The vegetative body of the fungi is generally
called as thallus and is filamentous in nature. It is highly branched
and thallus is called the mycelium. The long tubular branches are
called hyphae. The hyphae of most of the fungi are 5 to 10
micrometers (µm) in width. There The hyphae are usually colourless
en masse and generally have a whitish appearance in some species.
Pigments like green, black, yellow, blue, violet or red may sometimes

APSCHE : SEMESTER-I BOTANY TEXT BOOK 131


 
 


 

 
    

     

  

* * * * * * * *
be present in the cell wall or in the cell contents. There are also
certain specialised forms of the mycelia like rhizomorphs,
sclerotia, stroma, etc.

Know more: The cell wall may be in the form of carbohydrate


e.g., hemicellulose, pectose, callose or other related compounds.
De Bary has termed it as fungal cellulose. True cellulose is present
in some of the Phycomycetes only. Chitin is usually present in
Chytridiales, Blastocladiales, Mucorales, etc. Calcium carbonate
and other salts may also be deposited on the cell walls.

The cytoplasm contains vacuoles, nuclei and food reserves


in the form of glycogen or oil droplets. Due to absence of septa the
cytoplasm is continuous in the entire mycelium of the
Phycomycetes. Such a mycelium is termed aseptate and coenocytic
(multinucleate). In higher forms of fungi e.g., Ascomycetes,

132 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Basidiomycetes and Deuteromycetes the mycelium is septate and
multicellular with usually one but sometime more nuclei in each
cell. The septa are perforated and allow the nuclei and nutrients to
move from one cell to another cell through the plasmodesmata.The
septa of different classes of fungi have been observed by Moore
and Mc Alear (1962). They have observed a simple pore in the septa
of Euascomycetes, and Uredinales and a complex pore called dolipore
in the Basidiomycetes except the Uredinales. The edge of the septa
around the pore is swollen. The swollen rim is guarded on both
sides by a membranous cap like structure termed septal pore cap or
parenthosome. This prevents the migration of nuclei from one cell
to another cell.

In Basidiomycetes, there are two kinds of mycelia i.e.,


primary and secondary. The former consists of uninucleate cells
and is called monokaryotic, whereas the latter consists of
binucleated cells and is called dikaryotic. In some fungi, genetically
different nuclei come to lie in one cell. Such mycelia are termed
heterokaryotic.

Ultrastructure of a Fungal Hypha: The cell wall is composed of


microfibrils of fungal cellulose and chitin. The plasma membrane is
a typical three-layered unit membrane. Occasionally organelles
called lomasomes occur in between the cell wall and the plasma
membrane. They are believed to be concerned with the synthesis of
non-cellulosic polysaccharides. The cytoplasm contains small
vacuoles lined by the tonoplast. Endoplasmic reticulum is
widespread. Mitochondria, ribosomes and dictyosomes are scattered
within the cytoplasm. The nucleus has a double-layered membrane
with nucleolus and nuclear pores (Fig.3.1.1).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 133


Figure 3.1.1 Structure of Hypha

Flagella: The fungal flagella have the typical 9+2 arrangement of


fibrils. The flagella are of two kinds.The whiplash type which is a
long flagellum ending into a blunt tip or an end-piece.It has smooth
surface, and the tinsel type, which is usually a small flagellum with
fine hairy appendages called flimmer hairs or mastigonemes.

Modifications of mycelium:
Plectenchyma : The mycelium of fungi consists of a network of
loose hyphae. In many higher fungi, however, the hyphae grow
together in groups, intertwine, adhere and form a loose or compact
mass of tissue-like structure termed plectenchyma. Plectenchyma
can be of two kinds (Fig. 3.1.2).

Prosenchyma Pseudoparenchyma

Figure 3.1.2 Other forms of Fungal tissues

134 APSCHE : SEMESTER-I BOTANY TEXT BOOK


(a) Prosenchyma: The hyphae are loosely interwoven and are
nearly parallel to one another. The hyphae can be observed as
elongated cells.
(b) Pseudoparenchyma: The hyphae are very compactly arranged
and appear as parenchyma like oval cells. The typical elongated
nature of the hyphae cannot be distinguished.
Nutrition and Growth : Fungi obtain their food either by infecting
living organisms as parasites or by attacking dead organic matter
as Saprobes. Fungi which live on dead matter and are incapable of
infecting living organisms are called obligate saprobes. Those
fungi capable of causing diseases or of living on dead organic matter,
according to circumstances, facultative parasites (or facultative
saprobes), and those which cannot live except on living protoplasm
are called obligate parasites. Some fungal hyphae show a
widespread association with the roots of higher plants and this
association is known as mycorrhiza. In these associations both
plant and fungus derive the benefit. Many fungi are capable of
synthesizing vitamins they require for their growth and
reproduction. Some, however are deficient in thiamine or biotin or
both and must obtain these or their precursors from the substratum.
Fungi usually store excess food in the form of glycogen and oil.
Know more : Fungi differ from plants, they require simple food in
order to live; and are incapable of manufacturing their own. If
carbohydrates given from outside preferably in glucose, sucrose or
maltose forms, most fungi can synthesize their own proteins by
utilizing inorganic or organic sources of nitrogen and various mineral
elements essential for their growth.

Activity: Collect the soil samples from rhizosphere of your


nearby fields. Observe the fungal organisms which are
beneficial to plants.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 135


Reproduction : The fungi show three types of reproduction namely,
vegetative, Asexual and sexual. Prosenchyma and
pseudoparenchyma are the constituents of several kinds of somatic
and reproductive structures of fungi. They are described below:
1. Stroma: The stroma is a compact pseudoparenchymatous
somatic tissue. The sporophores of a parasitic or saprophytic fungus
get embedded into it.

II. Sclerotium: In some higher fungi like Claviceps, the mycelium


after vigorous active growth forms hard and compact masses of
irregularly shaped structures. They may be spherical, cylindrical or
football like. They are generally black or purple outside and grey-
white inside. They have rich amount of food reserve and are
perennating structures; germinate when favourable conditions
reappear.

III. Bulbils: These are small sclerotia composed of a few layers of


cells. They are often present in large numbers.

IV. Rhizomorphs: A number of fungi produce more or less firm


cortical layer made up of long strands of parallel hyphae forming
compact masses. They resemble the finer roots of a tree and their
diameter is, uneven. This is advantageous for the fungus, since
single hyphae are easily damaged. They creep over long distances,
often through ver y unfavourable environments to new
environments, ex: Armillaria mellea, the Honey Mushroom. They
attack roots of many kinds of trees and may often be found
underneath the dead bark of the roots or even the trunks.

V. Sporophores: From the vegetative prostrate hyphae arise a


number of erect hyphae called the sporophores. They are of various
types.

136 APSCHE : SEMESTER-I BOTANY TEXT BOOK


a. The sporangiophores are sporophores which form the spores
endogenously in spherical terminal structures called the
sporangia. The enclosed non-motile spores are called
sporangiospores but, if the spores are motile, they are
termed zoospores.
b. If the sporophores cut off their spores exogenously, they
are called conidiophores and the spores as conidia. The
conidiophores may be simple or branched and bear conidia
singly or in chains.
c. In higher fungi, the hyphae forming the conidiophores show
a tendency to come together into groups or fructification.
The Coremia are fruiting bodies of certain fungi, consisting
of a loosely bound bundle of conidiophores. Each coremium
consists of a sterile stalk, terminating in a fertile head
composed of conidia-bearing hyphae, e.g. Stynamus
thyrosoides.
d. In rusts, sporophores are grouped into sori. Sometimes, the
sporophores join in groups of plectenchymatous structures
forming flask-shaped structures, the pycnia (pycnidia), in
whose interior they cut off the conidia, termed pycniospores
or stylospores. Pycnidia are provided with openings at their
apical ends, known as ostioles.
e. The widespread cushions or saucer-shaped fructifications,
called the acervuli in parasites and barrel or sausage-shaped
sporodochia (singular, sporodochium) in saprophytes.
A. Vegetative reproduction:
(i) Fragmentation: The hyphae break up into pieces of fragment
accidently or through external forces and each piece growing
into a new individual.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 137


(ii) Fission: This is the commonest and simplest method of
vegetative propagation to increase in the number
especially in some of the yeasts. This involves the
splitting of the cell into two daughter cells by simple
constriction or transverse wall.

(iii)Budding: In Saccharomyces (Yeasts), the parent-cell buds off


new cells which separate from the parent-cell at an early
stage and develop into new individuals (Fig 3.1.3).

Figure 3.1.3 A-D Budding, E-H Fission


Source: http://eagri.org/eagri50/PATH171/lec03.pdf

B. Asexual reproduction : Fungi reproduce asexually by means


of spores. The spores show great variation in their form and mode
of origin. Usually, they are oval or spherical in outline, yet in certain
cases they may assume spindle-shaped, club-shaped or even They
vary in colour from hyaline through green, orange, red, brown to
black; in number of cells from one to many: in size, from minute to
large and in the manner in which they are borne. In most cases, a
spore is bounded by two walls, of which the external, termed as
exine or exosporium is often thick and ornamented, while the inner,
intine or endosporium is usually thin and transparent. According
to their mode of origin, the following types of spores are recognised:
(i) Endospores(ii) Conidia (iii) Oidia(iv) Chlamydospores(v)
Pycniospores (vi) Ascospores (vii) Basidiospores (Fig 3.1.4).

138 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig 3.1.4 Different types of spores in Fungi

(i) Endospores: They are produced endogenously by the division


of the protoplasm within an enlarged cell, the sporangium, borne
at the end of a special hypha, called the sporangiophore. The
sporangia, though chiefly spherical or oval in outline, may be
cylindrical or club-shaped. The endospores are usually non-motile
and bounded by cell-wall (Rhizopus, Mucor). The non-motile
endogenously produced spores are called sporangiospores or
aplanospores.In the algal fungi Phycomycetes naked endospores
furnished with vibratile cilia are produced under moist conditions
in sacs called zoosporangia These are called zoospores or
swarmspores and they swim freely in water on liberation, after the
rupture of zoosporangia (Albugo, Pythium, Saprolegnia, Achlya). The
number of spores produced by those fungi is enormous.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 139


(ii) Conidia: These are non-motile spores produced nakedly
(exogenously) by constriction at the end of special hyphal branches,
called the conidiophores. They are produced singly (Pythium,
Phytophthora) or in the chains (Aspergillus, Pencillium). The tip of
the conidiophore is spherical (Aspergillus) and gives out a number
of small pegs like branches, the sterigmata, bearing chains of
basipetal conidia while in Pencillium the sterigmata bearing chains
of conidia are simply the ends of repeatedly dichotomous, brush
like conidiophore. In Aspergillus, the sterigmata cutting more and
more conidia at the tip from below resulting in the form of a chain
of conidia, i.e., the oldest conidium is at the tip and the youngest is
at the base, while in the latter (Penicillum), the arrangement of
conidia may be the same or they may be acropetal, i.e., first-formed
conidium buds off a new conidium and this process is repeated,
resulting in the formation of a chain of acropetal conidia with the
youngest conidium at the apex and the oldest at the base.
(iii) Oidia: In several fungi (Mucor), the hyphae forming the
mycelium often become divided by transverse walls into a large
number of short segments, each of which is capable of germinating
and developing into a new plant. These segments, which may remain
united in chains or become free from one another, are known as
oidia.
(iv) Chlamydospores: They are produced like oidia, but they differ
from the latter in being thick-walled and coloured black or brown.
They are terminal or produced at irregular intervals along the
hyphae. On germination, they produce a short hypha bearing either
sporangia or conidia.
(v) Pycniospores: These are small conidia like bodies produced in
flask-shaped cavities called the pycnia o pycnidia (Puccinia).

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(vi) Ascospores: These are uninucleate, unicellular, non motile,
usually eight in number produced in sac like structure called ascus,
characteristic of Ascomycetes.
(vii) Basidiospores: They are characteristic of Basidiomycetes,
produced exogenously by a club shaped structure called basidium,
on short-stalked outgrowths, the sterigmata. Usually four
basidiospores are produced by a basidium.
(viii) Uredospores and Teleutospores: In the rusts, special forms
of binucleate summer-spores, called the uredospores are produced
in clusters called the uredesori or uredinia. Each spore is borne singly
at the tip of a stalk. They serve for dispersal during the summer
and, in certain cases (Puccinia in Australia), they perennate
throughout the winter, re-infecting the summer host-plant (wheat)
thus propagating the fungus in the absence of acidia or the sexual
stage. In the same way many other fungi (Fungi) imperfecti
propagated by the asexual spores with no nuclear fusion in the life-
cycle for many generations. Teleutospores are formed in Puccinia
and other fungi for perennation.
C. Sexual reproduction : The process of sexual reproduction in
fungi involves the union of two compatible nuclei (Fig. 3.1.5) and
consists of three typical phases. The nuclei may be carried in motile
or non-motile gametes or in gametangia or in somatic cells of the
thallus.
I. Phases of sexual reproduction: Three typical phases occur in
sequence during the sexual reproduction. They are Plasmogamy,
Karyogamy and Meiosis.
A. Plasmogamy: In plasmogamy, anastomosis of two cells or
gametes and fusion of their protoplasts take place. In the process
the two haploid nuclei of opposite sexes (compatible nuclei) are
brought together but eh nuclei will not fuse.

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B. Karyogamy: The fusion of two haploid nuclei brought together
as a result of plasmogamy is called karyogamy. This stage follows
immediately after plasmogamy in many of the lower fungi or may
be delayed in higher fungi. In higher fungi, plasmogamy results in
a binucleate cell containing one nucleus from each cell. Such a pair
of nuclei is called dikaryon. These two nuclei may not fuse until
later in the life history of the fungus. Meanwhile, during growth
and cell division of the binucleate cell, the dikaryotic condition may
be perpetuated from cell to cell by conjugate division of the two
closely associated nuclei and by the separation of the resulting sister
nuclei with two daughter cells. Nuclear fusion, which eventually
takes place in all sexually reproducing fungi, is followed by meiosis.
C. Meiosis : Karyogamy results in the formation of a diploid (2n)
nucleus. Meiosis reduces the number of chromosomes to haploid
and constitutes the third phase of the sexual reproduction. This
nucleus undergoes a reduction division to form two haploid nuclei
each with ‘n’ chromosomes, they again undergo mitotic division to
form four nuclei. In ascomycetes, another nuclear division takes
place resulting in the formation eight nuclei. The nuclei get
surrounded by a small amount of cytoplasm and secrete a wall to
become spores. In a true sexual cycle, the above three phases occur
in a regular sequence and usually at specified points.
II. Methods of sexual reproduction : The fungi employs five
methods i.e., 1. Planogametic copulation, 2. Gametangial contact
(Gametogamy), 3.Gametangial copulation (Gametangiogamy) ,
4. Spermatization, and 5. Somatogamy to bring the compatible
nuclei together for fusion.
1. Planogametic copulation or conjugation : This involves the
fusion of two naked, free gametes, one or both of which may be
motile. The motile gametes are called planogametes. Depending
on size and motility of the fusing gametes, there are three types of
planogametic copulation.

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i. Isogamy: When the two fusing gametes are of the same shape
and size, they are called isogametes and their fusion isogamy.

ii. Anisogamy: This occurs only in one genus – Allomyces a chytrid,


the two planogametes are morphologically similar but different in
size.

iii. Heterogamy: It involves the fusion of a motile male gamete


with a non – motile female gamete (oosphere, contained in an
oogonium) the motile gamete, the antherozoid, enters the
oogonium and fertilizes the egg or oosphere. Heterogamy occurs
only in one genus Monoblepharis.

2. Gametangial contact: In this method the male gamete


(antheridium) and the female gamete (oogonium) come in contact
and one or more nuclei from the male gamete enter the female
gamete, oogonium dissolved in the intervening wall through a pore
or through a fertilization tube. Gametangial contact is common in
some Ascomycotina where antheridia and female organs ascogonia
may or may not be well defined.

3. Gametangial copulation: This is a process of fusion of entire


contents of the two mating gametangia. There are two types.

a. Mixing of entire protoplasm of male and female gametangia in


which two gametangia meet and their entire contents fuse in the
female gametangium leading to formation of a zygote. The zygote
forms a resting sporangium (Chytridiomycetes).

b. Isogamous copulation: Two morphologically similar gametangial


hyphae come in contact, the wall at the point of contact dissolves
and the contents mixes in the cell thus formed the zygospore
(Rhizopus).

4. Spermatization: Some fungi causing rusts produces numerous


minute, non-motile uninucleate, male cells called spermatia. They

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are produced in spiral receptacles called spermagonia or pycnia.
Special receptive hypha (or trichogyne) is there to which insects,
wind or water to the female gametangium usually become attached.
A pore develops at the point of contact and the contents of
spermatium pass into the particular respective hyphae. Plasmogamy
and initiation of the dikaryotic stage of the cell is initiated.

Fig. 3.1.5 Different types of sexual methods for nuclear fusion in fungi

Activity: Collect the damaged and spoiled vegetables from


the market and observe under microscope. List out the
fungal organisms based on their characters.

144 APSCHE : SEMESTER-I BOTANY TEXT BOOK


5. Somatogamy: No sex organs are produced and somatic cells
function as gametes in somatogamy . Hyphae anastomose and the
nuclei of opposite matting type are brought together in one cell
(e.g., Ascomycotina and Basidiomycotina fungi).
Life Cycles of Fungi : Fungi display different types of life cycles
centred around the sexual reproduction. Seven basic types of life
cycles in fungi are generally recognised. A life cycle involving
dikaryons is found only in fungi and nowhere else.These are:
1. Asexual cycle: the entire group of “fungi imperfecti” shows this
type of life cycle. There is no alternation of haploid and diploid
nuclear phases. The diploid (2n) phase is lacking.
2. Haploid cycle: The life cycle is completely haploid (n), the diploid
phase is restricted only to the zygote nucleus. This is the most
common type of life cycle found in majority of lower fungi and
Ascomycotina.
3. Hapliod cycle with restricted dikaryon: This type of life cycle
is found in higher Ascomycotina forming (n + n) ascogenous hyphae,
which are completely dependent on the haploid mycelium. This is
predominantly a haploid life cycle but it differs in the separation of
plasmogamy from karyogamy in space and time. The gametic nuclei
pair to form dikaryons which multiply by conjugate mitotic divisions
in the ascogenous hyphae. The dikaryons finally undergo karyogamy
and meiosis in the ascal primordia.
4. Haploid – dikaryotic cycle: Most of the Basidiomycotina,
excluding the smuts, exhibit this type of life cycle in which the
dikaryophase (n + n) is more extensive and also independent of the
haploid phase. The cycle comprises of two roughly equivalent phases
and terminates in a single diploid nuclear generation represented
by the diploid zygote nucleus.
5. Dikaryotic cycle: The complete life cycle is passed in dikaryotic
phase, and both the haploid and diploid generations are represented

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by single nuclear generation. The immediate products of meiosis
like the ascospores or the basidiospores fuse to initiate the
dikaryophase, which persists until karyogamy occurs. Examples are
the rusts and smuts.
6. Haploid – diploid cycle: This type of life cycle, which is
characteristic of algae, occurs only in two groups of fungi like the
Blastocladiales and Endomycetales (Ascocybe grovesii) and
Saccharomyces cerevisiae. In this type, haploid and diploid phases
are equally extensive and important.
7. Diploid cycle:This type of life cycle, that is completely diploid
except for the immediate products of meiosis, is reported in a
number of yeasts, Myxomycetes, Blastocladiales and the Oomycetes.

Parasexuality in fungi: Some fungi (Deuteromycetes) do not go


through a sexual cycle but derive many of the benefits of sexuality
through parasexuality. This is a process in which plasmogamy,
karyogamy and haploidization takes place, but not at specified
points in the thallus or the life cycle of an organism. Parasexual
cycle is ver y important in Deuteromycetes where sexual
reproduction does not take place. Some fungi, which reproduce
sexually, also exhibit parasexuality.Pontecorvo and Roper from the
University of Glasgow in Aspergillus nidulans, the imperfect stage
of Emericella nidulans first discovered parasexuality in 1952. Since
then it has been reported in number of fungi .

Know more: The phenomenon of existence of different kinds of


nuclei in the same individual is known as heterokaryosis. The
individual which exhibits heterokaryosis is called heterokaryon or
heterokar yotic. It has been demonstrated in numerous
Ascomycetes, Basidiomycetes and Fungi Imperfecti. In a
heterokaryotic individual, each nucleus is independent of all other
nuclei, but the structure and behaviour of the individual appear to
be controlled by the kinds of genes it contains and the proportion
of each kind

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3.1.2 Ainsworth Classification of fungi :

Earlier fungi were placed under division Thallophyta of the


sub kingdom Cr yptogamia of the kingdom Plantae.
Fitzpatrick (1930) divided the thallophyta into sub-divisions
Myxothallophyta (including Myxomycetes) and
Euthallophyta (which included lichens, bacteria, algae and
fungi).

Criteria for the classification of Fungi : Microscopic features


in the classification of fungi play an important role, since these were
not modified by external factors, because macroscopic characters
will be affected by environment. The nature of the asci and basidia
were taken as fundamental features in the classification of the
ascomycetes and basidiomycetes. The basidia in most basidiomycete
species are single-celled and four-spored and there are variations
within this form. There is also considerable variety in spores, they
may be smooth or ornamented, vary in shape from spherical to
highly angular, be white or coloured when viewed in mass and one-
celled or septate.
Geoffery Clough Ainsworth (1905-1998) thought that the
fungi have evolved from algae by loss of chlorophyll and thus are
properly placed in the ‘plant kingdom’. Some Mycologists believe
that the fungi had a common ancestry, with the protozoa but split
off at a very early stage of organic evolution. Ainsworth Published
fungal system of classification as a paper in a journal in 1966. It
was modified and published in “Ainsworth and Bisby’s Dictionary
of the Fungi” (1971) and the Fungi an advanced Treatise (1973). In
the Ainsworth system of classification, the fungi are treated as a
separate kingdom. Ainsworth (1971) has upgraded the fungi up to
kingdom level. In this the fungi treated in a separate kingdom, or a

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  

      



            
         
            
    


      
      
  

        
      
       
 
        




     



148 APSCHE : SEMESTER-I BOTANY TEXT BOOK


sub-kingdom of plant kingdom. Ainsworth divided the fungi into
two divisions namely two Myxomycota and Eumycota.
Division Myxomycota:
organisms. Possess either a plasmodium, a mass of naked,
multinucleate protoplasm, which feeds by ingesting particulate
matter and shows amoeboid movement, or pseudoplasmodium, an
aggregation of separate amoeboid cells. Both are of a slimy
consistency, hence they are also called ‘Slime moulds’. It includes
four classes. Namely Acrasiomycetes, Hydromyxomycetes ,
Myxomycetes and Plasmodiophoromycetes.
a. Acrasiomycetes: Free living assimilatory phase of amoebae
unite as a pseudoplasmodium before reproduction.
b. Hydromyxomycetes : Net like plasmodium is present.
c. Myxomycetes : Presence of free living saprobic
plasmodium.
d. Plasmodiophoromycetes: Presence of parasitic
plasmodium within host cells
Division Eumycota:
to recognize five subdivisions under this division.
A. Mastigomycotina:
aquatic. Four classes are included in this, each characterized by their
distinctive type of zoospores.
a. Chytridiomycetes : Posteriorly uniflagellate zoospore,
flagellum is whiplash type.
b. Hyphochytridiomycetes : Anteriorly unif lagellate
zoospore, flagellum is tinsel type.
c. Oomycetes : Biflagellate zoospores, posterior flagellum is
whiplash type and anterior flagellum is tinsel type. They
consist of cellulose cell wall.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 149


d. Plasmodiophoromycetes : Biflagellate heterokontean
swarmers (zoospores), flagella are whiplash type.
B. Zygomycotina:
are non-motile spores. Sexual reproduction takes place by the
complete fusion of two multi-nucleate gametangia producing a

also known as conjugation fungi. Cell wall is made up of chitin and


chitosan. It includes two classes.
a. Zygomycetes: Mostly saprophytic; sometimes weak
parasites or mycoparasites some attacking insects but then
developing mycelium inside instead of only attached to the
inner lining of digestive tract; zygospores generally spherical
in shape.
b. Trichomycetes : Mostly commensals with the guts of
arthropods; hyphae attached to inner lining of digestive
tract; rarely on external parts of aquatic living arthropods;
zygospores where known bipolar or biconical.
Hyphae are septate, vegetative body is
haplophase. It has five classes. This subdivision includes forms such
as yeasts, brown moulds, green moulds, pink moulds, cup fungi,
and edible morels. In this group of fungi asexual reproduction takes
place by various types of non-motile spores such as oidia,
chlamydospores and conidia. Sexual reproduction takes place by
means of gametangial copulation (yeasts). gametangial contact
(Penicillium) and by somatogamy (Morchella). The ascomycetes or
sac fungi are characterized by the development of spores

In advanced ascomycetes, groups of asci get aggregated to form

150 APSCHE : SEMESTER-I BOTANY TEXT BOOK


three types i.e., cleistothecium (closed and spherical ascocarps.
Eurotium Neurospora),
and apothecium (cup shaped ascocarps. eg. Peziza). This sub division
includes six classes namely-
a. Hemiascomycetes : Ascocarp and ascogenous hyphae
absent; thallus mycelial or yeast like; asci naked.
b. Loculoascomycetes : Ascocarp and ascogenous hyphae
present; thallus mycelia. Ascus bitunicate; ascocarp an
ascostroma.
c. Plectomycetes : Ascocarp and ascogenous hyphae present;
thallus mycelia. Ascus unitunicate; if bitunicate; then
ascocarp an apothecium. Asci scattered at various levels
within a cleistothecium; ascospores aseptate.
d. Laboulbeniomycetes : Ascocarp and ascogenous hyphae
present; thallus mycelia. Ascus unitunicate; if bitunicate;
then ascocarp an apothecium. Asci regularly arranged
forming a hymenium at the base or periphery of ascocarp.
Ectoparasitic on Arthropods; thallus reduced; ascocarp a
perithecium; ascus inoperculate.
e. Pyrenomycetes : Ascocarp and ascogenous hyphae present;
thallus mycelia. Ascus unitunicate; if bitunicate; then
ascocarp an apothecium. Asci regularly arranged forming a
hymenium at the base or periphery of ascocarp. Not
ectoparasitic on Arthropods. Ascocarp a perithecium; ascus
inoperculate.
f. Discomycetes : Ascocarp and ascogenous hyphae present;
thallus mycelia. Ascus unitunicate; if bitunicate; then
ascocarp an apothecium. Asci regularly arranged forming a
hymenium at the base or periphery of ascocarp. Not
ectoparasitic on Arthropods. Ascocarp an apothecium, often
massive; hypogeal or epigeal, ascus inoperculate or
operculate.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 151


D. Basidiomycotina:
vegetative body is dikaryophase. It includes the highly evolved
This group got its name from the basidium, the club shaped

mushrooms, toadstools, puffballs and bracket fungi . ‘The mycelia


of this group are of two types. Primary and secondary. Primary
mycelium multiplies by oidia, conidia like spores and pycnidiospores.
Distinct sex organs are absent. Fusion occurs between two
basidiospores or between two hyphal cells of primary mycelia.
Advanced forms of basidiomycetes produce fruiting bodies
basidiocarps.
microscopic to large ones. This sub division includes three classes
namely –
a. Teliomycetes : Basidiocarps absent, basidium arising from
thick walled probasidium, a teleutospore, teleutosori on host
tissue, parasitic on vascular plants.
b. Hymenomycetes : Well developed basidiocarp present
basidia arranged in a hymenium saprophytic , rarely
parasitic. Basidiocarp gymnocarpous orangiocarpous,
hymenium exposed throughout its development, i.e .,
Gymnocarpous, hymenium enclosed in the f irst bo dy, i.e.,
Angiocarpous
c. Gastromycetes : Well developed basidiocarp present
basidia arranged in a hymenium saprophytic , rarely
parasitic. Basidiocarp angiocarpous basidium holobasidium
E. Deuteromycotina: Three classes are included under this. They
are the so-called ‘ Fungi Imperfecti’. It is a group of fungi known
only from their asexual (imperfect or anamorphic) or mycelial state.

152 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Their sexual (perfect or teleomorphic) states are either unknown
or may possibly be lacking altogether. This sub division includes
three classes namely –
a. Blastomycetes: Without pseudomycelium or
pseudomycelium with yeast-like budding cells; true
mycelium absent or underdeveloped

b. Hyphomycetes : Mycelium developed assimilatory budding


cells absent. Sterile mycelium, spores borne on sporophores,
sporophores may be grouped together, but pycnia and
acervuli are not formed.

c. Coelomycetes : Mycelium developed assimilatory budding


cells absent. Spores or conidia formed in pycnidia or acervuli.

Know more:
 Deuteromycota is an informal group of unrelated fungi that
share a common character of asexual reproduction
 Ascomycota is characterized by the formation of an ascus,
a sac-like structure that contains haploid ascospores.
 Basidiomycota is characterized by showy fruiting bodies
that contain basidia in the form of clubs.

Activity: Take the different types of spores of fungi and draw a


table showing different classes of fungi that you have studied along
with examples.

Know more: The fruiting bodies of a basidiomycete form a ring in


a meadow commonly called “fairy ring.” The best-known fairy ring
fungus is Marasmius oreades. The mycelium is underground and
grows outward in a circle. As it grows, the mycelium depletes the
soil of nitrogen, causing the mycelia to grow away from the center
and leading to the “fairy ring” of fruiting bodies where there is
adequate soil nitrogen. (Credit: “Crop circles”/Wikipedia
Commons)]

APSCHE : SEMESTER-I BOTANY TEXT BOOK 153


Structure, reproduction and life history of
Chapter - 3.2 (a) Rhizopus (Zygomycota),
(b) Puccinia (Basidiomycota)

Learning outcomes: At the end of the chapter you should be


able to
1. Explain the structure, reproduction and life cycles in
Rhizopus and Puccinia.
2. Compare and contrast the features of fungi from two
different classes in the syllabus.

Kingdom : Fungi
Division : Mycota
Subdivision : Eumycotina
Class : Zygomycetes
Order : Mucorales
Family : Mucoraceae

3.2.1. Rhizopus :

Occurrence : Rhizopus is a type of fungus that belongs to the class


Zygomycetes, because of the production of zygospore in the sexual
reproductive phase. Rhizopus species are cosmopolitan in distribution.
Primarily Rhizopus species live on dead and decaying matter.
Majority of the Rhizopus species are saprobic and few are parasitic
viz. R. artocarp, R. arrhizus etc. It grows on a variety of substrates like fruits,
vegetables, bread, jellies etc. In 1820, Ehrenberg first used the term Rhizopus.
It is also called bread, black or pin mould. Nearly 8-10 species of Rhizopus
are reported. It differs in properties from other moulds by producing
sporangiospores instead of conidia. The main identifying feature of
the Rhizopus species is the presence of rhizoids at the base.

Mycelium of Rhizopus : The mycelial hyphae of Rhizopus species


are typically non-septate and white in colour. The hyphae

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differentiate into three distinctive parts namely stolons, rhizoids
and sporangiophores (Fig 3.2.1). Stolons and rhizoids are formed
by the differentiation of vegetative hyphae. Reproductive hyphae
formed into sporangiophores.

Mycelium is aseptate, coenocytic, tubular, multinucleated,


and consists of cell organellae like Golgi body, mitochondria,
vacuoles etc. (Fig.3.2.2) Cell wall is non-cellulosic and made of
chitin. Food material is reserved in the form of glycogen and oil
droplets. Moisture or water availability is necessary for the growth
of Rhizopus. Extra cellular digestion is present.

Stolons: It is the internodal region and in the form of a runner


which are aseptate and branched. It is the aerial hyphae that grow
horizontally, and it is found attached to the substratum.
Rhizoids: It is the nodal region that eventually forms by the contact
of stolon to the surface of the substratum. It shows much branched
structure. Its main function is to absorb all the nutrients from the
substratum.
Reproductive hyphae: Unbranched,elongated, and columellate
reproductive structures are called sporangiospores. These are stalked
hyphae that grow vertically from the stolon. Spherical or globose
sporangium with the columella attaches at the tip of a
sporangiophore.
Growth : The growth of mycelium appears rapid. It occurs as dense
cottony colony. Generally, the colour of the mycelium is white, and
after sporulation it turns into grey or golden brown.
Reproduction : Rhizopus reproduces by vegetative (fragmentation),
asexual (sporangiospores) and sexual (zygospores) methods.
Vegetative Reproduction : Sometimes the thallus of the Rhizopus
breaks accidentally or due to some other factors into the few

APSCHE : SEMESTER-I BOTANY TEXT BOOK 155



        
      





     

  


   
       
  



Fig. 3.2.1. A. Rhizopus on bread


B. Mycelium of Rhizopus

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Fig 3.2.2. A) Rhizopus hyphae B) Ultrastrucutre
Source:https://www.onlinebiologynotes.com/Rhizopus-stolonifer-morphology-and-
reproduction-of-black-bread-mold/

fragments, after which each fragment give rise to a new thallus on


favourable conditions.
Asexual Reproduction : The asexual reproduction can take place
through both sporangiospores within the sporangia and
chlamydospores. The vegetative hypha grows to form terminal,
aseptate sporangiophore that is long and slender. The nuclei and
cytoplasm push more and more towards apical side and
consequently the apex swells up and forms the swollen tip. After
that, differentiation of sporangia occurs that gives rise to the
formation of spore sac in between the sporangia and columella. A
septum is formed to separate into two regions. The swollen part
enlarges and form young sporangium. The cytoplasm of the
sporangium differentiates into the denser peripheral region with
more nuclei (sporoplasm) and the central columella region with few
nuclei and more vacuoles (columelloplasm). In the centre of the
protoplasm, vacuoles are arranged in semi crescent manner and
united to form swollen vegetative structure called columella which
is absolutely a sterile structure. It gives rise to the large and round
sporangium by pushing its cytoplasmic material to the peripheral
wall. Nucleus in the sporoplasm divides rapidly and later the

APSCHE : SEMESTER-I BOTANY TEXT BOOK 157


   
 

       



    

     

    

 
 
 

* * * * * * *
protoplasm which surrounds the columella begins to divide in small
multinucleate protoplasmic bits from outside to inside.
Each protoplasmic bit contains 5-10 nuclei and develops a
wall surrounding it to form a spore. Each spore is oval or triangular,
unicellular, multinucleate and non-motile, it is called sporangiospore
(aplanospore). On the maturation of sporangiospores the outer wall
of the sporangium bursts during the extreme conditions and spores
are dispersed and disseminated by wind. The sporangiospores
remain in dormant or resting state. On return of favourable
conditions these sporangiospores form germ tube or undergo
germination to develop new vegetative hyphae.

158 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Chlamydospores: During unfavorable condition like the old
mycelium becomes aseptate and the protoplasm of each cell becomes
rounded and thick walled by the accumulation of insufficient food
material and water. It is surrounded by a thick and nutrient rich
wall for perennating and to survive in adverse conditions. On getting
suitable conditions, when this chlamydospore gets moisture, it
forms a germ tube or undergoes germination process to develop a
new thallus. (Fig.3.2.3)

Fig 3.2.3 Rhizopus Asexual reproduction


Source: https://www.onlinebiologynotes.com/Rhizopus-stolonifer-morphology-and-
reproduction-of-black-bread-mold

Activity: Collect the molds from vegetable market and record the
morphological characters by observing under microscope. Identify
them to which class they belong.

Sexual reproduction : In Rhizopus, sexual reproduction occurs by


the method of gametangial copulation. Sexual reproduction takes
place during unfavourable condition by means of gametangial
copulation. Rhizopusis either heterothallic (Rhizopusstolonifer) or

APSCHE : SEMESTER-I BOTANY TEXT BOOK 159


homothallic (Rhizopus sexualis). The (+) and (-) thalli come in contact
with each other. They grow towards each other in opposite direction
to form protuberances or outgrowths. These are called
progametangium and the hyphae are called zygophores. The apical
regions of two progametangia come in close contact with each other
and cytoplasm of each progametangium push more and more
towards the tips and swell up with dense cytoplasm. Now the apical
region separated by septa and the apical region is called gametangia
and the basal region is called suspensor.

Fig 3.2.4 Rhizopus Sexual Reproduction


Source: https://www.onlinebiologynotes.com/Rhizopus-stolonifer-morphology-and-
reproduction-of-black-bread-mold/

Each gametangia contain dense granular multinucleate protoplast


called as aplanogamete or coenogamete. Now the wall of the
coenogamete/ gametangia at the point of contact dissolve which is
called plasmogamy and the nuclei of opposite strains paired and
fused to form diploid nuclei called karyogamy and ultimately forms
the diploid zygospore. Then, karyogamy occurs by which the

160 APSCHE : SEMESTER-I BOTANY TEXT BOOK


gametes of both the (+) and (-) thallus fuse with each other to form
a zygote. Zygospore enlarges and become thick walled to resist
adverse environmental conditions. It is a resting spore and
germinates during favourable conditions. There are two layers,
namely an outer layer which is ornamented and inner layer which
is smooth that surrounds the zygospore. The inner wall of zygospore
develops into promycelium and consists of germsporangiophore and
germ sporangium. Reduction division occur in germ sporangium
and contain haploid nuclei. Cytoplasm gathered to behave like a
spore. Each haploid spore is released and germinate to give rise to
new mycelium. (Fig. 3.2.4)

Know more: R. oryzae is used in the production of cortisone


and lactic acid which are used for biosorption of heavy metals in
waste water.

R.stolonifer produce lactic acid, fumaric acid and cortisone. It


also act as parasite causing soft rot diseases in fruits and
vegetables and therefore preventive fungicides are often sprayed

Rhizopus is used in the preparation of tempeh, a soybean ‘curd’


food consisting of crushed soybeans partially decomposed by
Rhizopus and held together by fungal hyphae.

Know more: A group of molds called mucorales causes a serious


rare fungal infection called mucormycosis, formerly called
zygomycosis. Mucormycosis is not a contagious disease and cannot
be spread from one person to another. These molds seriously threat
people who have other health problems or take medicines that lower
the body’s ability to fight germs and sickness.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 161


Life cycle : Rhizopus shows haplontic life cycle like many other fungi.
and some of the green algae. In this, the multinucleate filamentous

reproduction, it forms a promycelium consisting of germ-


sporangiophore and germ-sporangium. The nuclei of germ-
sporangium undergo meiosis to produce haploid spores (n). Each
haploid spore is released and germinate to give rise to new mycelium
(Fig.3.2.5).

Fig 3.2.5 Rhizopus Life cycle


Source: https://www.brainkart.com/article/Rhizopus—Fungi_32856/

162 APSCHE : SEMESTER-I BOTANY TEXT BOOK


3.2.2. Puccinia (Basidiomycota)

Kingdom : Fungi
Division : Basidiomycota
Class : Basidiomycetes
Order : Uredinales
Family : Pucciniaceae

Puccinia consists of more than 4000 different species, many of which


are plant pathogens. They cause several types of rusts in both
monocotyledons and dicotyledons. These are called stem rusts (P.
graminis), brown rusts (P. recondita), guava rust (P. psidii) etc.
Infection of P. graminis results in retarded growth and reduced seed
size in wheat. In south India, it appears at the end of November
while in north India, it appears in March.

Know more : Puccinia graminis is an obligate parasite on a wide


variety of hosts and commonly effects wheat plant. It also effects
many other species of grasses such as oat, barley, rye etc.

Structure of the mycelium: The vegetative mycelium consists of


branched hyphae, which are septate with simple pores. It is inter-
cellular in nature, grows between the cells of the host. The haustoria
produced by this parasitic fungus penetrate through the host cell
and absorb nutrients. The mycelium is of two types i.e.,
monokaryotic and dikaryotic based on nuclear behavior. (Fig.
3.2.6)

Dikaryotic mycelium: It is found in primary host, wheat. Each


cell of this mycelium consists of two nuclei belong to two opposite
strains (+ and -). It develops from dikaryotic aeciospores produced
on secondary host (Barberry), or dikaryotic urediniospores from

APSCHE : SEMESTER-I BOTANY TEXT BOOK 163




   



       
      
  

   
         
 
   

      
 

 


Fig. 3.2.6 Monokaryotic and Dikaryotic mycelia


Source: https://www.slideshare.net/vivekaiden/structure-and-reproduction-of-puccnia-and-
fuserium

164 APSCHE : SEMESTER-I BOTANY TEXT BOOK


primary host itself. It again produces urediniospores in uredinia
and teliospores in telia on entering the reproductive phase.).

Monokaryotic mycelium: This mycelium is seen in secondary


(alternate), barberry. It develops by the germination of haploid
basidiospore on the leaf of host plant. Each cell in this mycelium
consists of a single nucleus.

Dikaryotisation: It occurs in the receptive hyphae of flask shaped


spermagonia (pycnidia) that develop on the upper surface of the
alternate host, when haploid hyphae of two opposite strains (+ and
-) come in contact.

Find: Pucciniais an obligateparasite and cangrow on living cells of


suitable hosts. Explain different species of Puccinia that attacks
on different plants causing rusts

Reproduction: Puccinia species are heteroecious and require two


hosts to complete its life cycle. For instance, Puccinia graminis tritici
is parasitic on wheat (Triticum aestivum) and the barberry (Berberis
vulgaris) as secondary or alternate host. Hence, it reproduces and
completes its life cycle on those two hosts. The dikaryotic mycelium
of Puccinia produces the following spore stages on the leaves and
stems of wheat.

Uredial stage and urediniospores: The uredinia are pustules that


produce urediniospores. They are elongated brick red coloured
structures formed between the veins of leaves and stems.
Aeciospores liberated from infected barberry plant causes infection
on healthy wheat plants by germination. An aeciospore produces a
germ tube and enter the wheat leaf or stem tissue through the
stomata. The dikaryotic mycelium aggregate below the upper
epidermis. It produces vertically growing slender stalk like hyphae

APSCHE : SEMESTER-I BOTANY TEXT BOOK 165


        
 

    


 
     

  
  
     


   
   

    
       
        
 
        


      
 
  

   
166 APSCHE : SEMESTER-I BOTANY TEXT BOOK
arranged in a palisade-like layer and form sori called uredinia. The
tip of each hypha in a uredinium swells to produce a single binucleate
spore called urediniospore.

The urediniospores are formed in groups after a week of


infection, burst through the host epidermis. A single uredinium may
contain nearly 50,000 to 4,00,000 urediniospores. Each
urediniospore is a stalked structure bearing a swollen oval body at
its tip. Its body is surrounded by spiny wall consisting of four germ
pores. Uredeniospores spread through air, on falling upon the leaves
of wheat plant germinate within a few hours. Air borne
urediniospores cause heavy and repeated infection on wheat crop.
Hence this stage of Puccinia is also called the repeating stage (Fig
3.2.7).
Telial stage and teliospores: The telia are pustule like sori
producing teliospores. They appear as black streaks along leaf
sheaths and stems of infected plants. Puccinia produces teliospores
instead of urediniospores either in the same sorus or in a new sorus,
when wheat crop about to reach harvesting stage late in the growing
season. Gradually more teliospores are produced and the number
of urediniospores is reduced. Finally, the sori are left with only
teleutospores. They exert pressure on the epidermis and rupture
to expose the teliospores. The telial stage is known as the perfect
stage because karyogamy and meiosis takes place in teliospores. By
the time the crop matures, the leaves become dry and the black
pustules appear more prominent on the stem. Hence this disease is
also called as black stem rust.
Teliospores are spindle shaped, stalked and two celled
structures. The wall of teliospore is smooth and thick. A single germ
pore is present at the apex of the upper cell and below the septum
in lower cell. A teliospore consists of two nuclei i.e., one ‘+’ strain

APSCHE : SEMESTER-I BOTANY TEXT BOOK 167


Fig3.2.7. Stage I Uredinia on wheat leaf


       
   
      
   
   


Fig.3.2.8. A. Telia on wheat. Fig.3.2.8. B. A telium bearing teliospores


Source: https://www.plantscience4u.com/ Source:https://www.alamy.com/elementary-
2016/11/5-stages-in-life-cycle-of-Puccinia.html botany-botany-fig-157

168 APSCHE : SEMESTER-I BOTANY TEXT BOOK


and one ‘-’ strain. On maturation of the spore, karyogamy of two
nuclei in each cell results in the formation of a diploid nucleus. These
teliospores remain dormant on the straw and stubble of harvested
crops. Under favorable conditions like high atmospheric humidity,
moisture and low temperature; after a resting period they germinate
in situ to produce basidiospores (Fig 3.2.8 A & B).
Basidiospores: These spores represent the stage IV of the fungal
pathogen. Each cell of teliospore that contains a diploid nucleus is
considered as a probasidium. During germination of a teliospore, a
short hypha of limited growth called as promycelium (epibasidium
or metabasidium) grows out through germ pore of each cell. The
diploid nucleus in each cell undergoes meiosis and four haploid
nuclei are produced, which migrate into promycelium. Septa are
formed between the nuclei dividing the epibasidium into four
uninucleate haploid cells. Each cell of the promycelium gives out a
sterigmata, and one basidiospore is formed at the tip of each
sterigmata. Of the 4 basidiospores formed from one cell of a
teliospore, two are of ‘+’ strain and other two are of ‘-’ strain. The
uninucleate haploid basidiospores do not infect wheat. These are
carried away by the wind to the alternate host, barberry and
germinate on its leaves (Fig 3.2.9A & B).

Fig.3.2.9. A. Germinatingteliosporeandformationofbasidiospores

APSCHE : SEMESTER-I BOTANY TEXT BOOK 169


Fig:3.2.9. B. Formation of basidiospores
Source:http://go.microsoft.com/fwlink/p/?LinkId=255141

Spore stages on barberry plant: The basidiospores when fall on


the upper surface of barberry leaf germinating immediately. During
germination, a germ tube has given out which penetrates through
the epidermis. The germ tube elongates, divides inside the leaf of
alternate host developing into hyphae. Those hyphae grow between
cells lying in between both lower and upper epidermal layers. The
hyphae consists of uninucleate haploid cells (primary mycelium) of
‘+’ or ‘-’ strain. Several basidiospores of opposite strains may infect
the same berberis leaf. Thus, haploid mycelia of two different strains
(+ or -) are formed. They remain haploid for some time and the
fusion between the hyphae of two strains (‘+’ or ‘-’) may occur at a
later stage.

Spermogonial (Pycnidial) stage: After about four days of the


infection on barberry plant, the haplo-mycelium collects and forms
dense mats beneath upper and lower epidermal layers. The mycelial
mat beneath the upper epidermis trasforms into primordium of

170 APSCHE : SEMESTER-I BOTANY TEXT BOOK


spermogonium; while the mat beneath the lower epidermis
develops into primordium of aecidium (protoaecidium). After 7 to
10 days of infection, each primordium of spermogonia matures into
a small flask shaped spermogonium or pycnidium. The minute
yellowish specks of pycnidia appear on the dorsal surface of the
leaf. Each spermogonium opens on the upper surface of the host
leaf through a pore like structure, ostiole. The wall of spermogonium
consists of three kinds of hyphae namely periphysis, receptive
hyphae and spermatiophores. Periphysis are the long, delicate and
sterile hyphae develop from the spermogonial wall and appear as
vertical projections at the fringe the ostiole. Receptive (Flexuous)
hyphae arise from the lateral wall of the spermogonium. They are
slender, delicate, cylindrical, septate, simple, branched or un-
branched with blunt ends. They are present in between periphysis
and sometimes it is difficult to distinguish those two.
Spermatiophores (Pycnidiophores) are slender, short, vertical and
uninucleate hyphae that arise from the base of the spermogonium.
Each spermatiophore produces several small uninucleate spermatia
(pycnidiospores) at its tip by abstraction method. The spermatia
are small, unicellular, hyaline, oval to spherical and smooth walled
structures. They fill the spermogonial cavity and are exuded from
the ostiole in a droplet of liquid, sticky, thick and sweet nectar.

According to Craigie and Butler (1927), the spermatia


function as male cell while receptive hyphae represent the female
sex organs. The spermatia may be ‘+’ or ‘-’ in their sexual nature
depending on the mycelium produced by the basidiospores. Insects
are attracted by nectar, help in dispersal of spermatia from one
spermogonium to another on the same leaf or different leaves; which
results in the transfer of ‘-’ spermatia to ‘+’ receptive hyphae or vice-
versa. Now the spermatization takes place when the spermatia of +

APSCHE : SEMESTER-I BOTANY TEXT BOOK 171


or - strain come in contact with the tip of the receptive hyphae of
opposite strain. The intervening wall of spermatium and receptive
hypha dissolves at their point of contact. Eventually, the nucleus of
spermatium passes downwards through septal pore and form a
binucleate cell. This pairing of nuclei of opposite strains is called
dikaryotization (Fig 3.2.10).

Fig. 3.2.10 A) Section of infected Berberis Leaf showing upper Pycnia and Lower
aecidium, B. Stages of Spermatization, two aeciospores showing disjunctor cells
Source: http://go.microsoft.com/fwlink/p/?LinkId=255141

Aecidial (aecial) Stage: The haplomycelium forms primordium


of aecidium (protoaecidium), which develops into aecidium after
the process of dikaryotization. The spermatial (male) nucleus forms
a second male nucleus by mitotic division, which moves to the next
cell through septal perforation. All the cells of primary mycelium
are dikaryotized when the male nuclei produced by successive
mitotic divisions pass down. The sporophores are formed when
dikaryotic basal cells of the protoaecidium arrange themselves
vertically above the lower epidermis. Each binucleate basal cell cuts
off a chain of binucleate cells called aecidiospore mother cells in
basipetal succession towards the lower epidermis of the host. These
cells further divide transversely to form a large cell, aecidiospore

172 APSCHE : SEMESTER-I BOTANY TEXT BOOK


and a small sterile cell, disjunctor (intercalary cell). The latter
dissolves to set free the aeciospores. Some of the basal cells lying at
the periphery of protoaecidium mature into a one celled thick
protective layer called peridium along with the development of the
aeciospores. This entire cup shaped structure is known as aecium.
The developing aeciospores exerts pressure and rupture the
peridium to liberate aeciospores. Each aeciospore is an orange yellow
coloured, unicellular, thin walled, polyhedral, and binucleate
structure (Fig 3.2.11).

Fig:3.2.11 Section through Aecium showing aeciospores


Source:https://4.bp.blogspot.com/-rI5E3rljWL0/XBx3p-0CSfI/AAAAAAAAAmQ/
vALaOJ7J2AkGwO-j5pYXisaN4G6qPZfmwCLcBGAs/s1600/6.%2BAeciospores.jpg

Aeciospores are capable of immediate germination, but


cannot infect barberry plants. The aeciospores are disseminated by
wind and germinate on wheat leaf by producing germ tubes (primary
hyphae). It forms the dikaryotic mycelium ultimately when further
development of the germ tube takes place in the uredinal stage.
This mycelium produces uredospores and later teleutospores on
wheat. Thus, Puccinia completes its life cycle on two different hosts
and by producing 5 types of spores. (Fig 3.2.12A&B).

Know more: In some Pucciniaspecies one or two spore stages are


missed in its life cycle and are known asmicrocyclic.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 173


Fig 3.2.12 Symptoms on wheat plant A) Uredial stage and B) Telial Stage

Life Cycle of Puccinia: The life cycle of Puccinia graminis tritici is


called as heteroecious and macrocyclic; as the fungus requires 2 hosts
and comprises of five different types of spores produced in five
sequential stages (0, I, II, III and IV). They are stage - 0 (Spermatia)
and stage-I (Aeciospores) on secondary host, Berberis vulgaris (Fig
3.2.13); Stage II (Urediniospores), III (Teleutospores) and IV
(Basidiospores) on primary host, Triticum aestivum (Fig 3.2.14).

Fig 3.2.13 Symptoms on Barberry Plant


Source:http://go.microsoft.com/fwlink/p/?LinkId=255141

Self -analysis: What are the differences in the spores of Puccinia?


Discuss the following points: Macrocyclic rust, heteroecious
rust, Dikaryotization, sexual reproduction and Types of spores in
the life cycle of Puccinia

174 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig.3.2.14 A A Life cycle of Puccinia graminis tritici
Source: http://go.microsoft.com/fwlink/p/?LinkId=255141

Fig.3.2.14 B Life cycle of Puccinia graminis tritici


Source: http://go.microsoft.com/fwlink/p/?LinkId=255141

APSCHE : SEMESTER-I BOTANY TEXT BOOK 175


Know more : Some important Rust Diseases of crop plants
S.no Causalorganism Disease Crop
1 P.graminishordei StemRust Barley
2 P.purpurea Rust Sorghum
3 P.penniseti LeafRust Bajra
4 P.helianthii Rust Sunflower
5 P.carthami Rust Safflower
6 P.arachidis Rust Groundnut
7 P.alli Rust Onions

LIFE CYCLE OF PUCCINIA


Host Growth Stage Characteristic feature
Wheat Uredo Stage Formation of single celled, binucleated
uredospores on stalk. Uredospores form germ
tube that apply pressure on epidermis breaking
it and forming uredinia. Uredospore germinate
and forms mycelia.
Telial Stage Teleutospores formed from uredospore in
(Black stage) unfavourable condition of growth. Each cell
has single germ pore and two nuclei.
Kar yogamy leads to formation of diploid
nucleus. Spores rest until favourable condition.
Basidial Stage Formation of haploid basidiospores by meiosis.
Ejection of spores that are carried out by wind
to secondary host. Infects barberry forming
haploid mycelia.
Barberry Pycnidial Stage Formation of specialised Pycnia or spermatia.
Pycnia produces haploid pycniospores and
receptive hyphae. Dispersed by insects,
Pycniospores fertilise receptive hyphae of
another plant forming dikaryotic mycelia.
Aecial stage Formation of dikar yotic aeciospores from
mycelia. Chain like aeciospores are carried by
wind to infect cereal host.

176 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Chapter -3.3 Economic uses of fungi in food industry,
pharmacy and agriculture.

Learning outcomes : At the end of the chapter you should be


able to to:

1. Explain the application of fungi in industrial,


pharmaceutical and agricultural fields.
2. Discuss the significance of fungal products and processes
in human welfare.

Fungi are used as foodstuffs. Many mushrooms are edible and


served in dishes. The fungi, like yeasts are responsible for
fermentation. Fungi influence the well-being of human
populations on a large scale because they are part of the nutrient
cycle in ecosystems. They show an important role in nutrient
recycling of the ecosystem as well. Fungi help to control the
population of damaging pests having the capacity of animal
pathogens. They are very specific to the insects they attack, and
do not infect other animals or plants. Currently in fungi lot of
investigation has been carrying out in search of potential microbial
insecticides, though several are already available in the market.
Many saprophytic fungi cause the sufficient damage and decay to
the food stuff, the paper wood pulp and gives a death blow to the
paper industry. They are vitally important to human life on many
levels, although we often think of fungi as organisms that cause
disease and rot food.

Can you imagine without the fungal partner in root systems, how
trees and grasses would survive?
Do you know how Mycorrhizal fungal inoculants are available as
soil amendments for farming?
Do you know gardening supply stores for the promotion and act as
supporters of organic agriculture?

APSCHE : SEMESTER-I BOTANY TEXT BOOK 177



 
 


   
     
      




   
        
       
         
   

 
       
   

 
   

    

178 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fungi exist everywhere in the environment and they are
important for human beings. Some of them are useful as food and
some form the basis of medication. They have been making
important contributions in managing diseases in humans and
animals. Other fungi such as mold are less desirable and spores
cause diseases. Discoveries of penicillin by Alexander Fleming and
the other antibiotics greatly changed the medical world for the
betterment of health. Ongoing research continues in several areas
to find ways that fungi can help for human welfare. Fungi serve as
both useful and harmful in many activities.
I. Beneficial effects of fungi:
A. Fungi in food industry: Fungi are used in many important
food industries to produce a variety of products.
a. Edible fungi: Some fungi are delicious in taste are used as food.
Most popular fungi as food are mushroom and morels. They have
very high percentage of protein either equal or more than animal
dietary protein. They are also rich source of vitamins. The common
field mushroom (Agaricus campestris) morel (Morchella esculenta),
Catherella cibarius, puff balls (Lycoprdon and Clavatia) and Boletus
edulis are the most important edible fungi.
b. Cheese Production: Cheese is the product formed by controlled
process when water is removed from milk. Cheese making involves
the following steps:
 From milk casein protein has been coagulated and then
separating the milk into solid curds and liquid whey.
 Drained away liquid whey, the curds are salted, shaped and
left to ripen in a controlled environment.
Penicillium
They provide a characteristic texture and flavour to the cheese.
There are two types of mould refined chesses. One is Camembert

APSCHE : SEMESTER-I BOTANY TEXT BOOK 179


 

    


********
and Brie types (soft cheese) and the second is Roquefort Gorgonzola
and Stilton cheese. These cheeses are made from two species of
Penicillium, P. camemberti, in Camembert cheese and P. roqueforti in
Roquefort cheese. These cheeses are among the favorites among
gourmets.
Find: Identify different types of Mushrooms, their uses and
products

c. Brewing and fermentation: In the late 1850s’ for the brewing


industry, the denoting discovery of Louis Pasteur led to the
Saccharomyces
cerevisiae. The common alcoholic beverages wine and beer are
products of fermentation by the activity of different species of
Saccharomyces. Most of the human cultures practiced for millennia
to ferment grains to produce beer, and fruits to produce wine.

Product Substrate for Fermentation % of Alcohol

Wine juice of grapes or other fruits 10-12 % alcohol

Beer malting of the starch or barley grains 3 – 8 % alcohol

Whisky fermented mash of cereals or potatoes 50% alcohol

Brandy distilled from wine 60 –70 % alcohol

Rum distillation of fermented sugarcane 40% alcohol


juice or molasses

180 APSCHE : SEMESTER-I BOTANY TEXT BOOK


FACTS: During the process of fermentation process a large amount
of CO2 is liberated which is used by the green plants for the synthesis
of food by photosynthesis.

Know more: Wild yeasts are acquired from the environment and
used to ferment sugars into CO2 and ethyl alcohol under anaerobic
conditions. Yeast cake in the large-scale production is called
‘microbial farming’. It is rich in protein and hence the yeast cakes
are eaten directly as health giving food. Yeast cakes are extensively
used in baking and brewing industries. In this, large number of
yeast cells with some inert substances are mixed such as, starch
and then compressed to form cakes. They are also used in
laboratories to demonstrate fermentation in sugary solution.

B. Fungi in pharmacy: Antibiotics are naturally produced by fungi


to kill or inhibit the growth of bacteria limiting their competition
in the natural environment. Important antibiotics such as penicillin
and the cephalosporins are isolated from fungi. Many secondary
metabolites of fungi are of great commercial importance and are
used as valuable drugs.
a. Drugs: Cyclosporine, an immunosuppressant drug isolated from
Cyclospora sp. It reduces the risk of rejection after organ transplant
and also act as the precursors of steroid hormones. Psilocybin
extracted from Psilocybe semilanceata Gymnopilus junonius have
hallucinogenic properties and are used by folkore people in their
cultural festivals
b. Enzymes: Various fungi have been used in the preparation of
many important and useful enzymes. Saccharoymces cerevisiae
produces a sucrose hydrolyzing enzyme known as Invertase and
helps to hydrolyze the sucrose into the mixture of glucose and
fructose. Many of them are commercially available Digestin,

APSCHE : SEMESTER-I BOTANY TEXT BOOK 181


Polyzime, Taka diastase, etc. are used in starch dextrinization and
desiring of textiles. Amylase is produced by the Aspergillus niger and
A. oryzae, which contains two starch splitting components.
Tyrosinase is obtained from Neurospora crania.

ANTIBIOTICS SOURCES USES


Penicillin Penicillium Used clinically for tonsillitis, tetanus,
notatum pneumonia, rheumatic fever,diphtheria
and many other diseases.
Griseofulvin Penicillium Used against my cosis
griseofulvum
Cephalosporin Cephalosporium Used against Gram +ve and Gram –ve
acremonium bacteria, typhoid
Clavicin Aspergillus Used against fungal diseases
clavatus

c. Vitamins: Fungi are rich source of many vitamins. Vitamins B


complex is obtained from yeasts. Other vitamins obtained from
yeasts and moulds are vitamin D, Ergosterol, Riboflavin, etc.
d. Antibiotics: Several antibiotics have been obtained from fungi.
Their name, sources and applications are listed below in table.

C. Fungi in Agriculture: Fungi are widely used as natural agents


in agriculture next to bacteria. They are useful to the agriculture in
several ways.
a. Decomposition of death and decay material: Many
saprophytic fungi decompose plant and animal debris (organic
waste) result in the formation of humus, which is needed for the
growth of plants as natural fertilizer in the form of compost and
litter. Because of the ability to produce a wide variety of extracellular
enzymes, they are able to break down all kinds of organic matter,
decomposing soil components and thereby regulating the balance
of carbon and nutrients to maintain soil health.

182 APSCHE : SEMESTER-I BOTANY TEXT BOOK


b. Biofertilizers: Soil is a primary source of fungal growth, and
produce a wide range of bioactive metabolites for improvement of
plant growth. Fungi are used as biofertilizers as they supply
inorganic nutrients to plants such as ammonium, nitrate, and
phosphate.
The mycorrhizal relationship between fungi and plant roots
plays essential role in the productivity of farm land. The plant
obtains mainly phosphate and other minerals such as zinc and
copper from the soil. The fungus in turn obtains nutrients such as
sugars from the plant root. This mutual beneficial relationship is
called mycorrhiza. These are used to bridge gaps in the soil to
transport nutrients relatively far distances back to the plants.

Fungal Plant type Fungi-mediated Beneficial effects on


species/strain response plant species
AM fungi Dead vegetation Degrade of dead Nutrient mobilization
in soil organic matter
Phanerochaete Wood Decomposing wood Phosphorus
velutina translocation
Pleurotus sp. Wood Wood decay Nutrient mobilization
Trichoderma sp. Arabidopsis sp. Auxins dependent Higher biomass
mechanism production and
increased lateral roots
formation
Ectomycorrhizal Higher plant Phenolic Plant protection
fungi species compounds
degradation
Ectomycorrhizal Agricultural Stomatal physiology Improved water
fung i and AM crops and water relation potential status and
fungi increased
photosynthesis rate

Soil-beneficial fungi on different physiological and catabolic


processes in various host plant species is as follows. Gibberella
fujikuroi produces a growth hormone known as Gibberrelin which
acceleratesplantgrowth.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 183


Product Microorganism Used Agriculture application
Gibberelins Fusarium monoliformae Plant growth hormone
Zeralenone Fusarium graminearum Growth promoter in cattle
De Vine Phytophthora palmivora Control of milk weed vine
Chontral Chondrostereum purpureum Control of hard woods
Rotstop Phanerochaete gigantea Control of butt rot of conifers

b. Biological control: Some of Fungi used as biocontrol agents


against pathogenic bacteria and fungi, insects and pests
(Biopesticide), and namatocides. They are safe for other non-target
organisms and infect only specific species, with long-term results
on target pests. These can infect a wide range of insect hosts.
Ascomycetes are fungi used for the control of insect pests usually
found in the soil and can cause natural outbreaks on their own when
environmental conditions are favourable. The entomopathogenic
fungi have the capacity to reduce or eradicate target insect groups
such as Coleoptera, Diptera, Hemiptera, Hymenoptera, Lepidoptera
and Orthoptera. Some fungi such as Empusa sepulchrasis,
Metarrhizium anisopliae, Cordyceps melothac etc. are used to control
the insect pests.
Some fungal inhabitants of the soil help to fight against
disease caused by the soil borne fungi. For example, Trichoderma
lignorum and Gliocladium fimbriatum inhibit the growth of the
mycelium of Pythium. They suppress the causative agent of damping
off disease of the seedlings, and promote the growth of plants. There
are other fungi which are used against soil borne fungal pathogen
such as Trichoderma viride and T. harzianum.

Activity: Collect the soil samples from the rhizosphere of your


surrounding farms and prepare the fungal cultures. Observe under
microscope for beneficial and harmful fungi

184 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Know more: Nematocides are
predacious fungi trap and destroy the
nematodes (eel worms). They form loops
on the mycelium (A) which act as nooses.
When nematodes try to pass through
this loop they are trapped. After that
mycelium absorbs nutrition from the
captive by sending a special hypha.
Predacious fungi also produce sticky
conidia. When nematodes pass through
the soil, conidia stick to their bodies.
After those conidia germinates and
produces hyphae which then penetrate
into the tissues of the host and absorb
Fig. 3.3.3. Control of Nematodes
by Nematophagous Fungi nourishment. Eg: Beauveria bassiana is
Source: http://go.microsoft.com/ used as a nematicide against borers,
fwlink/p/?LinkId=255141
thrips, and aphids.(Fig. 3.3.3)

II. Harmful activities of fungi:

Apart from the beneficial effects, Fungi has harmful effects as


mentioned below
a. Toadstools: Certain mushrooms are poisonous ones and called
as toadstools. Some of the mushrooms like Amantia muscaris, Boletus
satanas, Entoloma lividium and Russula spp are poisonous. They are
popularly called “death cups” or “destroying angels” as they are fatal
and cause death.
b. Spoilage of food stuffs: Some fungi cause spoilage in food
stuffs. Citrus fruits are spoiled by Penicillium digitatum, Mucor,
Aspergillus, Penicillium, Oidium and Fusarium responsible for the
spoilage of milk and milk products. Aspergillus responsible for bread
spoilage. Oidium lactis responsible for fishy odour of butter. Mucor

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sp., Penicillium, Neurospora, Fusarium, Aspergillus etc. are responsible
for the spoilage of meat.
c. Diseases: In agriculture causing various diseases on different
crops like damping off, wilts, rusts, smuts etc.
d.Aflatoxins: Some fungi such as Aspergillus flavus, A. fumigatus,
A. parasiticus and Penicillium islandicum produce a most potent
carcinogenic agent called Aflatoxins on dried foods and groundnut
meal. Aflatoxin prevents the transcription by binding with the DNA.
They cause liver cancer in animals and human beings.
e. Ergot: It is the most useful drug obtained from sclerotia of
Claviceps purpurea. The fungus is parasitic on rye grass and cause
disease called ‘ergot of rye’. The ergot contains alkaloid ergotinine
and many other substances like ergotonic acid, cornutine,
spacelotoxin, etc. The drug stimulates muscles of uterus and used
as abortifacient to assist child birth. It also used to stop bleeding
and in many other uterine disturbances.

Know more :

a. Death caps: The poisonous mushrooms are called “death


cups” or “destroying angels” as they are fatal and cause death.
e.g., Amantia muscaris, Boletus satanas, Entoloma lividium and
Russula spp.
b. Brewer’s yeast: Saccharomyces cerevisiae is commonly
called as Brewer’s yeast It is used in common alcoholic
beverages such as wine and beer the products of fermentation.
c. Roquefort or Gorgonzola and Stilton cheese: It is named
after its origin in the natural environment such as the caves
of Roquefort, France, in which Penicillium roqueforti is the
principal mold. Sheep milk cheese is stacked in order to
capture the moulds responsible for the blue veins and pungent
taste to the cheese

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A general account on symptoms of plant diseases
Chapter -3.4
caused by Fungi & Blast of Rice

Learning Outcomes : At the end of the Chapter you should be


able to :
· Describe the symptoms of plant diseases caused by fungi
· Summarize the various diseases caused by fungi
· Describe the Blast of rice, disease cycle and control

Introduction: Economically useful plants which are grown for food


and other commercial purposes are attacked by many different
diseases. They are caused by different pathogens like bacteria and
virus. Similarly, many crop, fruit and other economically useful
plants are also attacked by Fungi and many diseases are affected.
These diseases cause for tremendous economic losses.
Fungi can cause localized or general signs and/or symptoms.
Fungal infections cause general necrosis of host tissue and often
cause stunting, distortions and abnormal changes in plant tissue
and organs in the majority of cases. The most distinctive and easily
identifiable characteristics of fungal infections are the physical
presence of signs of the pathogen. Significant clues for proper
identification and diagnosis of a disease include signs in hyphae,
mycelia, fruiting bodies and spores of the fungal pathogens.
Microscopic to macroscopic fruiting bodies of fungi are there. They
possess many shapes and configurations with individual
characteristics. Accurate identification of the disease will be done
by the fruiting bodies, along with spores, and mycelium. The
symptoms like wilts, rusts, spots, smuts and rot are common in
fungal infections whether alone or in combination with other fungal
pathogens. “Bulls-eye-like” typical fungal leaf spot are seen
consisting of roughly concentric rings that may display zones of

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


    

   

        







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Fig 3.4.1 Symptoms of different types of fungal diseases in different host plants.

different yellow, red or purple colours. The spots are not restricted
by the leaf veins as can be seen in bacterial leaf spots.

3.4.1 Symptoms of plant Diseases caused by fungi:

A.Damping off disease: A rapid collapse and death of young


seedling due to either the seed rots before emergence or the seedling
rots at the soil line. It falls and dies. Several soil-born fungi cause
this disease. This disease is caused by a species of Pythium,
Rhizoctonia and Fusarium. It has found in commercial crops like
tomatoes, cotton, mustard, peas, beans, tobacco etc (Fig. 3.4.1.a).
B.Blights: It can happen because of both biotic and abiotic
disorders. Rapid generalized browning and death of leaves, floral
organs, stems and branches are seen. Cherry Brown Blight (Monilinia
fructcola and M. laxa) ,Tomato Early Blight ( Alternaria solani) and
Tomato and Potato Late Blight (Phytophthora infastans) are some of
the examples of fungal blight.. This disease causes great damage to
the potato tubers. In 1845, in Ireland and destroyed the entire
potato crop. Egg plants, tomatoes, etc also will be infected (Fig.
3.4.1b).
C. Dieback: It is progressive death of shoots and twigs generally
starting at the tip of the infected plant.Eg.Brown Rot of Cherry
(Monilinia sp.) causes Shoot Dieback of Apple and Poplar Shoot
Dieback (Venturia populina) (Fig.3.4.1c)

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  

       
 

* * * * * * * * *

D. Downy mildews: White, grey, bluish, or violet downy patches


of mildew form mostly on the undersides of leaves in damp weather.
Affected leaves often wilt, wither, and die early.It is caused by
members of the oomycete genus Sclerospora, but other pathogens
include species of Bremia, Peronospora, Phytophthora, Plasmopara, and
Pseudoperonospora. In France, Downy mildews of grapes caused by
Plasmopora viticola destroyed all the vine yards and caused heavy
losses to the crop. Bordeaux mixture is used in treatment of the
disease (3.4.1d).
E. Scab: Scab characterized by crustaceous lesions on fruits, tubers,
leaves, or stems. It often found in apples, peaches, crabapples,
cucumbers, pecans, cereals, and potatoes.Apple scab is caused by
the fungus Venturia inaequalis (3.4.1e).
F. Smut: Smut is characterized by fungal spores accumulated in
soot like masses called sori. They are formed within blisters in leaves,
stems, seeds, flower parts, and bulbs. It reduces the yield and quality
of grain of corn, wheat, oat and other cereal crops, corn
smut(Ustilago maydis), Sorghum (covered kernel) smut (Sporisorium
sorghi).
G.Wilt: Disease fungi generally enter through the roots and
interfere with the water conducting strands of the plant. As the
infection spreads up into the stems and leaves it restricts water flow
causing the foliage to wilt and turn yellow. It is caused by Fusarium

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oxysporum. Fusarium wilt is a soil-borne pathogen that attacks
potato, tomato, eggplant and pepper plants.
E. Ergot disease of rye: It forms a poisonous sclerotia in the rye
kernel which is known as ergot of rye by Claviceps purpurea. It is
found in cereal crops especially in rye. Ergot is highly poisonous to
man which can cause hallucinations, insanity and finally death.
F.Galls: Enlarged parts of plant organs, usually caused by excessive
multiplication or enlargement of plant cells.Camellia Leaf Gall
(Exobasidium camelliae),Plum and Prune Black knot (Apiosporina
morbosa),Enlarged roots that look like clubs (spindles) e.g. Clubroot
of Crucifers (Plasmodiophora brassicae).
G. Powdery mildews: It is caused by a powdery growth on the
surface of leaves, buds, young shoots, fruits, and flowers.Powdery
mildew is caused by many fungal species like genera Erysiphe,
Microsphaera, Phyllactinia, Podosphaera, Sphaerotheca, and Uncinula.
H. Leaf Curls: Curling,t hickening, distortion by distortion and
coloration of leaves occur by pathogen attack.Almond Leaf Curl,
Peach Leaf Curl, maple leaf curl caused by Taphrina deformans.
I. Leaf Spots: Leaf spots are very common in both biotic and abiotic
plant disorders. Fungal leaf spots often take the form of localized
lesions consisting of necrotic and collapsed tissue. Leaf spots can
vary in size and are generally round and concentric, but can be ovoid
or elongated on both leaves and stems of the host. Many different
crops are affected by species of the fungal genera Alternaria,
Cercospora, Colletotrichum, and Myrothecium.
J. Rusts: Rusts are plant diseases caused by pathogenic fungi of
the order Pucciniales (previously known as Uredinales). These are
obligate plant pathogens that only infect living plants. Each spore
can typically infect only one kind of plant which is very host specific.
Hemileia vastatrix (Coffee rust) and Puccinia graminis (Stem rust of
wheat).

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K. Canker: A localized necrotic lesion on woody tissue, often
sunken is the characteristic feature.Apple European Canker (Nectria
galligena), upper Twig Canker (phomopsis sp.)
L. Anthracnose: An ulcer like lesion is seen that can be necrotic
and sunken. These lesions can be found on the fruit, flowers and
stems of the host.Apple Anthracnose of stems and or leaves
(Cryptosporiopsis sp. Formally Pezicula sp.) Dogwood Anthracnose
(Discula distructiva) .
M. Rots: Soft and dry root rots: Rot is disintegration of fleshy leaves,
roots, tubers and fruit.This disease is common in fruit crops such
as apricots, cherries, plums and peaches. Red rot disease of
sugarcane caused by Colletotrichum falcatum.

Famines due to fungal pathogens: The two important famines of


the world namely Great Irish Famine (1845–49) that occurred in
Ireland due to Phytophthora infestans causing late blight potato crop.
Bengal famine due to Helminthosporium oryzae causing rice blast.

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3.4.2 Blast of Rice
Classification
Class : Deuteromycotina
Order : Moniliales
Family : Moniliaceae

Rice blast is a serious fungal disease of rice (Oryza sativa L.) that is
threatening global food security. It has been extensively studied
owing to the importance of rice production, consumption, vast
distribution and destructiveness across the world. Taxonomically
it is described as Magnaporthe oryzae based on recent phylogenetic,
molecular and morphological data, isolates of the fungus from rice
and closely related isolates from other grasses like Eragrostris
curvula, Eleusine coracana, Lolium perenne, and Setaria spp. The
isolates from Digitaria sanguinalis (crab grass) are distinct and should
be described as Magnaporthe grisea.

Know more: Magnaporthe oryzae causing Rice blast was previously


described by the name Pyricularia oryzae (= P. grisea)

SYMPTOMS :Rice blast can cause total crop failure if it infects


aboveground tissues of rice plants at any growth stage. The fungus
may attack all of the ground portions of rice plant at various stages
of growth: the leaf, collar, node, internode, base or neck, and various
parts of panicle, and often on the leaf sheath.The neck blast infects
the panicle causing seed failure to fill, or causing the whole panicle
to fall over as it rots.The pathogen produces lesions on leaves (leaf
blast), leaf collars (collar blast), culms, culm nodes, panicle neck nodes
(neck rot), and panicles (panicle blast). They vary in color and shape
depending on varietal resistance, environmental conditions, and age.
The lesions frequently remain short in size (1 to 2 mm) and grey to
deep brown color on resistant cultivars. Symptoms of collar node

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infection consist of general area necrosis when the two tissues come
together. Collar infections may kill the whole leaf and can spread
to and around the sheath a few millimeters later, when the plant is
damaged in the collar region.The lesions are often greyish brown
discoloration of panicle branches which may break at the lesions
over time.Neck blast infection results in development of triangular
purplish lesions accompanied by expanding on either side of the
neck.The panicles become white when young necks are infected and
later infection caused incomplete grain filling and poor grain quality
(Fig 3.4.2).

Fig. 3.4.2. Severe leaf blast symptoms

Biology of rice blast pathogen : The fungus that causes rice blast
is called Magnaporthe oryzae (formerly Magnaporthe grisea). It
produces sexual spores (ascospores) in structures called asci, and is
classified in the newly erected family Magnaporthaceae in
Ascomycetae. These asci found within specialized structures called
perithecia. The mycelium of M. oryzae is septate and the nuclei
within the mycelium and spores of this fungus are haploid.
Sexual Reproduction : The sexual, or teleomorphic stage of the
rice blast pathogen has not been found in the field but can be
produced in the laboratory if isolates of opposite mating type found

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paired. Hyaline, fusiform or spindle-shaped with tapering ends
ascospores with three septa are formed. Unitunicate asci are present.
This fungus is considered to be heterothallic with a bipolar mating
system. It is mating controlled by two different alleles at a single
locus with additional genes controlling the sexual cycle.
Asexual Reproduction: The asexual stage of Magnaporthe oryzae
is described by the name Pyricularia oryzae and it is the most
common spore form of the fungus. These spores, called conidia,
are produced abundantly on lesions and in culture on specialized
stalks, called conidiophores. The conidia are usually three-celled,
and produced on the apex of a conidiophore. Sporulating colonies
on agar plates may take on a fleecy greyish appearance.
Disease cycle of rice blast: Rice blast fungus can infect all above
portions of rice plants including leaves, stems, neck, nodes, and
panicles at all stages of growth and development due to its polycyclic
nature. When airborne conidia land on rice plants, these adhere to
the surface through the sticky mucilage produced during hydration
from an apex compartment of conidium tip. Conidia germinate
when enough humidity is present on the host plant surfaces. Germ
tube emerges from conidium’s tapering end and grows over the plant
surface. The germ tube is swollen and enlarged and to form an
appressorium after sometime. Fungus requires hard surface to form
appressoria structure. Appressoria contains chitin and melanin like
molecules in host cell wall and the presence of glycerol enhances
turgor pressure to allow penetration. The appressoria passes via
stomata into the rice plant. The blast fungal hyphae expand into
the tissues of plants and ultimately develops lesions. Plant tissue
invasion and colonization is intended by the fungal hyphae which
invades the plasma membrane as well as epidermal cells. Specialized
feeding structures or feeding hyphae are formed during early tissue
invasion to help colonize the tissues and obtain nutrients from living

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plant tissues. The hyphae move into various plant cells through
plasmodesmata. The blast lesions are evident within 3 to 4 days of
infection. The blast fungus sporulates quickly under high humidity
and releases conidia in plenty. The conidia are usually transmitted
by wind or rain splash to neighboring rice plants, starting another
disease cycle.
Favorable conditions for the pathogen Low night temperature
(22 to 28°C), High relative humidity (> 95%),New
deposit,Extended leaf wetness period (>10 hrs),Cloudy and drizzling
weather,Soil fertility (High N),Degree of host susceptibility and
Straw of the previously infected crop heaped nearby.
MANAGEMENT:
Preventive measures: Use of resistant varieties like Gauthami,
IR-36, IR-64, Parijatha, Rasi,Sasyashree,Salivahana, Simhapuri,
Srinivas, Tikkana. Burn previously blast affected straw and
stubbles.Use of disease-free seeds.Use of balanced rates of nutrition
based on soil test- split applications of nitrogen (Excessive N use
can promote excessive luxuriant crop growth which increases the
relative humidity and leaf wetness of the crop canopy. This can lead
to increased infection).
Cultural practices: Avoid stagnating of water in the field.Use
IR-64/MTU-1005, Jaya and Jyothi varieties.Do not use high dose
of nitrogenous fertilizers.
Biological control: Seed treatment with Pseudomonas
fluorescence 10g / 1 of water for 30 min.Seedling root dip treatment
with Pseudomonas fluorescence (4g / 1 of water) for 20 min.Foliar
spray of Pseudomonas fluorescence (4g / 1 of water) at 20-25 days
after transplanting
Chemical control: Seed dress with either tricyclazole 75 WP or
carbendazim 50 WP @ 2g/kg of seed.Spray 1g of carbendazim (780g/
acre) or 1 ml edifenphos (780ml/acre) or 0.6g of tricyclazole (120g/

196 APSCHE : SEMESTER-I BOTANY TEXT BOOK


acre) in 1 liter of water, depending on severity repeat spray at 10-
12 days of first spray. Seed treatment with Pseudomonas
fluorescence

Fig 3.4.1 Disease Cycle of Blast of Rice

Fact to Know: The asexual stage of Magnaporthe oryzae is


described by the name Pyricularia oryzae and it is the most common
spore form of the fungus.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 197


Lichens- structure and reproduction; ecological
Chapter -3.5
and economic importance

Learning Outcomes : At the end of the chapter you should be


able to:
· Discuss the general characteristics of lichens.
. Explain the ecological and economic value of lichens.

Lichens are the most extraordinary group of plants found


in the plant kingdom. They are unique in being formed by the
intimate association of two very different plants, one of which is
filamentous fungus and the other an alga. They are associated in
such a way that they appear to be a single plant. The fungal
component always envelops the algal component of the association
and the plant resulted from the combined growth of the two is more
or less constant in form and internal structure.

FACTS: In most of the lichens the fungus is called ascolichen,


a member of Ascomycetes

A lichen is a plant consisting of a fungus and an alga. Thus, the algal


cells of the association enveloped by the fungus, two separate plants
form with each other to appear a single plant. The combined growth
of both partners results in a constant and internal structure of the
lichen. The term lichen’ was first used by Theophrastus to denote a
superficial growth on bark of trees. Lichens were first discovered
by Tulsane (1852). The science of studying lichens is lichenology
and one who studies this science is known as lichenologist.

External Forms: The fungal component in four genera of lichens


is a member of Basidiomycetes and in rest it is an Ascomycetous
member and as such they are called Basidiolichens and Ascolichens.

198 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Basidiolichens have only four genera, all are restricted to tropical
regions, while ascolichens are restricted to temperate regions. The
algal components may belong to Myxophyceae or Chlorophyceae
algae. They may be chief filamentous Trentepohlia or non-
filamentous. Some epiphyllous lichens common in tropics have been
observed with a curious parasitic filamentous alga, Cephaleuros.
Strigula, very common in tropics. In many, however, it is with blue-
green alga. eg.. Nostoc, Gloeocapsu and Rivularia.One lichen has been
reported in which a filamentous fungus is associated with an
autrotrophic purple bacterium rather than with an alga Upholf
stated that the bacterium is of a species that produces a pigment,
which gives the lichen a reddish appearance. Sessenguth (1926),
opined that these bacteria are not an essential component of the
lichens.

The accountancy for the consistency in the different kinds


of lichens is always due to the formation of the same fungus and
the same alga. It has always same habit and internal structure. This
association is mainly physiological. The “dual hypothesis” of the
lichen was proposed by Schwendner in 1867. This states that the
various blue green and green algal components or gonidia are really
algae and that the complete lichen in all cases represented a fungus
living parasitically on an alga. He further described that, though
eventually the alga is destroyed, it is at first excited to more vigorous
growth by contact with fungus and in the course of a generation it
may become changed beyond recognition both in size and form. In
support of this theory of the composite constitution of the thallus
Schwendener pointed out the wide distribution and frequent
occurrence in nature of the algae that become transformed to lichen
gonidia.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 199


 

      


  
   
   
    
  

       

 
     

   


 
    
          

    
        

    
  


200 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Rees (1871), was the first of those who attempted to make
and proved synthetic culture. For this purpose, spores from the
apothecium of Collema taken and sowed them on pure culture of
Nostoc. As a result, a lichen thallus has formed but did not succeed
in producing any fructification. It has further observed that the
hyphal filament from the germinating spore died off, when no
Nostoc was present. Next synthetic cultures were made by Barnet
(1873), Treub (1873), Borzi (1875) and Stahl (1877), with a series
of lichen spores. Bonnier (1886-1889), made interesting series of
synthetic cultures between the spores of lichens germinated
carefully in sterilized conditions and algae taken from the open. He
observed that in a few years not only the lichen thalli were produced
but, in many cases, apothecia were also developed. Even till the
present time it has not been possible to isolate the fungus and grow
it to maturity in pure culture except in a single instance in the
basidiolichen Cora where the fungus, Thelephora, is known to grow
in nature apart from the lichen. Alga, on the other hand, can grow
independently when isolated.

Habit and Habitat: Lichens have been described as “perennial


aerial plants of somewhat low organization.” In the form of
spreading encrustations, horizontal leafy expansions, or upright
strap-shaped fronds or pendulous filaments, they take possession,
of the tree trunks, palings, walls, rocks or even soils that afford
them a suitable and stable foothold (A.L. Smith, 1920). A few of
the largest genera are: Lecidia, Buellia, Lecanora, Parmelia, Physcia,
Collema, Sticta, Cladonia, Ramalina and Usnea. Direct light, moderate
or cold temperature, constant moisture and pure atmosphere favour
their growth. Polluted, smoky atmosphere as found in industrial
area is not favourable for growth.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 201



  
       

 
   
 

 

                          
 

  

       

* * * * * * *
This group has about 400 genera and 15,000 species and is
treated separately instead of its connection with fungi or algae.
Lichens are worldwide in distribution and are found in a great variety
of habitats like the surfaces of bare rocks, barks of trees and on soil.
Many lichens thrive and multiply in habitats where other vegetation
is practically nonexistent. In many habitats these are exposed to
the greatest extremes of temperature and humidity and can undergo
extreme desiccation for long periods and revive when conditions
become favourable. Lichens form the pioneers of vegetation and
are of geological importance in colonisation of rocky habitats.

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Lichens are the first plants to appear on the boulders of the rocky
cliffs. Growth of a lichen is accompanied by a disintegration of the
part of the rock beneath it. When a lichen dies its decaying remains
together with the rock particles from a soil in which other plants
may grow. The first successors are generally mosses but sooner or
later colonisation by flowering plants occurs.
In India, they are much more common in the Eastern
Himalayas (with an annual rainfall of 133 inches or more) and higher
hills of Peninsula than in the Western Himalayas with much less
rainfall. They are found there is abundance on tree trunks, especially
on the shaded sides; they occur on exposed rocks, where no other
vegetation could grow, they occur on old logs and on the ground in
woods. In fact, in any position capable of sustaining plant life. they
are likely to develop. In damp forests of the tropical and subtropical
regions, they often hang in long festoons from trees. In the arctic
zones they develop as cushion shaped masses on the grounds, where
they often form an important food of the reindeer. G.L. Chopra has
published a monograph on the lichen flora of Darjeeling the Sikkim
Morphological structure: Based on the morphological structure,
the lichens are of three types i.e., Crustose, Foliose and Fruticose.
(1) Crustose: These lichens have flattened thalli and closely
adherent to the substratum. Thalli of majority of such lichens has
more or less leathery texture and is internally differentiated with
the algal component always restricted to a definite portion of the
thallus. Some lichens on the centre of the thallus or a lichen may be
attached by several rhizines.
e.g., Haematomma, Lecidia, Lecanora
(2) Foliose (Foliaceous) : The lichens, which have flat and leaf
like thalli, with lobed or irregular margins. Some part of their thallus
usually adheres more or less firmly to the substratum by means of

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Fig:3.5.1 Habit of lichens A) Crustose : Caloplaca
B) Foliose: Xanthoparmelia substrigosa C) Fruticose: Usnea
Source: http://go.microsoft.com/fwlink/p/?LinkId=255141

strands of hyphae called the rhizoids, e.g., Peltigera, Physcia, Parmelia,


etc. Some foliose lichens like Gyrophora are attached by smell central
holdfasts to the ventral surface. In some large foliose differentiated
into layers of tissues and known as homomerous. Thalli of most
foliose and fruticose lichens are differentiated into several layers of
tissues, and therefore known a heteromerous.
3. Fruticose: These lichens have much branched, cylindrical,
ribbon-like, flattened or netimes filamentous thalli. They are much
branched, and appear shrubby, and so the name fruticose is given
to them. They may be erect in Cladonia or pendant in Usnea. Air
thalli are attached to the substratum by the basal portion only which
is composed of strands of densely packed hyphae.

Fig: 3.5.2 V.S. of Lichen thallus

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Internal structure: The typical heteromerous thallus shows, in
a transverse section the following strata or regions, an upper
cortex of densely packed hyphae, a gonidial layer with algal cells,
medulla with loose fungal hyphae and a lower cortex with densely
compacted hyphae.
1. Upper Cortex: It forms the upper surface which is generally
thick and protective. The fungal hyphae in this region grow more
or less vertically and are compactly interwoven to produce
pseudoparenchymatous layer. The fungal hyphae are compactly
arranged so that there is no intercellular space between the hyphae.
It is the blue green or green zone which lies
immediately beneath the upper cortex. It consists of a network of
loosely interwoven fungal hyphae with the algal cell of green alga,
intermixed with fungal hyphae. The Algal region is the

It contain the central core of the thallus. In certain


regions there is less compact and consists of loosely interwoven
hyphae with large space between them. The fungal hyphae in this

directions. The central hyphae of the medullary region usually run


longitudinally.
It forms the lower surface of the thallus and is
composed of densely compacted hyphae. They may run
perpendicular to the surface of the thallus or parallel to it. Bundle
of hyphae often arise from the surface of lower cortex and penetrate
the substratum to function as anchoring organs. In some lichen
species the lower cortex is absent. In the simplest of lichens, the
algal cells or gonidia, are scattered uniformly among the fungal
elements and in these cases the thallus is described as
homoiomerous. The majority of lichens, however possess a stratified

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thallus, in which the gonidia are found as a definite layer or layers
embedded in pseudoparenchy matous fungal hyphae. Such thallus
is described as heteromerous as found in Parmelia and Physcia.
The term hypothallus is sometimes applied to the lower
layer, when specially modified, in immediate contact with the
substratum. One or more of these layers may be missing in many of
the species. For instance, in the crustose species, the lower cortex
may be absent and the medullary region lies in intimate contact
with the substratum. The bodies of some crustose lichens are partly
or wholly embedded in the bark, disintegrated rock, or soil upon
which they grow. In the latter case, all that appears above the surface
of the substrate may be the fruiting bodies of the lichens. In the
foliose lichens, all the four regions are found and the rhizines grow
out from the lower cortex and fix the plant body to the substrate.
In the cylindrical species of the fruticose lichens, the thallus is
radially symmetrical and there is a central region of hyphae,
surrounded by a zone containing algal cells and, this in turn, by an
outer zone of compact hyphae. The lichen is attached to the
substrate by a definite basal portion composed of strands of densely
packed hyphae.
Outgrowths of the thalluThe surface of the thallus of a lichen
often exhibits outgrowth In the form of hairs, warts etc. In addition,
there are certain structures peculiar to a lichen thallus, which may
be:
1. Isidia: Many lichens have corolloid outgrowths from the f surface
of the thallus. Isidia consist of an external cortical layer and internal
algal layer. The alga is the same as that in the thallus. T primary
function of an Isidium seems to be that of increasing the
photosynthetic surface of the thallus. Sometimes, they become
detached from the thallus and serve as vegetative reproductive
bodies.

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2. Cephalodia: In certain species of lichens, peculiar gall like
growths has developed in the interior of the thallus which cause
slight projection on the upper or lower surface of thallus. These
structures are known as the cephalodia. They are distinguished from
Isidia by possessing as gonidia, algal cells foreign to the ordinary
part or the thallus. The function of these peculiar structures is
unknown. Zukal has suggested that they may play the part of water
absorbing organs.
3. Cyphellae: They are small, circular depressions, or roundish
cavities, scattered over the undersurface of certain lichens. Each
has a definite rim and open into the medulla of the lichen. Probably,
their function is to aid aeration.
Reproduction: There are three methods of reproduction in lichens:
1. fragmentation 2. soredia 3. formation of fungal spores.
Fragmentation: under suitable conditions any separated portion
of the body of a lichen may develop independently. Progressive
growth and death of older portions result in the production of new
independent plants. Several portions of a thallus may develop into
a new plant provided they contain both symbionts. This is of
frequent occurrence in pendent forms like “Californian Spanish
Moss”, Ramalina reticulata where detached portions of the pendent
thallus are carried to other tress by wind and these develop into
new plants.
Soredia: The commonest method of propagation is by soredia. The
soredia are formed in a large number of lichens. A soredium consists
of one or more algal cells enclosed by a few fungal hyphae. They
may develop over the entire surface of a thallus or in localised
pustule like areas, the soralia. They arise usually in the gonidial layer
of the thallus by division of the gonidia and development around
the hyphal region. Their increase in number leads to the rupture of
the enclosing cortical layer and the soredia escape from the thallus
as a powdery mass. They are provided with both fungal and algal

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elements and they are able to develop directly, under suitable
conditions, into a new thallus. In certain species of lichens, this is
the only known method of reproduction, for not only some forms
are nearly always without spore formation and in the others, the
spore represents only the fungal component of the thallus, and Its
success in the development of a new lichen thallus depends on the
chance meeting, at the time of germination, with the appropriate
algal component.
Spore formation: In lichens, of the two components, it is the
fungus alone that produces the fruit bodies and the spores and the
alga takes no part in the process. With the exception of a few tropical
Basidiolichens, all known lichens belong to the division Ascolichens,
in which the fungal element belongs to Ascomycete. So,
consequently the fruit bodies or fructifications produced in
Ascolichens are either apothecia or perithecia and the spores
produced are ascospores. In basidiolichens, the fungus belongs to
the class Basidiomycetes, and the fruit body produced is
characteristic of that class and the spores produced are
basidiospores.
In Ascolichens, two chief types of fruit bodies are found,
the apothecium and the perithecium, the first when the fungus
belongs to Discomycetes and the second when it belongs to
Pyrenomycetes. In the simplest or Lecideine type, the apothecium
solely consists of fungal tissue and differs in no striking way from
that found in the Ascomycetes, but in the Lecanorine type, the rim
of the apothecium has the same structure as the rest of the thallus
including algal cells. The asci are separated by paraphyses and the
whole hymenial surface resembles that found in Pezizales. Each
ascus is of the typical Ascomycete form, and each usually has eight
spores. Lichen apothecia vary greatly in size and not infrequently.
The hymenlum is of different colour from the thallus and is
sometimes quite brilliantly coloured. The perithecia where they

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occur also resemble those of the Ascomycetes. The hymenium may
line the entire inner surface or be restricted around the upper part
only.
An ascospore may develop into hypha which branches and
elongates until its food supply is exhausted. It will die, if the hypha
does not come into contact with the algae of the species with which
it is ordinarily associated.
Sexual Reproduction : Sexual reproduction takes place by the
production of specialized male and female reproductive Structures.
Male reproductive structure is called spermogonium and the female
reproductive Structures is called carpogonium.
(a) Spermogonia: In certain species of lichen Spermogonia are
reported as pycnidia like structure. It is a flask-shaped receptacle
immersed in a small elevation on the upper surface of the thallus.
It opens by a small pore called an ostiole at the surface. The cavity
of the spermogonium is filled with sterile and fertile hyphae. The
fertile hyphae produce minute rounded cells at their tips which are
the male cells and are called spermatia. They are non motile and are
produced in large number. The spermatia are set free in a slimy
mass which oozes out through ostiole.
(b) Carpogonia: The carpogonium is a cellular filament consisting
of two portions namely the lower coiled portion and upper straight
portion. The coiled portion constitute the ascogonium which is
Multicellular and has uninucleate cells. In certain species they are
multinucleate. The ascogonium lies deep in the medullary region
of the thallus. The carpogonia either develop from the hyphae or
from the medullary region or from the hyphae deep in the algal
layer of the lichen thallus. The straight, elongated and separate
upper portion of the carpogonium is called the trichogyne. The
terminal portion of the trichogyne ends in a long cell which project
beyond the surface of the thallus.

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At the time of fertilization, several spermatia are lodge to
sticky tip of the trichogyne. A few empty spermatia are found at
the tip at the trichogyne. This is due to migration of protoplast
into the trichogyne. But it has not seen the actual migration of the
male nuclei down the trichogyne.
(c) Formation of ascocarp and ascospores- After fertilization
several ascogenous hypha develop from the basal portion of the
ascogonium. Asci are produced at the ends of these freely branched
ascogenous hyohae . Two nuclei in a young ascus fuse to farm a
diploid nucleus. This diploid nucleus divides and redivides to farm
8 haploid nuclei. The asci are uninucleate or binucleate. These asci
bearing Structure are ascocarps.
Normally eight ascopores are formed in each ascus. They
vary in colorur , shape, size and Structure. Ascopores are unicellular
, transversely separate or both transversely and longitudinally
separate. On liberation each ascospore germinates by producing
hyphal branch. This hyphal branch comes in contact suitable alga,
ultimately lichen thallus is formed after the combined growth of
the fungus and the alga.
Development of ascocarp and Sexuality in lichens : The
question of sexuality of lichens has been hotly disputed in common
with that of the rest of the Ascomycetes. In some cases, the ascocarps
develop as a result of a previous fertilization by spermatia. In other
cases, the ascocarps develop without such a union, while in still
other cases, the reduction goes still further and the ascogenous
hyphae. Instead of developing from ascogonia, they are derived
directly from the vegetative hyphae.
Economic Importance of Lichens : Many lichens have been highly
used in the arts, food and medicine.
Medicine: During the Middle Ages and, even in some quarters, to
a much later period, lichens were extensively used in medicine in
various European countries. Peltigera canina was long regarded as a

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sovereign cure for hydrophobia; Platysma juniperinum was lauded
as a specific for jaundice. Above all, even today, in some parts of the
world, lichens are believed to be effective in alchemy.
Species of Cladonia and Cetraria are used in intermittent fever while
Evernia furfuracia is used in cough. Acaraspora smaragdula secretes a
compound possessing antibiotic properties.
Usnicacid obtained from lichens has antibacterial activity
and in combination with streptomycin has proved very effective
against tuberculosis. Some lichens also act as allergens and cause
allergy.
Food: Many lichens for supplying food for man and beast. This is
due to their containing starchy substances and in some cases a small
quantity of saccharine matter of the nature of marmite. One of the
most nutritive species is Cetraria islandica “Iceland moss”, which
after being boiled is reduced to a powder and made into cakes or
boiled and eaten with milk, by the poor Icelander, whose sole food
in often constitutes. Similarly, Cladonia rangiferina and C. sylvatica
the familiar “reindeer moss” are of considerable value as food for
man and reindeer and cattle. It forms dense tufts, sometimes 12"
in height, and is abundant in extremely cold regions, where other
vegetation is practically not existingt and where it may be buried in
snow for long periods without injury.
Two species constitute an important part of the vegetable diet. A
leathery Ubicara known as “rock tripe” has often been eaten by
travellers when they face starvation in arctic regions. Lecanora have
been used as food in the barren plains and mountains of Western
Asia and Northern Africa. They are considered to have constituted
a part of the material called “manna” by the Israel people. They are
still known as the ‘bread of heaven’, apparently because they are
scattered by wind and deposited at considerable distances from their
source. In India, a species of Parmelia known in Telugu as rathapu”
or “rock_flower has been used as curry by natives in the Karnataka

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and Tamil Nadu and is cherished as a delicacy. It is collected during
April-May and forms a profitable business.
Another nutritious lichen is the “Tripe de Roch” of the Arctic
regions, consisting of the several species of the Gyrophorel when
boiled is often eaten by Canadian hunters and Red Indians. But the
most familiar esculent lichen of all is the manna lichen two species
of Lecanora, which in times of drought and famine has served as
food for a large number of men and cattle in the barren plains and
mountains of Western Asia and Northern Africa. This is the material
referred to as Manna” by Israelites and still known as the “bread
from heaven” apparently because they are sometimes picked up by
winds and deposited at considerable distance from their source. In
Scandinavia and Russia, an alcoholic spirit has been distilled from
Cladonia rangifetina, and extensively consumed especially in seasons
when potatoes were scarce and dear. Formerly also Sticta pulmonaria
was much employed in brewing instead of hops.

Find: Enumerate different types of lichens used as food

Art: Poisons, drugs, perfumery and alcohol have been prepared from
certain lichens. Dhup and Havan Samagri are prepared from certain
species of lichens, e.g., Ramalina, Evernia, etc. having sweet scented
thalli. The most important species of the lichens which yield dyes
is species of Roccella, Lecanora, and Parmelia, etc. Roccella tinctoria,
and R. Montaguei yield a rich purple dye known as “archil “. “perelle”
is prepared from Lecanora parella and used in the preparation of a
red or crimson dye. Inferior to this is ‘’cudbear” derived from
Lecanora tartarea. Yellow dyes are again derived from Parmelia
caperata. At one time, some species of lichens were also used for
supplying a gum as a substitute for gum aribic. In 17th Century,
species of Usnea, Ramalina and Cladonia rangiferina etc., were used
in the art of perfumery. “Orcein”, a purified extract or archil serves

212 APSCHE : SEMESTER-I BOTANY TEXT BOOK


as a stain for microscopic preparations. The most note-worthy of
colour substances is orchil or cudbear obtained from species of
Roccella and Lecanora. Orchil was formerly used in the textile
industry for dyeing wool and fabrics Orecin, a purified extract of
orchil is used as a stain in the laboratories for microscopic
preparations. Litmus solution is made by grinding up the plant
bodies of species of Roccella and Lecanora and extracting the
colouring matter. Litmus, used as an indicator for acidity, is also
derived from Roccella. Many lichens yield colouring substances
when soaked in alkaline solution like caustic potash.
Lobaria pulmonaria, the lungwort, besides being used for the
treatment of lung diseases, has also been employed in tanning, as a
substitute for hops in brewing and in perfume making.
Usnea being inflammable, easily catches fire in nature
causing forest fires. Some are reported to be radioactive.
Lichens also harm the host tree on which they grow. They
also reduce the market value of glasses and marble stones due to
etching of the surface by the action of their acidic secretions.

Activity: Take the different types of spores of fungi and draw a


table showing different classes of fungi which u have studied
along with examples

Self- Analysis

 Draw the forked diagram of Ainsworth classification briefly


explaining characters of Classes of Fungi
 Study about the diseases caused by fungi in plants, animals
and human beings
 Study about the preparation of various biological agents of
fungi for the disease control of plants
 Study about the diseases caused by fungi in plants, animals
and human beings

APSCHE : SEMESTER-I BOTANY TEXT BOOK 213


 Study about the preparation of various Biological agents of
fungi for the disease control of plants
Collect and use your knowledge to answer about a symptom
of a fungal infection that causes plant tissue
 Study the different types of lichens available in Andhra
Pradesh and their economic importance
Collect the lichen samples and photographs from your locality
and observe under microscope

Self-Assessment

1. Which polysaccharide is 5. True mycelium lacking in


usually found in the cell the following class of
wall of fungi? Deuteromycotina
a. starch a. Myxomycetes
b. glycogen b. Zygomycetes
c. chitin c. Phycomycetes
d. cellulose d. Blastomycetes
2. Which of these organelles is 6. Puffballs, mushrooms and
not found in a fungal cell? bracket fungi belong to
a. chloroplast this class
b. nucleus a. Myxomycetes
c. mitochondrion b. Basidiomycetes
d. Golgi apparatus c. Phycomycetes
3. Perfect state absent in the d. Blastomycetes
following Division 7. The internodal aerial and
a. Deuteromycotina arching hyphae in
b. Zygomycotina Rhizopusare
c. Phycomycetes a. Stolons
d. Myxomycetes b. Rhizoids
4. Spores are found in c. Sporangiophores
pycnidia or acervuli in this d. Columella
class 8. Hyphae that form
a. Coelomycetes progametangium are
b. Chytridiomycetes a. Sporangiophore
c. Myxomycetes b. Columelloplasm
d. Zygomycetes c. Zygophores
d. Sporoplasm

214 APSCHE : SEMESTER-I BOTANY TEXT BOOK


9. The spore neither infect b. Podaxon podaxis
primary host nor the (Khumb)
alternate host. c. Lycoperdon and
a. Uredospore Clavatia (puff balls)
b. Pycniospores d. All of the above
c. Basidiospore 15. Ergosterol is a product of
d. Aecium many molds and yeasts
10. The spores are incapable of and is the source of
infecting Barberry plants a. Vitamin D
but they can infect wheat b. Vitamin C
plant c. Vitamin A
a. Uredospore d. Vitamin B
b. pycniospores 16. Powdery mildew is caused
by many fungal species like
c. Basidiospore
genera
d. Aeciospores
a. Puccinia
11. The spores unable to infect
b. Aspergillus
the wheat plant but infect
c. Erysiphe
the alternative host
d. Plasmophora
Berberis vulgaris
17. Red rot disease of
a. Basidiospores sugarcane is caused by
b. Aeciospores a. C o l l e t o t r i c h u m
c. Chlamydospores falcatum
d. Teliospores b.
12. The mycelium produced by c. Aspergillus niger
germination of d. Pencillium notatum
basidiospores is
18. Coffee rust is caused by
a. Dikaryotic
a. Fusarium oxaliformis
b. Coenocytic
b. Aspergillus niger
c. Monokaryotic
c. H e m i l e i a
d. Tubular vastatrix
13. Ergot toxin is producedbythe d. Taphrinadeformans
following fungus
19. Californian Spanish
a. moss is
b. Aspergillus niger a. Parmelia
c. Pencillium notatum b. Cladonia
d. Fusarium oxaliformis c. Ramalina
14. Edible Mushrooms are reticulata
a. Agaricus campestris d. Humaria
(dhingri) granulata

APSCHE : SEMESTER-I BOTANY TEXT BOOK 215


20. Hematomma, Lecidia are https://biologyreader.com/
a. Foliose lichens Rhizopus.html
b. Fruticose lichens http://go.microsoft.com/fwlink/p/
c. Filamentous ?LinkId=255141
lichens .https://biologyreader.com/
d. Crustose lichens Rhizopus.html
http://go.microsoft.com/fwlink/p/
Interactive links ?LinkId=255141
http://go.microsoft.com/fwlink/p/ Video source:
?LinkId=255141 https://www.youtube.com/
https://www.youtube.com/ watch?v=2NvrL_zekqw
watch?v=uAJtYSIX6MU https://www.youtube.com/
Online resources watch?v=x0j4EMBx_1s
http://go.microsoft.com/fwlink/p/ https://www.youtube.com/
?LinkId=255141 watch?v=6vIF3zazwAw
http://go.microsoft.com/fwlink/p/ https://www.youtube.com/
?LinkId=255141 watch?v=Bp9rW5-ZXu8
http://bioweb.uwlax.edu/bio203/ https://www.youtube.com/
2011/olbrantz_chri/ watch?v=AeuP5IYP5HA
reproduction.htm https://www.youtube.com/
https://www.vedantu.com/biology/ watch?v=m0UMYrJPQac
Rhizopus https://www.youtube.com/
http://www.vpscience.org/ watch?v=qOjkmjJSI0s
materials/Fungi-Rhizopus.pdf https://www.youtube.com/
https://ohioline.osu.edu/factsheet/ watch?v=IxOfL1Jxa28
plpath-gen-7 https://www.youtube.com/
http://go.microsoft.com/fwlink/p/ watch?v=HD1V0r28l9U
?LinkId=255141 https://www.youtube.com/
https://herbarium.usu.edu/fun- watch?v=mFxh6IQqrcwhttps://
with-fungi/lichens www.youtube.com/
http://www.lichens.lastdragon.org/ watch?v=x0j4EMBx_1s
faq/lichenthallustypes.html https://www.youtube.com/
http://go.microsoft.com/fwlink/p/ watch?v=wFcgrY9oizc
?LinkId=255141 https://www.youtube.com/
https://classofbiology.com/ watch?v=j7U8CZlYqTI
reproduction-in-fungi/ http://go.microsoft.com/fwlink/p/
http://go.microsoft.com/fwlink/p/ ?LinkId=255141
?LinkId=255141

216 APSCHE : SEMESTER-I BOTANY TEXT BOOK


References : Basidiospore: The haploid spore
B.R. Vashishta, A.K .Sinha, 2007. which is products of meiosis, thin-
Botany for Degree Students, S. Chand walled and colourless, cannot infect
& Company, RamNagar, New Delhi the cereal host, but can infect the
O.P.Sharma,1989, TATA Mc Graw-Hill alternative host i.e., berberis plant.
Publishing Company Limited. Brewer’s yeast: It is
Mehrotra, R.S. & K. R. Aneja (1990) Saccharomyces cerevisiae used in
An Introduction to Mycology. New fermentation to prepare common
Age International Publishers, New alcoholic beverages such as wine
Delhi and beer
Cephalodia: Peculiar gall like
Kevin Kavanagh (2005) Fungi ; Biology
outgrowths developed in the
and Applications John Wiley & Sons,
interior of the thalluscausing slight
Ltd.,West Sussex, England
projection on the upper and lower
John Webster & R. W. S. Weber (2007) surfaces of the thallus
Introduction to Fungi,Cambridge Cephalodia: Peculiar gall like
University Press, New York outgrowths developed in the
interior of the thalluscausing slight
Smith, G.M (1955): Cr yptogamic projection on the upper and lower
Botany (Vol.I Algae, Fungi & Lichens) Mc surfaces of the thallus
Graw-Hill Book Co., New York.
cheese: It is Penicillium roqueforti,
Glossary : pr incipal mold. Which is
responsible for the blue veins and
Aecidiospore: Dikar yotic, Round,
pungent taste to the cheese.
yellowish red, unicellular, binucleate,
thick walled with six germ pores, Chlamydospores: A thick-walled,
produced in chains able to germinate intercalary or terminal, asexual
on the cereal host but not on the spore formed by the rounding of a
alternative host Biological Control: A cell or cells.
method of controlling pests such as Clamp connection: Clamp
insects, mites, weeds and plant connection is a bridge-like hyphal
diseases using other organisms connection characteristic of the
Anthracnose: Ulcer like lesions that secondar y mycelium in many
can be necrotic and sunken Basidiomycetes. They are found
during nuclear division and help in
Anthracnose: Ulcer like lesions that
dikaryotization of adjacent cells.
can be necrotic and sunken

APSCHE : SEMESTER-I BOTANY TEXT BOOK 217


Coenocytic hypha: single hypha that in oxygen-r ich and oxygen-poor
lacks septa and contains many nuclei environment
Columella: The crescent structure Foliose Lichen: The body is flat,
present at the tip of long erect stalk lobed or deeply incised thallus
sporangiophore of Rhizopusis is attached to the substratum by
known as columella. It gives support rhizenes
to the Sporangium which has asexual Foliose Lichen: The body is flat,
sporangiospores. lobed or deeply incised thallus
Crustose lichen: The body is in the attached to the substratum by
form of incrustations on the surface of rhizenes
the rocks, barks of trees Fruticose Lichen: The body is much
Crustose lichen: The body is in the branched, cylindrical to ribbon like
form of incrustations on the surface of thallus that may be erect or pendant
the rocks, barks of trees Fruticose Lichen: The body is much
Cyclosporine: An immuno- branched, cylindrical to ribbon like
suppressant drug which reduces the thallus that may be erect or pendant
risk of rejection after organ Haustoria: Modified hyphae on many
transplant and precursors of steroid parasitic fungi that penetrate the
hormones. tissues of their hosts, release digestive
Cyphellae: Small, circular depressions enzymes, and/or absorb nutrients
or roundish cavities of the lichen from the host
thallus Hemileia vastatrix: Pathogen of
Cyphellae: Small, circular Coffee Rust
depressions or roundish cavities of Hemileia vastatrix: Pathogen of
the lichen thallus. Coffee Rust
Death caps: Poisonous mushrooms as Heterothallic type: describes when
they are fatal and cause death only one mating type is present in an
Downy Mildews: White, powdery individual mycelium
growth on leaves, new shoots and other Heterothallism: It is a phenomenon
plant parts in which two different sexes
Ergot: Drug obtained from sclerotia of participate in sexual reproduction.
Claviceps purpurea Homothallic: describes when both
Facultative anaerobes: organisms mating types are present in mycelium
that can perform both aerobic and or hypha fungal filament composed of
anaerobic respiration and can survive one or more cells

218 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Homothallism: It refers to the gametangial copulation or markedly
possession of male and female sexes on unequal gametic fusion.
the same thallus. Orcein: A purified extract of lichen
Iceland mass: Nutritive species of uysed for microscopic observation
Cetraria islandica used in cakes and Orcein: A purified extract of lichen
food uysed for microscopic observation
Iceland mass: Nutritive species of Penicillin: Antibiotic extracted from
Cetraria islandica used in cakes and Penicillium notatum used against
food gram+ bacter ia and many other
Isidia: Corolloid outgrowths of lichens diseases.
consists of external cortical layer and Psiloc ybin: Extracts of Psilocybe
internal algal layer semilanceata and Gymnopilus junonius
Isidia: Corolloid outgrowths of lichens used for hallucinogenic properties.
consists of external cortical layer and Pycnia: It is a specialized structure
internal algal layer produced by haploid mycelium formed
Macrocyclic Rust: The life cycle from germination of basidiospore
involves 5 different spores with Pycnidiospore or spermatia:
different functions pycniospore leading to the production
Magnaporthe oryzae: The asexual of a dikaryotic mycelium by fertilization
stage of Rice blast of a receptive hypha of the opposite
Mycelium: mass of fungal hyphae mating type

Mycorrhiza: The relationship Pythium: Causative agent of Damping off


between fungi and plant roots which disease
is essential for the productivity of Reindeer moss: Cladonia sylvatica
farmland. which is used as food for man and cattle,
Obligate aerobes: organisms, such as reindeer
humans, that must perform aerobic Reindeer moss: Cladonia sylvatica
respiration to survive which is used as food for man and cattle,
Obligate anaerobes: organisms that reindeer
only perform anaerobic respiration and Rhizenes: Rhizoid like outgrowths of
often cannot survive in the presence the lichen thallus
of oxygen Rhizenes: Rhizoid like outgrowths of
Oospores: It is a sexually produced the lichen thallus Roquefor t or
spore develops from unequal Gorgonzola and Stilton

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Saprobe/ saprophyte: organism Toadstools: Poisonous mushrooms
that der ives nutr ients from are called toadstools.
decaying organic matter; Uredospore: The only type of spiny
Soredia: Small rounded bodies and brick red spores in the rust fungus
from one to several algal bodies life cycle that are capable of infecting
closely surrounded by fungal the host on which they are produced
hyphae which is containing two unfused,
Soredia: Small rounded bodies haploid nuclei in one cell formed on
from one to several algal bodies individual stalks
closely surrounded by fungal Wilt: The infection that spreads up
hyphae into the stems and leaves and restricts
Taphrina deformans: Pathogen of water flow causing the foliage to turn
Leaf Curl yellow

Taphrina deformans:Pathogen of Wilt: The infection that spreads up into


Leaf Curl the stems and leaves and restricts water
flow causing the foliage to turn yellow
Teleutospore: the only form of
dark brown or black stalked, two Zygospores: The structure formed in
celled, spindle shaped spore with zygosporangia of fungi, after the
thick, black and smooth wall fusion of specialized budding
undergoes karyogamy and meiosis structures, from mycelia of the same
in which is able to over winter (homothallic fungi) or different mating
independently of a host. types (heterothallic fungi)

220 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Unit - 4
ALGAE - The Autotrophic Thallophytes

CONTENTS LEARNING OUTCOMES:


At the end of this unit learner should be
1. General
characteristics of able to:
Algae and Fritsch 1. Summarize the important criteria for
classification. identification algae belong to different
classes.
2. Thallus
2. Appraise the Fritsch classification and
organization and
life cycles in Algae. also the diversity of algae.
3. Explain the range of thalli and life
3. Spirogyra and cycles in algae.
Polysiphonia
4. Compare and contrast the general
4. Economic characteristics of algae, with special
importance of reference to green and red algae.
Algae. 5. Judge the ecological-economic value
and impact of algae in human welfare.
6. Recognize the web-resources to acquire
additional knowledge on algae.

General Characteristics of Algae and Fritsch


Chapter -4.1
Classification
4.1.1 General Characteristics of Algae :

Learning outcomes: At the end of the chapter you should be


able to:
1. Describe, discuss and debate the prominence of
photosynthetic pigments, flagella and reserve food in algal
classification.
2. Memorize the Fritsch system of algal classification;
illustrate the typical features of different algal classes in it.

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
Algae are the primary producers in all aquatic ecosystems.They
perform about 50% of global photosynthesis and release 50% of
the total Oxygen on the planet earth. Thus, every second molecule
of oxygen inhaled by humans is produced by an alga; similarly,
every second molecule of carbon dioxide exhaled is reused by an
alga. Over 90% of the species of marine plants are algae. Algae
gave are directly responsible for all seafood and indirectly
responsible for all land foodas they were progenitors of land plants.
Vast numbers of phytoplankton cells are initially food for marine
and freshwater webs and expressed as pasturage of the seas. Fossil
records of algae date back to three billion years, indicating their
origin in the prolific Precambrian era. Fossil fuels came from
Cretaceous deposits of marine algae. It is a fact that about 3.5
billion years ago, the origin of blue green algae paved the way for
the beginning of other life forms by the production of oxygen in
the earth’s atmosphere. The evolution of eukaryotic organisms
began after the oxygenic photosynthesis by algae. Blue green algae
not only produce oxygen but also play a vital role as primary
producers in the food chains and food webs; they also fix
atmospheric nitrogen needed for other living organisms, thus
support life on the planet.Algal growth (cell numbers) has been
affected by season, temperature, amount of sunlight penetrating
the water column. The amount of available inorganic nutrients,
competition from other algae and aquatic plants, and duration of
water retention (residence time) in the water bodies may also alter
algal growth.

   


  



222 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Introduction: The branch of botany deals with algae is Phycology
or Algology. Algae (singular alga) are eukaryotic or prokaryotic,
photoautotrophic and predominantly aquatic plants. The algae are
thallophytes possess a simple and undifferentiated plant body,
thallus. Algae are primitive plants, but display high order of
diversity in their habit, colour (pigment), reproduction and
lifecycles. All classes of algae consist of Chlorophyll-a as primary
photosynthetic pigment. The reproductive structures are mostly
unicellular devoid of a sterile jacket and hidden, a characteristic
feature of cryptogams.
Based on the magnitude (size), algae are broadly classified
into microalgae and macroalgae. Microalgae are further
categorized into two general groups namely phytoplankton and
periphyton. Phytoplankton live suspended in lentic and lotic water
bodies, whereas Periphyton is attached to substrata like rocks,
sediment, plant stems, and aquatic organisms. Marine macroalgae,
seaweeds are generally benthic and rarely pelagic (Sargassum
muticum, S. fluitans, and S. natans).

Thallus and cell structure: In algae, the plant body is a thallus


devoid of differentiation into root, stem and leaves. The thallus
structurally ranges from simple unicellular to complex multicellular
form. The cells in all classes of algae are eukaryotic, except
Myxophyceae. Algal cells are flagellated (exceptRhodophyceae and
Myxophyceae) and motile; In Myxophyceae members, the
prokaryotic cells are devoid of membrane bound cell organelles.
Eukaryotic algal cell possesses well defined cell wall composed of
polysaccharides; plasma membrane made of lipo-proteins. It consists
of nucleus, mitochondria, Golgi bodies, endoplasmic reticulum,
plastids etc. The nucleus is surrounded by a bilayered nuclear
membrane, contains chromosomes and nucleolus in nucleoplasm.

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


    



        

  


 
      

       
 

    
        

      



       
224 APSCHE : SEMESTER-I BOTANY TEXT BOOK
Chloroplast (plastid) consists of a fluid space called stroma, and is
enveloped by two cell membranes. The stroma has thylakoids,
circular dsDNA, 70S ribosomes and reserve food material. The
thylakoids are membrane bound sacs; they consist of photosynthetic
pigments (refer pigments of the chapter)
Pigments: Algae have a great diversity of photosynthetic pigments,
perhaps no other group of plants display such diverse colours.
Pigments are one of the significant criteria for classification of algae.
Depending on the predominant pigment, different classes of algae
are green (Chlorophyceae), yellow (Xanthophyceae), diatoms
(Bacillariophyceae), brown (pheophyceae), red (Rhodophyceae),
blue-green (cyanophyceae) and so on and so forth. Pigments are
present in specialized plastids, chromophores. In general, the algal
pigments fall into 3 major categories, namely chlorophylls,
carotenoids and phycobilins.
A. Chlorophylls: Chlorophylls are green pigments. They are
hydrophobic, lipophilic and chlorin compounds, soluble in organic
solvents such as ethyl alcohol and acetone. They absorb red and
blue light and reflect green colour; impart green colour to algae and
also embryophytes.

B. Carotenoids: Carotenoids are orange (carotenes) or yellow


(xanthophylls) coloured pigments. They are hydrophobic, lipophilic
and tetra-terpenoid compounds, soluble in organic solvents like
acetone, chloroform and petroleum ether. These pigments are
present in close association with chlorophylls, act as accessory
pigments and protect chlorophylls from photo-oxidation. Carotenes
are hydrocarbons (C 40H 56), while xanthophylls (C 40H56O 2) are
carotenols. Carotenes absorb UV, violet and blue light; scatter orange
or red or yellow wave lengths. Absorption spectrum of xanthophylls

APSCHE : SEMESTER-I BOTANY TEXT BOOK 225


A. Chlorophylls:
Chlorophyll Molecular formula Taxa in which present
Chlorophyll-a C55H72O5N4Mg Common in all classes of
Algae
Chlorophyll-b C55H70O6N4Mg Chlorophyceae and
some Cyanophyceae
Chlorophyll-c1 C35H30O5N4Mg Phaeophyceae ,
Bacillariophyceae and
Dinophyceae
Chlorophyll-c2 C35H28O5N4Mg Phaeophyceae,
Bacillariophyceae
and Dinophyceae
Chlorophyll-d C54H70O6N4Mg Rhodophyceae and some
Cyanophyceae
Chlorophyll-e C54H70O6N4Mg Xanthophyceae
Chlorophyll-f C55H70O6N4Mg Some Cyanophyceae
(e.g., Stromatolites)

B. Carotenoids:
Algal class Dominant Dominant
Carotene Xanthophylls
Chlorophyceae, α-, β-carotene Lutein
Xanthophyceae α- carotene Diadinoxanthin,
diatoxanthin,
heteroxanthin,
vaucheriaxanthin
ester
Chrysophyceae β-carotene Fucoxanthin
Bacilloriophyceae e-carotene Fucoxanthin
Phaeophyceae α- and β-carotene Fucoxanthin
Rhodophyceae α-carotene Myxoxanthin
Cyanophyceae α- carotene, Myxoxanthin,
myxoxanthophyll

226 APSCHE : SEMESTER-I BOTANY TEXT BOOK


is similar to carotenes, but scatter yellow colour. Five types of
carotenes and 20 types of xanthophylls are reported in algae so far.

C. Phycobilins: Phycobilin pigments are mainly phycocyanin (blue)


and phycoerythrin (red). They are water soluble and bound with
proteins as phycobiliproteins. They are present in macromolecular
complexes, phycobilisomes. These accessory pigments absorb green,
orange, yellow and red lights; scatter blue (phycocyanin) or red
(phycoerythrin) light. Six types of phycobilins are reported in algae.

Flagella: Flagella are the microscopic extra cellular appendages


meant for locomotion, drive the cell in the water by their movement.
The flagella are one of the important characteristics to classify the
algae. The number, length, appendages and position of flagella vary
in different classes of algae. Flagella are found in vegetative or
reproductive cells or in both in most of the algal classes except in
the members of Rhodophyceae, Myxophyceae and

Algal class Phycobilin present


Rhodophyceae r-phycoerythrin and r-phycocyanin,
Myxophyceae c-phycocyanin and c-phycoerythrin,
Cryptophyceae Phycocyanin or phycoerythrin, but never both in
a species

Know more: Structure and value of algal pigments: Algal


pigments are being exploited commercially; they have several
niches of applications as natural dyes, antioxidants and precursors
of vitamins. A chlorophyll molecule consists of a head (tetra-
pyrrole ring) and a phytol tail (a hydrocarbon chain). Carotenoids
have a backbone chain with 40 carbons arranged into 8 isoprene
molecules.â-carotene from Dunaliella spp. and astaxanthin from
Haematococcus spp. are of great value as antioxidants and vitamin-

APSCHE : SEMESTER-I BOTANY TEXT BOOK 227


A precursors. Phycobilins are auto-fluorescent pigments,
chemically aliphatic with an open chain of 4 pyrrole rings.They
are frequently used in immunology research as chemical tags to
bind antibodies. Phycobilins are much related to chromophore in
phytochrome, a light detecting pigment of higher plants. They
are structurally similar to bilirubin, a piment in liver of mammals.
Extract of Porphyridiumcruentum is an ingradient of face creams,
useful to retain moisture and to impart colour, owing to its
phycobilins and polysaccharides.

        


 
        






        


  

Activity: Collect algal samples from the water bodies in your locality,
extract the photosynthetic pigments - chlorophylls and carotenoids
using acetone and petroleum ether;phycobilins using hot water;
separate them by paper chromatography in your laboratory

228 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Prochlorophyceae.In diatoms flagella are absent, but in some genera
of diatoms male gametes possess one or rarely two flagella.
Pleuronematic f lagella are of 3 types i.e., pantonematic,
pantoacronematic and stichonematic based on the arrangement of
flimmers. In pantonematic flagellum, mastigonemes are arranged
in two opposite rows, while stichonematic f lagellum has
mastigonemes on only one side. Pantonematic flagellum with
terminal fibril is called pantoacronematic.
A. Morphology of flagella: Based on external appearance flagella
are 2 types i.e., whiplash and tinsel (Fig.4.1.1). Whiplash or
acronematic flagellum is smooth, devoid of hairy outgrowths. Tinsel
or pleuronematic flagellum consists of one or more rows of lateral
hairs, flimmers (mastigonemata). The flimmers may be solid or
tubular; increase the diameter of the flagellum and also amplify its
stroke.
Based on length and morphology of the flagella, algae are
of following types: Isokont (e.g.,Chlamydomonas) possessing two
flagella of equal length and appearance; while anisokont (e.g.,
Chara), having two similar flagella of unequal length. A
Heterokont has two dissimilar flagella (one whiplash and other
tinsel). Members of Chlorophyceae are isokonts with two whiplash
flagella. Cryptomonas is an isokont with two pleuronematic flagella.
Phaeophyceae and Xanthophyceae members are heterokonts with
one whiplash and another tinsel flagellum.
B. Structure of the flagellum: The transverse section of a
flagellum under the Transmission Electron Microscope (TEM) has
9+2 arrangement. It consists of two central microtubules
surrounded by a ring of nine doublet microtubules, and bounded
by a plasma membrane. This membrane is continuous with the
plasma membrane of the cell (Fig.4.1.2).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 229


  

       

    


Know more:The flagellum in algae passes across a channel,


the flagellar collar in cell wall. The two central microtubules
culminate at a dense plate, after entering the cell body. On
the other hand, the nine peripheral doublets usually pick up
an extra structure after linger into the cell; thus, transform
into triplets. A basal body is seen at the base of the flagellum;
usually, some microtubular roots or striated fibrillar roots are
attached to it. The microtubular roots extend into the
cytoplasm. Groups of fibres with striations are present along
the entire length of the striated fibrillar roots. A flavin-like
auto-fluorescent substance occurs in the flagella of euglenoids
and brown algae; it appears to be photoreceptive, useful for
phototaxis. Thus, flagellum in such algae functions as sensory
organ, in addition to its normal function.

Fig. 4.1.1 Morphology of flagella in algae


(a) Acronematic (b) Pantonematic
(c) Pantoacronematic (d) Stichonematic

230 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig.4.1.2 Ultrastructure of an algal flagellum
Source: https://istudy.pk/algae-flagellation/

C. Number of flagella: Motile algal cells are biflagellate (e.g.,


Chlamydomanas) or tetraflagellate (e.g.,Tetraselmis Cladophora) or
multiflagellate (e.g., Oedogonium,Vaucheria) (Fig.4.1.3).
Chrysophytes and euglenoids typically consists of two flagella, in
which one is well developed and the other is vestigial.

(a) Biflagellate (b) Quadriflagellate (c) Multiflagellate

Fig.4.1.3 Number of flagella

D. Position of flagella: In Chlorophyceae, flagella are attached at


anterior end of the cell, apical; but in charophytes, two flagella may
be inserted laterally or sub-apically. The zoospore in Derbesia and

APSCHE : SEMESTER-I BOTANY TEXT BOOK 231


male gamete in Oedogonium are stephanokonts; consist of a ring of
sub-apical flagella at anterior end. In dianoflagellates, a flagellum
fits into transverse girdle, while another flagellum comes out from
the longitudinal sulcus. Chrysophyceae members have 2 flagella, in
which the tinsel flagellum at the anterior end is parallel to the cell
axis, while the whiplash flagellum is perpendicular to the tinsel
flagellum. The flagella in all members of Phaeophyceae and some
algae of Xanthophyceae are lateral in position (Fig.4.1.4).

(a) Apical (b) Lateral

Fig.4.1.4.a. position of flagella

Fig. 4.1.4.b. Types of flagella


and their position in different
classes of algae.
A. Crytophyceae,
B. Xanthophyceae,
Phaeophyceae, Chrysophyceae,
Raphidiophyceae,
C. Bacillariophyceae,
D. Prymnesiophyceae,
E. Charophyceae,
F. Dinophyceae,
G. Euglenphyceae,
H. Eustigmatophyceae,
I,J Chlorophyceae.

232 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Reserve food material: Reserve food material is an important
characteristic of algae and is specific to each class of algae. Thus, it
is a useful principle for algae identification and classification. In
general, reserve food is polysaccharide; but fats and oils are also
found in the cells of Bacillariophyceae, Xanthophyceae and
Dinophyceae.
Pyrenoids made of densely packed proteinaceous fibres are
seen within or on the surface of chromatophores in almost all algal
classes (except Cyanophyceae).
Pyrenoids are always traversed by chromophore lamellae
(except in Phaeophyceae). A chromatophore may contain one
(Chlamydomonas) or many (Cladophora) pyrenoids. The
photosynthetic reserves of algae generally accumulate in pyrenoids.
In Chlorophyceae, proteinaceous core of pyrenoid is encircled by a
layer of starch plates; found as conspicuous structure within the
chloroplast. In most of the algae, the pyrenoids may be transient
structures or do not associate with reserve food.
Class Reserve food
Chlorophyceae Starch (polymer of amylose and amylopectin)
Euglenophyceae Paramylum(laminarin like polysaccharide)
Xanthophyceae Oil or leucosin
Bacillariphyceae Leucosin and chrysolaminarin
Phaeophyceae Laminarin and mannitol
Chrysophyceae Chrysolaminarin (similar to laminarin)
Rhodophyceae Floridean starch (floridoside) and
mannoglycerate. (resembles glycogen in fungi
and animals).
Cyanophyceae Cyanophycean starch (resembles floridean starch)

Activity: Collect Spirogyra, Cladophora or some other green alga from a


local water body – stain with iodine, observe the reserve food and pyrenoid
under microscope.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 233


         
         
                  
      
       
  
   


* * * * * * * *
Reproduction: Algae reproduce by vegetative, asexual and sexual
methods.
A. Vegetative reproduction: Vegetative reproductionin algae
occur under favourable conditions. It takes place in various ways.
In unicellular algae (Chlamydomonas, Chlorella etc.,) cell division is
the prime method of reproduction. In filamentous forms (Zygnema,
Oedogonium, Cladophora etc.,) thallus breaks into fragments and each
of those pieces develops into an independent thallus. Multicellular
adventitious branches formed from the thalli of some Phaeophyceae
members (Dictyota, Fucus), and develop into new thalli on
separation. Protonema, tubular structures formed on thalli of
Cladophora and Chara get detached from parent thallus and advance
into new ones. Chara produces starch filled organs like tubers, bulbils
and amylum stars for vegetative reproduction. Filamentous green
algae like Ulothrix, Oedogonium and Cladophora produce thick walled
akinetes with copious reserve food during unfavourable conditions;
they undergo resting period and germinate into new thalli on return
of favourable conditions.
B. Asexual reproduction: Most of the algae produces asexual
reproductive organs, sporangia. The protoplasts in cells of those

234 APSCHE : SEMESTER-I BOTANY TEXT BOOK


sporangia may divide mitotically or meiotically to produce spores.
Asexual reproduction also occurs under favourable conditions; the
asexual spores germinate to form new individuals. In Oedogonium
and Ulothrix, motile and naked zoospores are formed from
zoosporangia; conversely non-motile tetraspores are formed from
tetrasporangia in Polysiphonia. Carposporangia in some red algae
produce haploid (Batrachospermum) or diploid (Polysiphonia) asexual
spores, carpospores. Those spores germinate and develop into new
thalli. In Vaucheria, Botrydium and Ulothrix vegetative cells
metamorphoses to form one to many aplanospores; which
germinate into new thalli after a period of rest. Hypnospores in
Pediastrum and autospores in Scenedesmus are also meant for asexual
reproduction. Some Rhodophyceae members (Porphyra,
Batrachospermum etc.,) produce naked and non-motile monospores
during asexual reproduction. The protoplasts of vegetative cells in
Asterocystis and Ectocarpus metamorphoses into neutral spores for
asexual reproduction.

C. Sexual reproduction: Algae belong to most of the classes


reproduce sexually, prior to the start of adverse conditions or as a
part their life cycle for alternation of generations. In general, male
gamete fertilizes the female gamete to form a diploid zygote. In
species with zygotic meiosis, reduction division in zygote gives
haploid spores that restore the gametophytic phase of life cycle.
Sexual reproduction is completely absent in Myxophyceae. In algae,
the fusion of gametes may be isogamous (Chlamydomonas),
anisogamous (Ectocarpus) or oogamous (Sargassum).

Life cycles in algae: Algae have diverse types of life cycles and
they may vary from one taxon to another within a class.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 235


4.1.2: Fritsch classification:

Linnaeus (1754) coined the term ‘Algae’to a group of plants including


the Hepaticae. A.L.de Jussieu (1789) separated the algae from other
plants and gave them a special status. S.L.Endlicher (1836) included
algae and fungi together in the division, Thallophyta.
Hecalledthallophytes as non-embryophytes, considered equivalent to
embryophytes, Thallophytes are a polyphyletic group, includes diverse
organisms like myxomycetes, bacteria, fungi, algae, and lichens.A.W.
Eichler (1883) kept Thallophyta, Bryophyta, and Pteridophyta
divisions in Cryptogamae, (non-flowering plants; plants with hidden
sex organs) a sub-kingdom of the plant kingdom. About 36,000 species
of algae are reported so far, but they represent 17% of the estimated
diversity.

Introduction: Felix Eugen Fritsch, a British Biologist, regarded


as The Father of Phycology. He proposed a comprehensive
classification of algae in the book, ‘The Structure and Reproduction
of Algae’, published in 1935. The important criteria for his algal
classification are pigments, flagella, reserve food material and mode
of reproduction. He considered algae as a division and divided into
following 11 classes:
1. Chlorophyceae: Green algae, mostly occur in fresh water and
some in marine water. The thallus may be unicellular, colonial,
filamentous or parenchymatous. Chloroplasts enclosed by double
membranes, devoid of ER; thylakoids present in stacks of 2-6 or
more. They possess Chlorophyll-a, Chl.-b and lutein as chief
photosynthetic pigments. True starch is the food reserve. Flagella
are generally whiplash type and isokontic. Reproduction is by
vegetative, asexual and sexual methods; sexual reproduction may
be isogamous, anisogamous or oogamous (Chlamydomonas, Vovox,
Oedogonium, Fritschiella).
2. Xanthophyceae: Yellow-green algae, mostly fresh water forms
and a few are from marine habitats. The thallus is globose or tubular

236 APSCHE : SEMESTER-I BOTANY TEXT BOOK


and coenocytic. Chloroplasts have two additional membranes of ER
the outer one is continuous with the outer nuclear membrane;
thylakoids are in stacks of three. The principal pigments are
Chlorophyll-a, Chl.-e, diadinoxanthin, diatoxanthin and
heteroxanthin. Photosynthetic products are leucosin (â-1,3-linked
glucan), fats and oils. Heterokonts with one whiplash and one tinsel
flagella. Reproduction is through vegetative and asexual methods;
sexual reproduction is rare, may be isogamous or oogamous
(Chlorogloea, Botrydium. Vaucheria).
3. Chrysophyceae: Golden-brown algae are unicellular naked
forms, live mostly in fresh water bodies of colder regions, a few
occur in marine water. Chloroplasts have two additional membranes
of ER, the outer one is continuous with the outer nuclear membrane;
thylakoids are in stacks of three. The major pigments are
Chlorophyll-a, Chl.-c1, Chl.-c2 and fucoxanthin. Heterokonts with
anterior tinsel and posterior whiplash flagella. Reproduction occurs
by formation of typical silicious statospores (stomatocysts); sexual
reproduction is seldom and is isogamous (Ochromonas,
Paraphysomonas, Chrysoamoeba).
4. Bacillariophyceae: Diatoms are ubiquitous, grow in fresh and
marine water and also terrestrial habitats. Chloroplasts have two
additional membranes of ER, the outer one is continuous with the
outer nuclear membrane; thylakoids are in stacks of three. These
unicellular algae contain Chlorophyll-a, Chl.-c1. Chl.-c2,
fucoxanthin, diadinoxanthin and diatoxanthin major pigments. The
major food reserve is chrysolaminarin and lipids. The male gametes
are flagellated, consist of a single hairy flagellum. Vegetative
reproduction by mitotic cell division is common. The sexual
reproduction in pennate diatoms is by isogamy of auxospores, while
in centric diatoms auxospores fuse in 5 different ways. Eg: Centric
diatoms (Thalassiosira, Cocconeis) and Pennate diatoms (Melosira,
Nitzschia).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 237



   

       






 

    
   

  
     

       
 
 

       
 

238 APSCHE : SEMESTER-I BOTANY TEXT BOOK


5. Cryptophyceae: Cryptomonads are unicellular algae, they occur
in marine, brackish and fresh water.Chloroplasts have two additional
membranes of ER; thylakoids present in pairs. Chlorophyll-a, Chl.-
c, α-caroene and diatoxanthin are the principal pigments. Starch
like carbohydrates are the reserve food material. Flagella are tinsel
type, may be equal and or slightly unequal. Binary fission is quite
common, sexual reproduction has not been reported (Cryptomonas,
Hemiselmis).
6. Dinophyceae: Dianoflagellates are single celled and marine
planktonic forms. Chloroplasts have two additional membranes of
ER not continuous with nuclear envelop; thylakoids are in stacks of
three. The principal pigments are Chlorophyll-a, Chl.-c2, peridinin
and neo-peridinin. Oil and starch are the food reserves. Two flagella
of unequal length lie in two grooves of the cell. Dinoflagellates
reproduce by binary fission; rarely sexual reproduction by isogamy
and production of planogametes are also reported (Ceratium,
Stylodinium, Peridinium, Pyrodinium)
7. Chloromonadineae: Raphidophytes, live in both fresh and
marine water. They are unicellular forms without cell walls and
contain numerous ellipsoid chloroplasts. Chlorophyll-a, Chl.-c1.
Chl.-c2, â-carotene and diadinoxanthin are the major pigments. Oil
is the food reserve and flagella are of equal size. Reproduction is by
longitudinal division; sexual reproduction is not yet reported
(Chlorinomonas, Psammamonas, Heliorapha, Fibrocapsa).
8. Eugleninae: Euglenoids are unicellular, mostly fresh water
forms. Cells contain many chloroplasts surrounded by 3 membranes.
The principal pigments are Chlorophyll-a, Chl.-b, β-carotene and
antheraxanthin. Paramylon is the principal food, chrysolaminarin
may also present. Flagella are usually two, hairy and unequal (one
is not emergent). They reproduce solely by binary fission; sexual
reproduction is not reported (Euglena).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 239


   

    
    
 
 


   
    


  
  

       


        
     
  



   

240 APSCHE : SEMESTER-I BOTANY TEXT BOOK


9. Phaeophyceae: Brown algae are exclusively marine and cold-
water loving forms; a few species reported from fresh water habitats.
Chloroplasts have two additional membranes of ER, the outer one
is continuous with the outer nuclear membrane; thylakoids are in
stacks of 2 to 6. Thallus is of filamentous or complex structure;
unicellular forms are absent. The major pigments are Chlorophyll-
a, Chl.-c1 and Chl.-c2 and fucoxanthin. Laminarin and mannitol
are the reserve food material. Reproduction occurs through
vegetative, asexual and sexual methods; sexual reproduction is
isogamous to oogamous in different species (Ectocarpus, Punctaria,
Sargassum, Laminaria).
10. Rhodophyceae: Red algae are predominantly marine; some live
in estuarine and fresh water bodies. Chloroplasts contain bi-layered
envelope; chloroplast ER is absent and thylakoids are unstacked.
Thallus is unicellular, pseudo-parenchymatous or parenchymatous.
The main photosynthetic pigments are Chlorophyll-a, Chl.-d and r-
phycoerythrin. The chief food reserve is floridean starch. Flagella
are absent. Reproduction is by vegetative, asexual and sexual
methods; sexual reproduction is oogamous (Bangia, Porphyra,
Palmaria, Gracilaria).
11. Myxophyceae (Cyanophyceae): Blue Green Algae (BGA) are
ubiquitous, live on every moist surface. Thallus is unicellular to
filamentous. Plastids are absent. The chief pigments are
Chlorophyll-a, myxoxanthin, myxoxanthophyll and c-phycocyanin.
Flagella are absent. Reproduction occurs by cell division; sexual
reproduction is not reported (Spirulina, Nostoc, Anabaena,
Gloeocapsa, Oscillatoria).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 241


    
  



 





       


       
     
   


242 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Prof. F.E.Fritsch (1879-1954), the British Phycologist published
two volumes on ‘The Structure and Reproduction of the Algae’;

publications are related to ecological, taxonomic, classificatory,


morphological and evolutionary aspects of phycology. He worked
in University of Munich, University College London, and the
Royal Botanic Gardens (Kew) in various capacities. He was
instrumental in establishing ‘Freshwater Biological Association
(FBA)’ in 1929; and ‘The Culture Collection of Algae and Protozoa’
in 1931. In 1912, Fritsch started to put illustrations of freshwater
algae onto foolscap sheets of paper; after the death of Fritsch,
Dr.J.W.G.Lund established it as ‘The Fritsch Collection of
Illustrations of Freshwater Algae’ consisting of about 20,000
collections.Prof.M.O.P.Iyengar (The Father of Phycology in India)
is one of the research students of Prof. F.E.Fritsch; who identified
a new alga, Fritschiella (named after).

Source:https://fmp.conncoll.edu/

Source: https://www.jstor.org/stable/769202

APSCHE : SEMESTER-I BOTANY TEXT BOOK 243


Chapter -4.2 Thallus Organization and Life Cycles in Algae

Learning outcomes: At the end of the chapter you should be able


to:
1. Explain the diversity in structure of thallus in algae and
understand evolutionary tendencies.
2. Discuss various types of life cycles in different taxa of algae
and compare with other groups of plants.

4.2.1.Thallus organization in Algae:

Thallus (plural: thalli) is derived from a Latinized Greek word,


thallos (= a green shoot). The thallus is a simple plant body
without root, stem, and leaves, as opposed to a complex plant
body in spermaphytes. Algae, fungi, lichens, and liverworts in
Bryophyta possess such simple plant bodies. The thallus in fungi
is often referred to as a mycelium. The gametophyte in
Pteridophytes with a low level of cellular organization is known
as prothallus.

Introduction: In algae both vegetative and reproductive plant


bodies are thalloid. The thalli of algae show greater diversity in their
habit, ranging from simple unicellular to complex multicellular
bodies. The unicellular algae are motile or non-motile. Whereas the
multicellular thalloid forms have many variations in nature and
arrangement of cells. The size of algal thalli ranges from one micron
(e.g., Micromonas pusilla) to 65 meters (Macrocystis sp.). The thallus
in algae are of three types i.e., unicellular, colonial and multicellular
based on the number of cells motility, and arrangement.

Unicellular habit: All classes of algae, except the Phaeophyceae,


contain unicellular forms. The cell of a unicellular alga performs all

244 APSCHE : SEMESTER-I BOTANY TEXT BOOK


the biological activities. Unicellular forms can be classified into
motile, rhizopodial, and non-motile types (Fig.4.2.1).

A.Motile forms: This type of algae moves with the help of flagella.
The unicellular algae in most classes are flagellated (except
Cyanophyceae, Rhodophyceae, and Bacillariophyceae). In general,
flagellated forms have a cell wall; but in Euglena, pellicle is present
instead of cell wall. In these forms number and size of the flagella
varies in different taxa.

Fig.4.2.1 Unicellular habit

(a) Chlamydomonas (b) Chlorogonium

(c) Chromulina (d) Tetraselmis

(e) Euglena (f) Chrysamoeba

(g) Synechococcus (h) Chlorella

(i) Cystodinium (j) Porphyridium


Source:https://commons.wikimedia.org/wiki/Category:Algae
https://libguides.astate.edu/oer/images , https://www.inaturalist.org/taxa/50863

APSCHE : SEMESTER-I BOTANY TEXT BOOK 245



  
    

  
  
 
 


Fig.4.2.2 Colonial habit (a) Pandorina (b) Synura (c) Volvox (d) Pediastrum
(e) Scenedesmus(f) Hydrodictyon (g) Phaeocystis (h) Dinobryon
Source:https://commons.wikimedia.org/wiki/Category:Algae
https://libguides.astate.edu/oer/images
https://www.inaturalist.org/taxa/50863

246 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Characteristic Example

Number 1(Chromulina), 2(Chlamydomonas) and 4(Tetraselmis)

Length Equal(Chlorophyceae) and unequal(Dinophyceae)

Morphology Isokontic (Chlorogonium)andheterokontic (Heterochloris).

B.Rhizopodial forms: These algae (e.g., Chrysamoeba, Rhizochrysis)


lack cell walls and flagella; but show amoeboid movement by means
of cytoplasmic projections. According to Fritsch (1935), flagellate
forms evolved into rhizopodial forms by losing flagella.

C. Non-motile forms: Fritsch (1935) named these algae as coccoid


forms. This type of thallus is reported in many algal classes. Round
(1973) categorized them under protococcoidal type. Movement in
these planktonic algae is by water currents only. Examples:
Synechococcus(Myxophyceae), Porphyridium (Rhodophyceae),
Chlorella (Chlorophyceae), Cystodinium (Dinophyceae), Characiopsis
(Xanthophyceae).

Colonial habit: A group of cells with similar structure and function


are aggregated in a common mucilage matrix to form a colony
(Fig.4.2.2). The colonial habit is either globose or net like. Volvox
is an example to globose colony, in which flagellated cells are
connected by cytoplasmic connections; to avoid breaking the colony
into pieces or segments. Hydrodictyonis a net-like colonial form, in
which 5-6 non-flagellated cells are connected forming pentagonal
or hexagonal rings. Colonial algae are classified into four types:

A. Coenobial colony: This type thallus has a definite number of


cells arranged systematically. The number of cells in this colony is
constant, does not increase with growth, but the size of cells and
colony increases. The colony may be motile with flagellated cells

APSCHE : SEMESTER-I BOTANY TEXT BOOK 247


     
    

         



          
      

        
 


       
    

 
    


 
    
  
 


248 APSCHE : SEMESTER-I BOTANY TEXT BOOK


(Pandorina, Eudorina, Synura); or non-motile with non-flagellated
cells (Scenedesmus, Pediastrum). Aggregation of similar motile cells
might have resulted in a coenobial colony during the course of
evolution.
B. Palmelloid colony: In this type of habit, non-motile cells
aggregate in an amorphous gelatinous or mucilage matrix. The
mucilage matrix may be secreted either by protoplasts of cells or by
gelatinization of the cell membranes. The number, shape, and size
of the cells of the colony are not determinate. Cells in this colony
perform independent functions, though aggregated in a common
mucilage matrix. This type of thallus organization is either
permanent or temporary. Permanent palmelloid colonies are seen
in Tetraspora (Chlorophyceae), Gleochloris (Xanthophyceae),
Phaeocystis(Chrysophyceae) and Microcystis (Myxophyceae).
Chlamydomonas spp. forms a temporary palmelloid colony.
C. Dendroid colony: Cells are attached by mucilage to form a
microscopic tree like (dendroid) structure The number, shape and
size of the cells are not specific. Cells have polarity and consist of
mucilage at their narrow bases. (Dinobryon, Chrysodendron,
Ecballocystis, Prasinocladus)
D. Rhizopodial colony: The cells in this colony are connected
through rhizopodia (Chrysidiastrum).
Filamentous habit: Cells are arranged one above the other in a
definite order to form a uniseriate filament (Trichome). This type
of forms has evolved from motile unicellular habits. In many cases,
the motile swarmer attaches to a substratum, secretes a cell wall
around it. Cells formed from cell divisions remain attached to each
other form a chain (filament). The filamentous thallus may exist in
four forms, unbranched, branched, pseudo-branched
orheterotrichous (Fig.4.2.3).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 249




      
 
    


                             

   
   

  


        
  


In Coleochete scutata, the prostrate system forms a discoid thallus,
the erect system is absent. In Draparnaldiopsis, the prostrate system
is suppressed and the erect system is elaborated. Stigeoclonium has
well developed prostrate and erect systems. Prof.F.E.Fritsch (1935)
states that heterotrichous algae belong Chaetophorales may be
the progenitors for land plants. Fritchiella tuberosa, living in moist
terrestrial habitats has well-developed prostrate and erect systems;
Prof. M.O.P.Iyengar (1932) opined that this alga might have led to
the evolution of Bryophytes.

250 APSCHE : SEMESTER-I BOTANY TEXT BOOK


A. Unbranched filamentous forms: Thallus is made of many
cells arranged in a single row. The filament attaches to the
substratum with a specialized basal cell, hold-fast (Ulothrix), or free-
floating (Zygnema). Members of Chlorophyceae (Oedogonium,
Mougeotia), Myxophyceae (Oscillatoria, Phormidium) show this type
of thallus.

Fig. 4.2.3 Filamentous habit


(a) Zygnema (b) Cladophora
(c) Scytonema
Source: https://alchetron.com/
Search#search=Zygnema
https://alchetron.com/Cladophora#cladophora-
9c88b8bc-61d3-4638-a20e-18550c22ff3-resize-
750.jpeg
http://cfb.unh.edu/phycokey/phycokey.htm

B. Branched filamentous forms: This type of algae is evolved


from uniseriate unbranched forms. This habit might have evolved
from formation of transverse septa subsequent to lateral
outgrowths in the cells of an un-branched. This habit can be seen
in Cladophora (Chlorophyceae), Phaeothamnion (Chrysophyceae),
and Callithamnion (Rhodophyceae).

C. Pseudo (false) branching forms: In this type of thallus, due


to the degeneration of an intercalary cell break the trichome into
two fragments and the two fragmentsprojectfrom mucilage sheath
give the appearance of branching (Scytonema and Tolypothrix of
Myxophyceae).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 251


D. Heterotrichous forms:This habit is highly advanced among
filamentous algal forms with a division of labour. Thallus is
multicellular, multiseriate and filamentous. The thallus consists of
a prostrate (horizontal) creeping system and an erect projecting
system (Fig.4.2.4). The prostrate system attaches to substratum
with rhizoidal filaments, it gives primary projecting system aerially.
Hence the thallus is called as heterotrichous.The upright system
may also contain secondary and tertiary projecting systems. This
type of differentiation in some filamentous forms led to the
evolution of terrestrial land plants. Heterotrichous habit is a
characteristic feature of order Chaetophorales in Chlorophyceae,
Phaeophyceae, Rhodophyceae and some members of Chrysophyceae
and Dinophyceae.

Fig.4.2.4 Heterotrichous thalli


(a) Coleochaete (b) Draparnaldiopsis
(c) Stigeoclonium
Source:https://www.uni-due.de/
biodiversitaet/lehre.php
http://www.outerhebridesalgae.uk/fwalgae/
fwa-species.php?id=441
https://commons.wikimedia.org/wiki/
Category:Algae#/media

Activity:
1. Collect macro and micro algae from fresh, estuarine, marine
habitats – observe the structural features of thalli with
naked eye and aid of microscope; draw diagrams on chart.
2. Debate evolutionary tendencies in thallus structure among
different classes of algae in a group discussion.

252 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Siphoneous habit: The thallus is unicellular filamentous and contain
many free nuclei (coenocytic). A large central siphon-like vacuole is
present in the thallus and thus named siphonous. Some members of
Chlorophyceae (Valonia, Bryopsis, Codium), and Xanthophyceae
(Botrydium, Vaucheria) possess this kind of thalli (Fig.4.2.5)

Fig.4.2.5 Siphoneous habit (a)


Botrydium (b) Bryopsis (c) Codium
Source: https://alchetron.com/Botrydium
https://www.inaturalist.org/taxa/
https://en.wikipedia.org/wiki/
File:Codium_sp.jpg

Pseudo-parenchymatous habit: These thalloid forms are originated


from filamentous algae by juxtaposing the branch system. This thallus is
of two types, uniaxial form and multiaxial form (Fig.4.2.6). In uniaxial
forms one main axis is present from which all other side branches are
formed. Batrachospermum and Dumontia. of Rhodophyceae show this
habit. On the other hand, in multiaxial forms, the thallus has many
threads nearby and look like multiaxial construction. (Nemalion,
Polysiphonia).

Fig.4.2.6Pseudo-parenchymatous habit (a) Batrachospermum (b) Nemalion


Source: https://www.uni-due.de/biodiversitaet/
https://www.centralcoastbiodiversity.org/rubber-threads-bull-nemalion-elminthoides.html

APSCHE : SEMESTER-I BOTANY TEXT BOOK 253


Fritsch (1935) opined that Chlorococcales members become
coenocyticpreceding reproduction to generate siphonaceous habit.

Parenchymatous habit: This type of thallus is formed due to the


septation of the filament in two or more than two planes. It appears either
simple foliose form (Ulva, Porphyra, Dictyota) or complex (Sargassum,
Fucus). In Laminaria, the thallus has an intricate form and divisible into
holdfast, stipe, and blades. The thallus in Macrocystis highly complex and
has sieve tube-like structures, a characteristic of higher plants (Fig.4.2.7).

Fig.4.2.7 Parenchymatous habit (a) Ulva


(b) Pophyra (c) Sargassum (d) Macrocystis
Source: https://commons.wikimedia.org/wiki/
File:Meersalat-Ulva-lactuca.jpg
https://upload.wikimedia.org/wikipedia/commons/
2/26/Porphyra_umbilicalis_Helgoland.JPG
https://commons.wikimedia.org/wiki/
File:Sargassum_-_1.jpg
https://commons.wikimedia.org/wiki/
File:CAS_Macrocystis_4.JPG

254 APSCHE : SEMESTER-I BOTANY TEXT BOOK


4.2.2. Life Cycles in Algae:

Introduction: Algae are primitive plants. Except in diatoms, some


brown and red algae, generally the main vegetative thallus in life
cycle is a gametophyte (n). The gametophyte is either dioecious or
monoecious. Male and female sex organs produce male and female
gametes. Fusion of gametes under the process of fertilization results
in a diploid zygote. Meiosis in zygote gives haploid spores, that
germinate into haploid gametophytes. Conversely, in some algae
zygote does not undergo meiosis immediately, develop into a diploid
sporophyte. The sporophyte produces haploid gametes asexually
through meiosis in spore mother cells (2n). Thus, alternation of
haploid gametophytic and diploid sporophytic phases are present
in the life cycles of algae. Asexual and sexual reproductions, apart
from vegetative reproduction are present in the life cycles of algae.
Sexual reproduction is completely absent in some classes of algae
(e.g., Myxophyceae); thus, alternation of generations is absent in
such forms. Based on the activity of zygote, site of fertilization and
meiosis, nature of gametophytic and sporophytic phases, life cycles
in algae are classified into 5 basic types.
A.Haplontic life cycle: This is the most primitive and simple life
cycle.Majority ofChlorophyceae members, few species of
Chrysophyceae, and Xanthophyceae show this type of life cycle.
Vegetative thallus is haploid gametophyte and either monoecious
or dioecious. It bears haploid sex organs and produces haploid
gametes. Fertilization of male and female gametes result in the
formation of a diploid zygote. The zygote undergoes meiosis,
immediately or after a resting period and form 4 meiospores
(asexual). In monoecious species (Oedogonium) 4 meiospores are
alike, germinate and develop into gametophytes. But in dioecious
species (some species of Chlamydomonas, Ulothrix), out of 4

APSCHE : SEMESTER-I BOTANY TEXT BOOK 255



 
       
        
   


   
 


     


Fig.4.2.8 Haplontic life cycle in Chlamydomonassp.


Source: https://sciencedoing.blogspot.com/2013/04/origin-and-evolution-of-sex-in-lower.html

256 APSCHE : SEMESTER-I BOTANY TEXT BOOK


meiospores, 2 develop into + strain and 2 into – strain
gametophytes. In this life cycle, haploid stages are dominant for
most of the time, zygote is the only diploid stage (Fig.4.2.8). Thus,
this life cycle is said to be haplontic.

B.Diplontic life cycle: This life cycle is evident in Bacillariophyceae,


Siphonales, Siphonocladales, and Dasycladiales of Chlorophyceae,
and Fucalesof Phaeophyceae . The vegetative thallus is a diploid
sporophyte. It produces diploid gametangia, in which gametes
formed after meiosis. The haploid gametes fuse to form a diploid
zygote, which re-establishes the sporophytic phase through mitotic
Sargassum
type. In this life cycle, diploid stages are evident most of the time,
and the haploid stage is restricted to gametes only (Fig.4.2.9). Thus,
this life cycle is considered diplontic.

C. Diplo-haplontic life cycle: In this type of life cycle,


Sporophyte(diploid) and gametophyte (haploid) plants alternate
with each other. Life cycle starts with diploid phase and alternate
with haloid phase. Due to the occurrence of diploid and haploid
phases. life cycle is diplo-haplontic. This type falls into two
categories, based on the morphological nature of gametophytes and
sporophytes.

i. Isomorphic diplo-haplontic life cycle: In this life cycle, both


the sporophyte and gametophyte are morphologically similar
(isomorphic) and develop alternately.Thus, the life cycle is known
as isomorphic (homologous or homomorphous) diplo-haplontic.
The sporophyte (2n) is an asexual thallus, produces haploid spores
by meiosis. Spore (n) germinates to produce haploid gametophyte.
Gametophyte is a sexual phase that produces gametes in gametangia
by mitosis. Fertilization of gametes result into the zygote (2n).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 257


Fig.4.2.9 Diplontic life cycle in Sargassumsp.
Source: https://wiki.nus.edu.sg/display/TAX/Sargassum+polycystum

  




 

       


 
  
 
       
  
   
     
258 APSCHE : SEMESTER-I BOTANY TEXT BOOK
Zygote germinates directly into a diploid sporophyte (Cladophora,
Ulva, and Ectocarpus) (Fig.4.2.10 a).
ii. Heteromorphic diplo-haplontic life cycle: The sporophyte
(2n) and gametophyte (n) are morphologically dissimilar
(=heteromorphic) and alternate in the life cycle. Hence it is named
heteromorphic (heterologous or heteromorphous) diplo-
haplontic life cycle. The sporophyte (2n) is macroscopic and grows
few to several meters in length. It bears sporangia and produces
haploid meiospores asexually. The spores germinate to form
microscopic gametophytes (sexual thalli). Sex organs developed on
gametophyte produce haploid gametes by mitosis. The diploid
zygote formed after fertilization produces the diploid sporophyte
(Laminaria)(Fig.4.2.10b Cutleria Urospora, game-
tophytes are much bigger than the sporophytes.

Fig.4.2.10 Diplo-haplontic life cycle (a) Isomorphic Diplo-haplontic in Ulva sp.


(b) Heteromorphic Diplo-haplontic inLaminaria sp.
Source: http://www.meghanrocktopus.com/bocas-arts-grant
https://www.plantscience4u.com/2014/05/diplohaplontic-life-cycle-in-algae.html

D. Haplobiontic life cycle: This type of life cycle occurs in many


Batrachospermum, Nemalion, etc. The life cycle
Batrachspermum
gametophyte produces gametangia, spermatangia (male) and

APSCHE : SEMESTER-I BOTANY TEXT BOOK 259





  



                               
          
   
 

Fig.4.2.11 Haplobiontic life cycle in Batrachospermum sp.


Source: http://comenius.susqu.edu/biol/205/day-10/laboratory_9-rhodophyta.htm

260 APSCHE : SEMESTER-I BOTANY TEXT BOOK


carpagonia (female). The spermatia (male gametes) and egg cells
(female gametes) are formed respectively from male and female sex
organs. The zygote (2n) formed after fertilization divides meiotically
and produces 4 meiospores (n) in the venter of carpagonim.
Meiospores germinate in the basal part of the carpogonium and
produce haploid gonimoblast filaments. Apical cells of those
filaments function as carposporangia producing haploid
carpospores. The gonimoblast filaments enclosed by many sterile
filaments form a carposporophyte (n). It is a parasite on the parent
gametophytic thallus. The carpospores germinate to produce a
heterotrichouschantransia stage. Erect filaments formed on the
chantrantia stage develop into new gametophytes. In this life cycle,
gametophyte and carposporophyte represent two haploid phases.
Thus, it is known as a haplobiontic or diphasic life cycle
(Fig.4.2.11
E. Haplo-diplobiontic life cycle: This life cycle is common in red
Gracilaria,Polysiphonia
male (spermatangia) and female (carpagonia) sex organs develop
on different thalli. The zygote (2n) formed in the venter of
carpagonium divides mitotically. Diploid gonimoblast filaments are
formed from the base of a placental cell, their uppermost cells behave
as carposporangia. Post-fertilization changes lead to the
development of a carposporphyte (cystocarp), which is a parasite
on the female gametophyte. Carpospore (2n) formed from cystocarp
germinate to produce diploid tetrasporophyte. Reduction division
in a tetrasporangium gives 4 haploid tetraspores. Out of the 4
tetraspores, 2 develop into male and other 2 into female
gametophytes. In such a life cycle, gametophytes represent the
haploid phase, while carposporophyte and tetrasporophyte
represent the 2 diploid phases. Thus, the life cycle is called haplo-
diplobiontic and triphasic (Fig.4.2.12).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 261


Fig.4.2.12. Haplo-Diplobiontic life cycle in Chondrus crispus
Source:https://commons.wikimedia.org/wiki/Category:Alga_life_cycle#/
media/File:Life_cycle_Chondrus.jpg

Mother of the Sea: Kathleen Mary Drew-Baker was a British phycologist


deciphered the life cycle of the red alga, Porphyra umbilicalis. She identified
Porphyra, previouslyconsidered as

journal; it revolutionized the commercial cultivation technology of nori in


Japan, Korea and China. In recognition of her research finding, Japanese
constructed a monument looking towards the Porphyra cultivation site in
Kumamato Prefecture. On the monument she was cited as mother of the
sea. Every year Japanese celebrate ‘Drew Festival’ nationwide on 14th April
in her honour.

https://en.wikipedia.org/wiki/Kathleen_Mary_Drew-Baker
https://mmwonderwomen.files.wordpress.com/2018/03/utoshi_17.jpg
http://www.fao.org/fishery/culturedspecies/Porphyra_spp/en

262 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Chapter -4.3 Spirogyra and Polysiphonia

Learning outcomes: At the end of the chapter you should be able


to:
1. Sketch the thallus structure, reproduction and life cycles
in Spirogyra and Polysiphonia.
2. Compare, contrast and infer the general qualities of green
and red algae.

Green Algae: Algae belong to class Chlorophyceae are of bright


grass green colour resembling embryophytes. About 14,000
species are being reported in this algal class. Of which, 90% are
fresh water and 10% are marine species. Species belong to
Oedogoniales and Zygnematales orders are exclusively freshwater.
Some members live predominantly in freshwater (Ulotrichales,
Coleochaetales) or marine (Caulerpales, Dasycladales,
Siphonocladales) habitats.

Red Algae: Algae belong to class Rhodophyceae are habitually


red in colour owing to the high amount of phycobilin, r-
phycoerythrin. About 7,000 species of red algae are recognized
so far, of which 95% of species occur in marine habitat. Other 5%
are found in fresh water bodies, mainly from warmer areas of the
globe. Porphyridium, a unicellular red algae dwells on the substrata
in freshwater brackish and marine environments and also moist
soils. Some red algae live as mangrove associates in tropical and
sub-tropical regions of the globe. Red seaweeds inhabit greater
depths (about 200 meters) than other macroalgae due to
chromatic adaptation. Unicellular red algae belong to
Cyanidiophytina are extremophiles living in sulphuric hot springs
and acidic habitats.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 263



    




       
       


         
 
     

        



     
    
   
       


  


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4.3.1 Spirogyra:
Kingdom : Plantae
Divison : Thallophyta
Class : Chlrophyceae
Order : Conjugales
Family : Zygnemataceae

Occurrence: Spirogyra is a filamentous green algal genus consisting


of 400 species. It is cosmopolitan and occurs in stagnant freshwater
bodies like ponds and pools, a few species live in lakes. S. crassata,
S. elongata, S. microspora, S indica and S. communis are common
species in India. The word Spirogyra derives from two Greek words
i.e., Speria (= coil) and gyras (= twisted). Spirogyra appears as a bright
green silky blanket in surface water of ponds (Fig.4.3.1). Hence, it
is called as pond silk, pond scum, or mermaid’s tresses. Most of
the species are free floating, some are (S.jogensis, S. adnata) attached
to substrata (adherent) with a specialized basal cell.

Fig. 4.3.1 Fig. 4.3.2


Spirogyra on surface water of a pond Thallus of Spirogyra
Source: https://eol.org/pages/11681/ Source: https://eol.org/pages/11681/
media?license_group=cc_by_sa&page=3 media?license_group=cc_by_sa

Thallus structure: Spirogyra thallus is a multicellular, uniseriate


and un-branched filament with a dome shaped terminal cell
(Fig.4.3.2). Often, the filaments appear together in a matted
blanket due to their slimy mucilage sheaths. The cells of the filament
are elongated and cylindrical present one above the

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      
   



 
        
 

 
 
     

    
        
      
 
    
  




      
  
  

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other(uniseriate). The lower cell in attached species (S.fluviatilis) is
called as holdfast or hapteron, which is colourless (non-
chlorophyllous) with irregular lobes. Except holdfast, all the cells
in a filament undergo divisions; they elongate to increase the length
of the thallus.
Cell structure: Each cylindrical cell in Spirogyra filament consists
of two parts namely cell wall and protoplast (Fig.4.3.3). The cells
in a filament are joined with one another by middle lamellae. The
cell wall has inner cellulose and outer pectose layers. The outer
pectose layer softens in water to form a slimy mucilage sheath

around the protoplast. The protoplast comprises of an outer plasma


membrane and central large vacuole surrounded by a thin layer of
cytoplasm. The cytoplasm is called primordial utricle (= peripheral
cytoplasm), because it is present around the large central vacuole.
Each cell contains 1-16 spirally coiled ribbon shaped chloroplasts
surrounding the vacuole; hence this alga owes its generic epithet,
Spirogyra. The margins of chloroplast are either smooth or serrated.
In each chloroplast, many pyrenoids are present in a row at regular
intervals. Each pyrenoid has a central protein core encircled by starch
sheath. A conspicuous nucleus with distinct nucleolus resides in
the centre of each cell, by means of strands from all sides of the
peripheral cytoplasm. Other cell organelles like mitochondria,
ribosomes, Golgi bodies, ER etc., remain scattered in the cytoplasm.

Activity: Collect the fresh sample of the Spirogyra from a fresh water
pond, fix it in FAA (Formaldehyde: Alcohol:Acetic acid in 10:50:05
percent ratio + 35% distilled water), make temporary slides, observe
under microscope to study structure of the thallus, cell and
reproductive stages. Take photographs or draw the diagrams.
Otherwise observe the said things using permanent slides available
in your Botany laboratory

APSCHE : SEMESTER-I BOTANY TEXT BOOK 267


(B)

(C)

Fig. 4.3.3 (a) Cell structure of Spirogyra


(b) Nucleus(c) Pyrenoids

(A)

 
   
  
       
 

 
    
 
    


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Reproduction: Spirogyra shows three types of reproduction namely
vegetative, asexual and sexual reproduction (Fig.4.3.4).

A. Vegetative reproduction: It is the most common type of


reproduction under favourable conditions. It occurs by means of
fragmentation. Mechanical injury or dissolution of the middle
lamella can lead to fragmentation. Occasionally the middle lamella
of one cell protrudes into an adjacent cell and causes breakage of
the filament. In either of the cases, cells in each fragment undergo
multiple divisions and elongation to form a new filament.

B. Asexual reproduction: It occurs rarely under unfavourable


conditions through production of asexual spores (akinetes and
aplanospores) or by azygospores.

i. Akinetes: They are thick-walled resting spores. In S. farlowii,


during harsh environmental conditions some cells of the filament
lose water and deposit thick cell walls around to form akinetes.
Akinetes germinate and produce new filaments under favourable
conditions.

ii. Aplanospores: They are thin-walled non-motile spores. In S.


aplanospora, the protoplast of each vegetative cell shrinks and a thin
wall develops around it, thus transforms into an aplanospore. On
return of favourable conditions, a new filament is formed by
germination of each aplanospore.

iii. Azygospores (parthenospores): In some Spirogyra species


(S.rhizoides, S.mirabilis) due to physiological conditions gametes fail
to fuse and act like asexual spores. Those haploid non-motile
gametes germinate to produce gametophytic filaments.

C. Sexual reproduction: In Spirogyra, sexual reproduction occurs


at the end of growing season, when the temperature elevates in the

APSCHE : SEMESTER-I BOTANY TEXT BOOK 269


Fig. 4.3.4 Asexual reproductive methods in Spirogyra

    



        

      


   


Know more: In Spirogyra, after completion of physiological


anisogamy all cells (male gametangia) in a conjugated filament
remain empty, while all cells (female gametangia) in the other
filament contain zygospores. In the case of isogamous scalariform
conjugation, cells in both the conjugated filaments remain empty
and zygospores are seen in conjugation tubes.

270 APSCHE : SEMESTER-I BOTANY TEXT BOOK


surrounding water. Spirogyra shows sexual reproduction by means
of Conjugation. Based on the nature and number of filaments that
participate in sexual reproduction conjugation in Spirogyra is of two
types; (i) Scalariform conjugation and (ii) Lateral conjugation.

i) Scalariform conjugation: It occurs in dioecious (unisexual or


(S.majuscula, S.ternata) and hence
it is also called as dioecious conjugation (Fig.4.3.5). Two filaments
of opposite strains (-male and +female) are attracted towards each
other and lie parallel under a common mucilage sheath. Opposite
cells of adjacent filaments act as gametangia and produce finger
like papillae towards each other. Elongated papillae fuse with one
another to form a horizontal tube called conjugation tube by the
action of enzyme cytase. As a result, the two filaments appear like a
ladder with many horizontal conjugation tubes under a
mucilaginous sheath. Hence it is called scalariform (Greek words,
scala = ladder; forma = shape) conjugation. When conjugation tubes
are forming, the protoplasts in conjugating cells lose water,
accumulate starch and transform as gametes called aplanogametes.
Aplanogametes of male (-) and female (+) strains are
morphologically similar and non-motile; but physiologically
different. The active male gamete (-) enters into female
gametangium by amoeboid movement, through conjugation tube.
Male gamete fuses with female gamete forming a diploid zygote.
Thus, the conjugation is said to be physiological anisogamy. In
some species of Spirogyra, both the aplanogametes from filaments
of + and – strains are active and alike (isogametes). Both the gametes
move opposite to each other and are fused in the conjugation tube
resulting in the formation of Zygote (2n). Thus, the conjugation is
said to be isogamy.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 271


Fig. 4.3.5 Stages of scalariform conjugation in Spirogyra
Source:https://eol.org/pages/11681/media?license_group=cc_by&page=2
https://eol.org/pages/11681/media?license_group=cc_by&page=3

ii.Lateral conjugation: It happens in monoecious (bisexual or


homothallic) species and so called as monoecious conjugation. In
this conjugation two adjacent cells of the same filament act as +
and – gametangia and produce aplanogametes. It is of two types:
(a) Indirect and (b) Direct lateral conjugation.

a. Indirect lateral conjugation: In this conjugation, two adjunct


cells in a thallus filament behave as male and female gametangia;
they produce papillae on both sides of the septum present between
+ and - cells. The two papillae unite and form a lateral conjugation
tube. The protoplasts of two conjugating cells transform into
gametes. Fertilization occurs as the gamete from male gametangium
moves and fuses with gamete in female gametangium. After the
completion of conjugation, female gametangium contain zygospore
while male gametangium remains empty (Fig.4.3.6). e.g. S.affinis,
S.tenuissim

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b. Direct lateral conjugation: This type of conjugation was
reported in attached species of Spirogyra (Spirogyra mirabilis,
Spirogyra jogensis). Two cells above the holdfast act as gametangia,
upper cell becomes male gametangium while cell next to holdfast
enlarges in size to form a barrel shaped female gametangium
(Fig.4.3.7). Conjugation tube is not formed. The male gamete
enters into female gametangium by making an aperture in the
septum with its beak like structure. A zygote is formed by the fusion
of male and female gametes.

Germination of zygospore: After completion of fertilization, the


zygote containing filaments settle down at the bottom of pond or

zygospore. Zygospores are liberated on decay of female gametangia


and remain dormant at the bottom of pond till the favourable
conditions return. The thick wall of zygospore wall is differentiated
in to 3-layers i.e., outer exosporium (cellulose), middle mesosporium
(cellulose and chitin) and inner endosporium (cellulose). The diploid
nucleus of zygospore undergoes meiosis during favourable
conditions to produce 4 haploid nuclei, out of which 3 degenerates.
The zygospore with one haploid nucleus gradually enlarges and burst
open to release a germ tube. A new filament is formed by repeated
transverse divisions in the germ tube (Fig.4.3.8).

Life cycle: Spirogyra is a green alga with multicellular and


unbranched filamentous thallus. The vegetative thallus that appears
for most of the time in its life cycle is haploid and gametophytic in
nature. Gametophytic plants reproduces by vegetative
(fragmentation) and asexual (akinetes, aplanospores, or
azygospores) methods. The sexual reproduction (conjugation)
results in the formation of a diploid phase, the zygospore (a

APSCHE : SEMESTER-I BOTANY TEXT BOOK 273


Fig. 4.3.6 Stages of indirect lateral conjugation in Spirogyra

Fig. 4.3.7.Stages of direct lateral conjugation in Spirogyra

Fig. 4.3.8

Zygospore germination stages


in Spirogyra

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temporary sporophytic phase). Zygospore formed from sexual
reproduction is the only diploid phase in the life cycle of Spirogyra.
Hence the life cycle of Spirogyra is haplontic (Fig.4.3.9).

Fig. 4.3.9 . A schematic diagram showing life cycle of Spirogyra

Watermelon snow: Chlamydomonas nivalis is a cryophilic (cold


loving) green alga and lives in freezing water. This alga imparts
red colour to snow in alpine and coastal polar regions worldwide
during summer months. This phenomenon is variously called
as red snow or pink snow or blood snow. This is due to the
accumulation of astaxanthin (red coloured xanthophyll) in
hypnospores of that alga. Walking on such snow often results
in attainment of bright red soles and pink trouser cuffs.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 275


Red or Blood rain phenomenon in Kerala, Tamil Nadu and
Sri Lanka:

The green algae belong to Trentepohliales order are aerial to sub-


aerial, need free water only to germinate. Their colour varies from
yellow-green to bright orange, depending on the amount of
carotenoid pigment produced. Species of Trentepohlia live on tree
trunks, wet rock or as symbiont in lichens. Air borne spores of
this alga mixed with rain water cause a red rain. This was reported
from some laces in Kerala, Tamil Nadu, and in eastern and north-
central provinces of Sri Lanka during June, November and
December months of 2012. This phenomenon was also reported
from many countries in the world where Trentepohliais habituated.

Green algae as experimental material to elucidate


photosynthesis:
 T.W.Englemann (1881-‘82) conducted experiments with
Spirogyra sp., Cladophora sp. and Proteus sp. (aerotactic
bacteria). He hypothesized that blue and red wavelengths
are effective for photosynthesis.
 O.Warburg (1923) estimated the quantum of light required
to fix a molecule of CO2and to release a molecule of O2using
Chlorella. He also demonstrated the inhibitory effect of
elevated O2 concentrations on photosynthesis using the same
alga.
 R. Emerson (1943 &1960) experiments on Chlorella,
revealed the two photochemical reactions during light phase
of photosynthesis.
 Melvin Calvin (1961) explained the mechanism of carbon
fixation, C3 cycle in photosynthesis through his classical
experiments on a unicellular green alga, Chlorella.
 In the present century, Chlamydomonas reinhardtii is being
used as an experimental material to work out molecular
biology of photosynthesis.

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Green algae with varied nutritional habits: Green algae are
photosynthetic and autotrophic. However, species of Cephaleuros,
Rhodochytrium and Phyllosiphon are parasitic green algae. The genus
Cephaleuros comprising 14 species, they live as host non-specific
parasites; infect about 400 species of seed plants from the tropics
and sub-tropics. Cephaleuros is sub-cuticular with fungus like
filaments and sterile hairs; it produces sporangiophores and
zoosporangia that resemble downy mildew fungi. The red patches
due to this algal infection on host leaves are referred as algal rust or
‘red rust’. In 19th century, Cephaleuro sparasiticus caused severe
damage to tea and coffee plantations in India. Trebouxia is the most
common photobiont (previously known as phycobiont) in lichens.
Some green algae are epizoic. For example, species of Cladophora
and Stigeoclonium are found respectively on snails and gills of fishes.
Species of Chlorella, Carteria and Tetraselmis live within the bodies
(endozoic zoochlorellae) of protozoans and invertebrates; the
relationship is said to be commensalism rather than symbiosis.

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4.3.2. Polysiphonia
Kingdom : Plantae
Divison : Thallophyta
Class : Rhodophyceae
Order : Ceramiales
Family : Rhodomeliaceae

Occurrence: Polysiphonia is a marine red algal genus consisting of


150 species. This alga is cosmopolitan in distribution, grows in
littoral and sublittoral zones. In India, 16 species of this genus are
reported from western and southern coasts, of which P. platycarpa
is the most common species. Species of this alga may grow as
lithophytes i.e., on rocks or stones (e.g., P. elongata), or epiphytes
(e.g., P. urceolata, P. terulacea) on Laminaria spp. or semi-parasites
(e.g., P. fastigiata) on Ascophyllum nodosum. The generic name
Polysiphonia is due to the thallus with many tubular filaments,
derived from the Greek words’ polloi (=many) and siphon (=pipe).

Polysiphonia: The thallus is filamentous, red or purple


red in colour. It has many inter-connected siphonous filaments.
The cells in the filaments are joined with one another by pit
connections forming tubular structures; thus, this alga is named as
Polysiphonia.

Fig. 4.3.10 Thallus of Polysiphonia


Source:https://www.researchgate.net/figure/Vegetative-structures-of-Polysiphonia-
ulleungensis-sp-nov-A-Holotype-specimen_fig1_272872707

278 APSCHE : SEMESTER-I BOTANY TEXT BOOK


The thallus is profusely branched and feathery in appearance (Fig.
4.3.10). It may reach the length of about 30 cm. Most of the species
are of heterotrichous habit with prostrate and erect systems.

a. Prostrate System: It is generally multiaxial and well developed.


Many unicellular rhizoids arise from its lower side (Fig.4.3.11).
The rhizoids in P. urceolata and P.nigrescens are lobed at their apices

P.elongata and P.violacea. In such species, rhizoids develop from the


basal cells of the erect system; they form massive attachment disc
by aggregation.

Fig. 4.3.11 Prostrate system in thallus of Polysiphoniashowing rhizoids and discs.


Source:https://www.researchgate.net/figure/Vegetative-structures-of-Polysiphonia-
ulleungensis-sp-nov-A-Holotype-specimen_fig1_272872707

b. Erect System: The erect filaments that develop from the


prostrate system constitutes a main axis and many branches
(Fig.4.3.12). The branches are of two types i.e., long branches and
short branches. The long branches are of unlimited growth, while
the short branches are of limited growth. The long branches develop
in a spiral or radial manner, whereas short branches borne in spiral
manner. The short branches develop on both the main axis and long
branches; they are also called as trichoblasts. Trichoblasts are
multicellular, uniseriate, dichotomously branched and colourless.
They are deciduous structures that produce sex organs.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 279


  



     

 
  
   
    
    

          
        
        



Fig. 4.3.12 Erect system in thallus of Polysiphonia showing branching pattern


Source:https://www.researchgate.net/figure/Vegetative-structures-of-Polysiphonia-
ulleungensis-sp-nov-A-Holotype-specimen_fig1_272872707

280 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Both the main axis and long branches contain a central siphon. It is
composed of many elongated cylindrical cells arranged in a vertical
row. It is surrounded by 4-20 peripheral siphons (Fig.4.3.13). Each
peripheral siphon is a multicellular and uniseriate structure. Hence,
the thallus appears as a polysiphonous structure. In most of the
species, cells of the pericentral siphons may divide by periclinal and
anticlinal divisions to give three layers of cortical cells. The main
axis and long branches at their apices contain central siphon only,
and so appear as uniseriate. Thallus grows by a dome shaped apical
cell located at the tip of the central siphon.

Polysiphonia: The cells have thick walls,


differentiated into outer pectic and inner cellulosic layers. All cells
in the thallus are connected with each other by pit connections
(cytoplasmic connections). The cytoplasm is present peripherally
around a large central vacuole. Each cell is uninucleate with many
discoid chromatophores devoid of pyrenoids. The cytoplasm
contains granules of floridean starch as food reserve. Chlorophyll-
a, Chl.-d, r-phycocyanin and r-phycoerythrin are the pigments
present in chromatophores.
Cyanobacterial lineage of red algae: Red algae are the closest
eukaryotic algae to Cyanobacteria (BGA). Both the groups share
common characteristics such as, lack of chloroplast ER, simple
plastids with unstacked thylakoids, phycobiliproteins and absence
of flagella.Based on these similarities, cyanophyte origin of
Rhodophyceae is presumed via Cyanidium like unicellular forms.

Reproduction: Poysiphonia has three morphologically similar


plants. They are male and female gametophytic plants (n) and
tetrasporophytic (2n) thalli. Reproduction in this alga occurs by

APSCHE : SEMESTER-I BOTANY TEXT BOOK 281


Fig. 4.3.13 Thallus of Polysiphonia showing cellular arrangement.

  


 
 



 

    
     


         

   
      

282 APSCHE : SEMESTER-I BOTANY TEXT BOOK


vegetative, asexual and sexual methods. Vegetative reproduction
happens due to fragmentation of the thallus.

A . Asexual Reproduction: Tetrasporophyte (2n) is an


independent, polysiphonous, and asexual thallus; it produces
haploid non-motile tetraspores (Fig.4.3.14 a & b). A pericentral
cell in each tier at the apex of the thallus functions as tetrasporangial
initial.This initial cell is smaller than other pericentral cells of any
particular tier. A vertical division in this initial cell forms an inner
and an outer cell. The inner cell directly acts as sporangial mother
cell; while the outer cell divides further to form two or more cover
cells.A lower stalk cell and an upper tetrasporangial cell are
formed by the transverse division in the sporangial mother cell.
The tetrasporangial cell enlarges in size and develops into a
tetrasporangium. Cell of the tetrasporangium gives 4

the sporangium ruptures to liberate tetraspores into the


surrounding water. Those spores develop into polysiphonous
Polysiphonia is heterothallic,
thus two male and two female thalli develop from the four
tetraspores produced by a tetrasporangium.

B. Sexual Reproduction: In Polysiphonia, sexual reproduction is


oogamous type. Gametophytes are dioecious in nature. Hence, male
sex organs (spermatangia) and female (carpogonia) sex organs
respectively develop on male and female plants.

a. Spermatangia: The male sex organs, spermatangia (antheridia)


are produced at the apex of a fertile trichoblast. In the
dichotomously branched fertile trichoblast, one arm develops into
a fertile axis (Fig.4.3.15a). The other sterile arm of the fertile
trichoblast may divide further in dichotomous manner. Except the

APSCHE : SEMESTER-I BOTANY TEXT BOOK 283


Fig. 4.3.14Polysiphonia tetrasporophyte showing (a) Formation of tetrasporangia

(b) Arrangement of tetraspores.



  
 

   

     


   

 

284 APSCHE : SEMESTER-I BOTANY TEXT BOOK


two basal cells, all the cells of fertile axis divide and form pericentral
cells on both the sides. The pericentral cells undergo several
divisions to cut off spermatangial mother cells. Each of which
divide further to produce 2-4 unicellular spermatangia
(Fig.4.3.15b). The cytoplast of each spermatangium transforms
into a non-motile male gamete, the spermatium. The spermatia
enter into water after the rupturing of the wall of a spermatangium.

b. Carpogonia: The fertile trichoblast on female plant is much


shorter as compared to the male one. A 5-7 celled long trichoblast
develops from an initial cell, which is 3-4 cells below the apical cell
of the central siphon. Three tiers of pericentral cells are formed by
vertical divisions in the lower most three cells of this trichoblast. A
pericentral cell in the middle tier present towards the mother axis
develops into a supporting cell.This supporting cell cuts off 4-celled
branch called procarp (carpogonial filament). The terminal cell of
this filament functions as carpogonium (female sex organ). The
carpogonium has a swollen base with a uninucleate egg and an
elongated trichogyne at its top (Fig.4.3.16). Later on, the
supporting cell cuts off two sterile initials, one at its base and other
on its lateral side. The basal sterile initial remains undivided. But
the lateral sterile initial immediately undergoes transverse division

ready for fertilisation.

c. Fertilization: The spermatia in water attach to the tip of a


receptive trichogyne.The walls of the successful spermatium and
trichogyne dissolve at their point of contact. The male nucleus
enters the base of carpogonium through trichogyne.The fusion of
male and female nuclei results in the formation of a diploid nucleus.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 285


Fig. 4.3.15 Polysiphonia male gametophyte showing
spermatangia and formation of spermatia
Source:https://upload.wikimedia.org/wikipedia/commons/3/30/
Polysiphonia_spermatangia_WM.jpg
      

      

  
        
       

     
    

Fig. 4.3.16Polysiphonia female gametophyte showing development of carpogonium

286 APSCHE : SEMESTER-I BOTANY TEXT BOOK


d. Post-Fertilization Changes: After the process of fertilization,
the supporting cell located just below the basal region of the
carpogonium cuts off an auxiliary cell. At the same time, the lateral
sterile filament increases in length to become 5-10 celled structure.
The basal sterile initial divides at this stage to form a two celled
filament. Simultaneously, a tubular connection (ooblast) is
established between the auxiliary cell and carpogonium. The diploid
nucleus in carpogonium divides mitotically into two nuclei, of which
one is transported to the auxiliary cell via ooblast and the other
one remains in the carpogonium. So that the auxiliary cell contains
its own nucleus (n) and a migrated nucleus (2n). The haploid nucleus
(n) in the auxiliary cell and trichogyne of carpogonium
degenerates.Later, carpogonial filament gradually shrivels, begin
to degenerate and finally disappears (Fig.4.3.17).

Fig. 4.3.17 Post-fertilization changes in Polysiphonia

The pericentral cells adjacent to the supporting cell begin


to form a sterile sheath. At first one gonimoblast initial followed by
many such initials are produced at the upper side of the auxiliary

APSCHE : SEMESTER-I BOTANY TEXT BOOK 287



          

  
      
      


        
 

       
   
 
 


n

    

     
 

 


288 APSCHE : SEMESTER-I BOTANY TEXT BOOK


cell. The diploid nucleus (fusion nucleus) in the auxiliary cell
undergoes mitotic divisions to form many diploid nuclei. These
nuclei enter into gonimoblast initials followed by formation of
partition walls.Each gonimoblast initial by repeated mitotic
divisions forms a gonimoblast filament. The terminal cell of the

the auxiliary cell, supporting cell, carpogonium and sterile filaments


form a large irregular shaped placental cell. This placental cell is of
nutritive function. Meanwhile, the pericentral cells, adjacent to the
supporting cell, divide and redivide to form a flask shaped protective
covering, the pericarp. The resultant structure, formed after all
these changes, is called as a cystocarp.It consists of the placental
cell, gonimoblast filaments and carpogonia covered by a sterile
pericarp.The cystocarp is parasitic on female gametophyte. It is an
urn-shaped structure with a small apical opening, the ostiole
(Fig.4.3.18). The diploid part of the cystocarp is carposporophyte.
Each carposporangium in this carposporophyte produces a single
carpospore (2n).Carpospores are liberated in to the surrounding
water through the ostiole of cystocarp.

e. Germination of Carpospore: The carpospores attach to any


solid surface in the marine waters, germinate to produce diploid
tetrasporophytic plants. The asexual spores, tetraspores formed on
this plant complete the cycle.

Activity: Collect the fresh sample of the Polysiphonia from nearest


marine waters, fix it in FAA (Formaldehyde: Alcohol:Acetic acid in
10:50:05 percent ratio + 35% distilled water), make temporary
slides, observe under microscope to study structure of the thallus,
cell and reproductive stages. Take photographs or draw the
diagrams. Otherwise observe the said things using permanent slides
available in your Botany laboratory.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 289


Fig. 4.3.18 Cystocarp
inPolysiphoniarepresenting
carposporophytic stagewith
carpospores.
Source: https://upload.wikimedia.org/
wikipedia/commons/7/70/
Polysiphonia_cystocarp_WM.jpg

Life Cycle: Life cycle of Polysiphonia consists of three distinct phases


i.e., diploid tetrasporophyte, haploid gametophytes and diploid
carposporophyte. Diploid tetrasporophytic plant produces
tetrasporangia. Four haploid tetraspores are produced in a
tetrasporangium after meiosis. Of which two tetraspores develop
into male gametophytes and the other two into female
gametophytes. Male gametes (spermatia) develop inside the
spermatangia on male gametophytic plants. Similarly, female
gametes develop inside the carpogonigia on female gametophytic
plants. Fusion of a spermatium with egg cell in carpogonium results
into a diploid zygote.

The zygote develops inside the auxillary cell. An urn shaped


cystocarp containing gonimoblast filaments and the carposporangia
representing carposporophytic stage develops on female
gametophyte. This parasitic cystocarp gives out diploid carpospores.
Carpospores produce the diploid tetrasporophytic plants on
germination. Thus, the life cycle of Polysiphonia is triphasic and
haplo-diplobiontic type (Fig.4.3.19).

290 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig. 4.3.19A schematic diagram showing life cycle of Polysiphonia

Red algal parasites:Parasitism is quite common among red algae.


Parasitic red algae are Adelphoparasite (parasitic on a closely related red
alga), Alloparasite (parasite on other red alga of different family, tribe or
order). Some genera are Promiscuous alloparasite (parasitic on many
hosts, in which at least one host is not closely related to the
parasite).Choreocolaxpolysiphoniae is a complete parasite on Polysiphonia
fastigiata, which is an epiphyte on a brown alga, Ascophyllum nodosum.
Species of parasitic red algae, Harveyella and Holmsella completely lack
pigments.
* *
Ocean acidification a threat to life of coralline algae: All red algae
of Corallinales and few members of Nemaliales orders respectively deposit
CaCO3, in the form of calcite and aragonite. They are the keystone species
supporting the life of other algae, coral animals, other invertebrates and
vertebrates in the marine habitat in many ways. Elevated levels of Carbon
oxides and other greenhouse gases resulting in warming of seawater and
ocean acidification. According to many research findings, the skeleton of
coralline algae is highly susceptible to dissolution under ocean
acidification conditions. Deterioration of the crest forming coralline algae
may disturb rather collapse the entire coral reef ecosystem.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 291


Chapter- 4.4 Economic Importance of Algae

Learning outcomes: At the end of the chapter you should be able


to:
1. Explain the economic products of algae and their utility
by human beings for various purposes.
2. Discuss the ill-effects due to some algae.
3. Appraise the worth of phycocolloids in industries and
scientific fields.

Introduction: Human beings utilizing algae for various purposes


from time immemorial. Historical evidences indicate that marine
algae have been eaten in China, Japan and the Republic of Korea for
several centuries. Algae are source of human food, animal feed,
biofertilizers, pharmaceuticals and nutraceuticals, bioenergy, and
many other valuable products. However, some algal species cause
detrimental effects to aquatic animals, higher plants and human-
beings. This chapter provides information on the beneficial and also
harmful impacts of algae.
I. Beneficial impacts of algae:
A. Algae as human food: People from different parts of the world,
in particular maritime countries like China, Japan, Korea, Taiwan,
Indonesia, Philippines, Malaysia etc., have been eating fresh
seaweeds as salad components or in the form of recipes. There is a
constant search for new food sources to feed the increasing human
population. Nutritionally rich algae appear to be one of the
promising sources to meet the ever-increasing demand for food. Now
a days marine algae are regarded as sea vegetables and consumed in
many countries. Seaweeds belong to 160 genera are being used as
traditional human diet. Red algae like Porphyra (nori, laver),
Asparagopsis (limu), Chondrus crispus (Irish moss) and Palmaria

292 APSCHE : SEMESTER-I BOTANY TEXT BOOK


palmata (dulse); brown algae like Laminaria (kombu), Alaria, and
Undaria (wakame); and green algae like Monostroma (Aonori),
Caulerpa, Codium, and Ulva (sea lettuce) are the most common
among edible seaweeds (Borowitzka,2008). Many of the edible algae
are quite rich in proteins, vitamins and minerals including iodine.
Microalgae such as Spirulina maxima, Chlorella vulgaris, Dunaliella
salina and Haematococcus pluvialis are commercialized as a food
source or nutritional supplements.
Porphyra, Spirulina and Chlorella are rich in protein (35-65%),
may address the hidden hunger and combat protein deficiency in
people from developing and under developed countries of the globe.
Brown algae and some members of red algae are rich in Iodine,
useful to control thyroid disorders. In many countries, the above
mentioned macro and micro algae are cultivated in an industrial
and commercial scale to harness the algal food products.
B. Algae as animal feed: The exploitation of seaweeds as fodder
for the livestock by Greeks dates back to the 1st century B.C. During
World War-I, there was a shortage of fodder supply; thus, seaweeds
were fed to livestock (cattle, pigs, horses, sheep, poultry) in
Ascophyllum
Rhodymenia palmata
in Britain, and horseweed or sheep’s weed in Norway. Dried and
processed seaweeds have been used as animal feed in Europe and
Sargassum, Fucus, Ascophyllum,
Laminaria
Denmark, Scotland, America, China, and New Zealand, the livestock
is nurtured with seaweed fodder. In many countries, factories are
being established to prepare suitable cattle feed from seaweeds.
Hens fed on seaweed meal found to have an increased iodine
content. Cattle provided with a commercial sea-weed meal from

APSCHE : SEMESTER-I BOTANY TEXT BOOK 293



      

      
          
  

      

  

   
 
   


  
    
  
  


      

     
              

294 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Maxicarp, shown to have increased milk production and butter-fat
content. Acadian, a kelp meal marketed by Morgrove Holsteins
Limited proved to boost the immune system in cattle and other
livestock.
In aquaculture, plankatonic or attached, fresh or marine
species of algae can be given as feed depending on the type of
ecosystem. Microalgae belong to Chlorophyceae (Chlorella,
Tetraselmis, Scenedesmus), Myxophyceae (Synechococcus, Anabaena,
Aphanizomenon, Planktothrix), Bacillariophyceae (Navicula,
Nitzschia, Melosira, Amphora) are used as feed for fishes and shrimps
in aquaculture ponds. Some filamentous (Rhizoclonium riparium and
Polysiphonia mollis) and macro algae (Enteromorpha intestinalis, Ulva
lactuca and Catenella repens) are also offered as feed to animals in
aquaculture. It is an established fact that vitamins and other
nutraceutical elements in captive or cultured fishes can be traced
back to the algae on which they feed. Further, in aquaculture ponds
algae absorb the Carbon dioxide and releases Oxygen during
photosynthesis; thus, maintain water quality in shrimp and fish
ponds. It is observed that rainbow trout, Oncorhynchus mykiss fed
by Haematococcus pluvialis contained higher concentration of
astaxanthin, a better flesh colour; gain high value. In large yellow
croaker, Pseudosciaena crocea immunity increased with the intake
of astaxanthin, a microalgal pigment. The colour of ornamental
fishes often attributed to the pigments of algae consumed by them.
C. Algae as biofertilizers: The green revolution gears up
agriculture sector to meet the food requirements of increasing
population on the globe. Organic and nutrient rich material of
biological origin, biofertilizers are being used for restoration of soil
health and to reduce pollution. Blue Green Algae (BGA) are capable
of fixing the molecular nitrogen and increase the soil fertility. BGA

APSCHE : SEMESTER-I BOTANY TEXT BOOK 295



     
     
 


  

     
  

  
     
  


     
      
     
        


 
          
     
 

296 APSCHE : SEMESTER-I BOTANY TEXT BOOK


like Anabaena, Nostoc, Tolypothrix, Aulosira etc., are used as algal
fertilizers; produced in large scale and supplied to farmers in
powdered form. Azolla containing Anabaena azollae is cultivated in
rice fields to achieve good produce. Those fertilizers enrich the soil
with nitrogen, helpful for reclamation of alkaline soils and barren
soils, apart from improving the water holding capacity of the soil.
Seaweed compost is rich in minerals like potassium. phosphate,
sulphate and trace elements.Seaweeds are used as fertilizers for
coffee plantations in Sri Lanka, for vegetable crops in France and
Ireland. For cultivation of groundnut and sweet potato in China,
for cultivation of rice in Japan seaweeds are added to the soil.
Sargassum and Turbinaria species are used as fertilizers for palm
plantations in coastal areas of Kerala, India. Algifet (Norway), SM3
(England), Seasol (Australia), Bioweed (India) etc., are seaweed liquid
fertilizers available in the market. They have beneficial effects on
vegetable (bhendi, brinjal, tapioca, cucurbits) and fruit (lemon,
papaya) crops; and tree plantations (coconut, areca palm).
D.Phycocolloids: Brown and red seaweeds consist of high
molecular weight polysaccharides (colloids) in their cell walls. Those
amorphous mucilaginous substances have good water holding
capacity. Because of viscous nature and sol-gel forming capacity,
the phycocolloids finding many scientific and industrial
applications.
a. Alginic acid or Alginates: Alginic acid is a cell wall
polysaccharide obtained from all brown seaweeds; it is present in
middle lamella and primary cell walls. Kelps like Macrocystis,
Laminaria, Nereocystis, Ascophyllum, Ecklonia, Lessonia and Durvellia;
rock weeds like Sargassum and Turbinaria are the important
alginophytes. Alginic acid reacts with cations like Na+, K+, Ca2+, NH4+

APSCHE : SEMESTER-I BOTANY TEXT BOOK 297


    

        
     

    
 

      


       

    


   


     
       
      

 



298 APSCHE : SEMESTER-I BOTANY TEXT BOOK


to form salts called alginates. The colour of alginates varies from
white to yellowish-brown, sold in granular or powdered form.
Alginates are used in the textile industry for printing
process, paper industry to make wrapping papers; used for making
antibiotic capsules due to non-toxic property. Alginates are used in
food industry to thicken ice cream, jams, puddings, sauces, custards,
fillings, and decorations for bakery products. They are also used for
making flame-proof fabrics and plastic articles. Sodium alginate is
used in production of dental creams and impression materials, shoe
polish, welding electrodes, and oil drilling muds. Calcium alginate
is used to make synthetic seeds and for immobilization of microbes.
Calcium alginate is used in different types of medical products

b. Agar-agar: It is an amorphous polysaccharide, which forms


supporting structure in the cell walls of red algae, belong to
Gelidiales and Gracilariales orders. Gelidium, Pterocladia,
Acanthopeltis and Ahnfeltia abroad; Gelidiella and Gracilaria in India
are important agarophytes. Agar is insoluble in cold water but readily
soluble in hot water. A 1.5% solution of agar is clear and forms a
solid and elastic gel on cooling to 32 to 39 °C; not dissolving again
at a temperature below 85 °C. It is sold in the form of flakes, strips
and powder.
Agar is used in food industry to make puddings, ice creams,

is used in fruit preserves, clarifying agent in brewing, and sizing of


paper and fabrics, cosmetics. Agar is the gelling agent in culture
media for microbes and tissue culture. Agarose gel is an ideal
material for separation of DNA and proteins through
electrophoresis. Agar is an impression material in dentistry,

APSCHE : SEMESTER-I BOTANY TEXT BOOK 299


    

     
         
 
      
      
     

 


                     

 

     
    

  

   
  β      
     
      



300 APSCHE : SEMESTER-I BOTANY TEXT BOOK


substrate for precipitin reactions in immunology and an MRI elastic
gel phantom.
c. Carrageenan: It is a cell wall polysaccharide obtained from the
red algae belong to Gigartinales order. Gigartina, Iridaea,Chondrus
crispus, Eucheuma and Kappaphycus abroad; Hypnea, Acanthophora
and Laurencia in India are the important carrageenophytes.
Carrageenan is available of 3 commercial types, kappa, iota and
lambda, sold in powdered form.
Carrageenan is used as thickening, gelling and stabilizing
agent. It is used in the baking, dairy industries and in clarifying
liquors; an ingredient in cosmetics, shoe-polishes, shampoos, hand
lotions, tooth paste etc. This colloid is also used in paper and textile
industries. Carrageenan is used in pharmaceuticals to make tablets
and pills; to immobilize cells and enzymes in biotechnology.

Chemical nature of algal colloids: Alginic acid obtained from


brown seaweeds is a polymer composed of alternating mannuronic
and guluronic acid molecules. Agar and carrageenan are obtained
from red seaweeds. Agar is a polymer of galactose, made up of
agaropectin and agarose sub units. Agarose is a linear
polysaccharide, while agaropectin is a heterogeneous mixture of
smaller molecules. Carrageenan is a sulphated galactose composed
of subunits, carrabiose; anionic in nature due to 15-40% ester
sulphate content.

D. Diatomite: The cell walls of diatoms, frustules are composed of


silicates. Diatomaceous earth (diatomite or kieselguhr) is the
fossilized frustules forming mounds of sedimentary rocks in marine
habitats. Some diatomaceous deposits reported to be about 90
meters height, spread over a length of 1.6 kilo meters at a depth of
600 meters in the ocean. Diatomite is white coloured, firm, soft
and light weight material. It can be cut into blocks or brick shaped

APSCHE : SEMESTER-I BOTANY TEXT BOOK 301



 
        
   
      





      
       

          
       


 
    

 

    
 
 
 

302 APSCHE : SEMESTER-I BOTANY TEXT BOOK


structures, cast-off for different purposes. Diatomite is used as filter
for clearing lubricating oils and aviation fuels, for refining sugar;
insulator in boilers, furnaces, refrigerators, for making soundproof
rooms. It is also finding applications as abrasive in scouring and
polishing powders like tooth powder, bleaching powder, glass
cleaners, paints and varnishes; absorptive in handling and packaging
of hazardous materials.
E. Pharmaceuticals and nutraceuticals: The therapeutic use of
algae was mentioned in Chinese ‘Ben Cao’ (herbal encyclopaedia),
Indian Ayurvedic medicine and Materia medica. Antihelminthic
drugs are prepared from the red seaweeds like Chondria, Digenia
and Alsidium in China. Fucoidan and sodium laminarin sulphate
from brown seaweeds, carrageenan from red seaweeds are used as
an anticoagulants of blood. Codium and Corallina are used in Unani
medicine, to cure kidney, urinary bladder and lung ailments.
Chlorellin, an antibiotic from Chlorella is effective against both Gram
+ve and Gram -ve bacteria. The saxitoxins obtained from some
dinof lagellates are used in neurological disorders and
neurobiological research. Agarol, a by-product of agar is used as a
laxative.
Poly-unsaturated fatty acids (PUFAs) from microalgae serve
as precursors to second messengers (eicosanoids, docosanoids),
exploited to improve the brain functioning in humans. Linoleic,
arachidonic and eicosapentaenoic acid are reported in many
seaweeds, finding the utility as a dietary supplement. β-carotene
from Dunaliella salina and Astaxanthin from Haematococcus pluvialis
have nutraceutical value as vitamin -A precursors and antioxidants.
Single cell proteins (SCPs) from Spirulina and Chlorella cure the
Kwashiorkor, a nutritional disorder due to lack of sufficient protein
in the diet.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 303



 
  
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  

      
   



       
      
   


********

F. Bioenergy:

Jatropha, corn and sugarcane. Biodiesel and bioethanol are obtained


respectively by converting lipid and carbohydrate fractions in algae.
Microalgae like Botryococcus braunii, Dunaliella tertiolecta, Nitzschia
spp., Phaeodactylum tricornutum etc.; seaweeds like Sargassum,
Enteromorpha, Gracilaria etc., are reported to be promising species
for bioenergy. Research findings indicate that Isochrysis sp. is a
possible feed-stock for jet fuel.

304 APSCHE : SEMESTER-I BOTANY TEXT BOOK


G. Minerals: Brown algae of Laminariales order, kelps are rich
source of potash and iodine; ammonia and charcoal are also obtained
from those algae. 3-6% of Bromine can be extracted from red algae
like Polysiphonia, Rhodymenia etc., Seaweeds are a rich source of
minerals like iron, zinc, copper, manganese, boron etc.,
H. Waste valorization: Domestic and industrial wastes can be
disposed and converted to some valuable products using algae.
Microalgae like Chlorella, Spirulina, Chlamydomonas, Scenedesmus
etc., and some macroalgae are cultured in sewage water to produce
biomass, which can be used to produce some products.
I. Soil reclamation: Some green algae and many BGA grow on the
alkaline or saline soils during rainy season; form a thick stratum on
the soil surface, checks the soil erosion and increases fertility by
adding organic material.
II. Harmful impacts of algae: Certain species of algae may be
destructive under certain circumstances.
A. Harmful algal blooms (HABs): Excess of inorganic nutrients
from agriculture run-off and domestic sewage etc., lead to the over
growth of planktonic algae in aquatic bodies, called algal blooms.
Some species of such bloom releases toxins and cause threat to
heterotrophic biota and even humans, said to be HABs. About 40
species of algae can produce potent toxins, find their way through
fish and shell fish to humans. For instance, fresh water algae,
Microcystis aeruginosa and Anabaena flos-aquae produce microcystin
(hepatotoxic) and anatoxin (neurotoxin) respectively, that are highly
toxic to fishes and humans. In marine water, Lyngbya gracilis secretes
debromoaplysia toxin that causes dermatitis in human beings.
Trichodesmium erythraeum, Gonyalaux polygramma and Noctiluca
scintillans grow so dense to form harmless red tides; but are
responsible for indiscriminate death of fishes and invertebrates due

APSCHE : SEMESTER-I BOTANY TEXT BOOK 305


to oxygen depletion. Chaetoceros convolutes, Chrysochromulina
polylepis and Prymnesium parvum are not toxic to humans, but cause
fish deaths by damaging or clogging their gills.
B. Parasitic algae: A few green algae belong to Chlorococcaceae,
Phyllosiphonaceae, Phyllosporaceae and Trentepohliaceae are
parasitic on higher plants.Cephaleuros (Trentepohliaceae) is a
parasite causing ‘red rust’ on fruit yielding plants like mango, guava
etc. The older leaves of the host infected by the said parasite show
moderate to severe damage. Another green algal genus,
Chlorochytrium (Chlorococcaceae) is an endophyte in the
intercellular spaces of hydrophytes such as Lemna and Hypnum;
causes little or no damage to host. A marked stunting is reported in
Hibiscus sabdariffa infected by Rhodochytrium (Chlorococcaceae).
Phyllosiphon (Phyllosiphonaceae) is a parasite on members of
Araceae,resulting in large chlorotic lesions on the leaves of host
owing to cell death. The dinoflagellate, Oodinium spp., are parasitic
on fresh water fishes and marine tunicates. Species of a colourless
green algal genus, Prototheca cause a disease, protothecosis
(algaemia) in dogs, cats, cattle and humans.
C. Algal weeds: Free floating (Spirogyra), epilithic (Cladophora)
and epiphytic (Oedogonium) algae alone or in combination with
aquatic vascular plants can obstruct the usage of ponds, canals and
shallow areas of lakes; impair swimming, fishing and boating. Pelagic
species of Sargassum, S.muticum, S.fluitans and S.natans are invasive
and noxious in oceans, obstruct the passage of ships due to
uncontrolled proliferative growth. Blue green algae like Oscillatoria,
Lyngbya and Phormidium attach to cement linings of swimming
pools, cause problems in recreation.
D. Obstruction in water supplies: Algal growth in water
reservoirs and water supplies are responsible for some direct or

306 APSCHE : SEMESTER-I BOTANY TEXT BOOK


indirect problems. Oscillatoria, Spirogyra and some microalgae clog
the water filters; thus, shortening the filter runs leading to economic
loss. Growth of algae in water supplies may cause discoloration,
abnormal taste and odour due to their metabolic and decomposition
products. Carbonic, oxalic and silicic acids produced by Anacystis

Activity:Take the agar, agarose, alginic acid and carrageenan


available in botany or microbiology or biotechnology laboratory
of your college; study their physico-chemical properties, list out
their applications in those scientific fields. Take face creams,
toothpastes, frozen dairy desserts, imitation orange pulp,
packaged foods etc., in cupboards and refrigerators at home; read
the lists of ingredients on the labels of those products, find the
names of algal products.

Economic value of algal colloids:


Ø The global production of alginates is 44,480 tons which earned 728
million USD in 2020. Norway, Chile, China and USA are the principal
producers of alginates. Algaia, Marine Biopolymers Ltd., DuPont de
Nemours Inc., Ingredients Solutions Inc., KIMICA,Ceamsa,Algea,
Shandong Jiejing Group Corp.,are key manufacturers of alginates.
Ø The global harvesting of 1,96,364 metric tonnes of agarophytes
accounts fora market revenue of 268 million USD in 2019. Japan, Spain,
Chile, South Korea and Moracco are major countries for agar
production. Marine Hydrocolloids, Global BioIngredients, Sobigel,
Agarmex, Iberagar, Agar Brasileiro, Mingfu Fujian Agar Co., BandV
Agar, Fuli Agar Factory, Fujian Wuyi Feiyan Agar are the prime
manufacturers of agar.
Ø Global carrageenan market is 780 million USD in 2020 from about
70,000 metric tonnes of carragenophytes harvested. North America,
Europe, Asia Pacific, Central and South America, Middle East and Africa
are potential regions of carrageenophytes. DuPont, CP Kelco U.S.
Inc.,Aquarev Industries, Marcel Trading Corporation, ACCEL
Carrageenan Corporation, Cargill Incorporated, Ashland, TIC Gums,
Inc. are the principal producers of carrageenan.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 307


Self-Assessment:
1. The following pigments are water 7. Siphoneous habit is seen in
soluble. A. Ulva and Valonia
A. Chlorophylls B. Vaucheria and Coleochete
B. Carotenoids C. Sargassum and Ulva
C. Phycobilins D. Valonia and Vaucheria
D. Phytochrome 8. In algae with haplontic life cycle
2. Fucoxanthin pigment is a dominant meiosis occurs in
pigment in A. Zygote
A. Chlorophyceae B. Gametangia
B. Phaeophyceae C. Sporangia
C. Rhodophyceae D. None of the above
D. Dinophyceae 9. Haplo-biontic life cycle is present in
3. Oil and leucosin are the reserved food A. Batrachspermum
material in B. B.Gracilaria
A. Xanthophyceae C. Cladophora
B. Myxophceae D. Chlamydomonas
C. Euglenophyceae 10. Gametes of Spirogyra are
D. Chrysophyceae A. Biflagellated
4. Flagellated forms are not present in B. Non-flagellated
the following algal classes C. Multiflagellated
A. Chlorophyceae and D. Uniflagellated
Rhodophyceae 11. In which species of Spirog yra,
B. Phaeophyceae and asexual reproduction occurs by
Myxophyceae azygospores?
C. Rhodophyceae and A. S. farlowii
Myxophyceae B. S. aplanospora
D. Chlorophyceae and C. S.mirabilis
Phaeophyceae D. S. adnata
5. Name the Father of Phycology 12. The number of new thalli formed
A. Carolus Linnaeus from germinating zygospore of
B. A.W. Eichler Spirogyra.
C. F.E.Fritsch A. One
D. M.O.P.Iyengar B. Two
6. Dinobr yon consists of following C. Three
thallus D. Four
A. Unicellular 13. The sexual reproduction in
B. Parenchymatous Spirogyra is of following type
C. Dendroid colony A. Conjugation
D. Heterotrichous B. Copulation

308 APSCHE : SEMESTER-I BOTANY TEXT BOOK


C. Binary fission 20. Identify the alga used as a source of
D. D.Cleistogamy Bio energy.
14. Trichoblasts in Polysiphonia are A. Botryococcus braunii
A. Multicellular and uniseriate B. Haematococcus pluvialis
B. Multicellular and C. Spirulina maxima
multiseriate D. Chlamydomonas nivalis
C. Multicellular and
Interactive links:
unbranched
https://www.saps.org.uk/secondary/
D. Multicellular and biseriate teaching-resources/235-student-sheet-
15. The ploidy and nature of cystocarp 23-photosynthesis-using-algae-
in Polysiphonia wrapped-in-jelly-balls
https://lakes.grace.edu/research/blue-
A. Haploid and gametophytic green-algae/
B. Diploid and sporophytic https://www.forbes.com/sites/
C. Haploid and sporophytic jeffmcmahon/2019/05/28/algae-
single-celled-savior-of-the-climate-
D. Diploid and gametophytic
crisis/?sh=429d6fc855df
16. Meiosis occurs in following form https://microbiologysociety.org/why-
of Polysiphonia. microbiology-matters/what-is-
A. Gametophyte microbiology/algae.html
Online resources:
B. Tetrasporophyte
http://deskuenvis.nic.in/pdf/
C. Carposporophyte PhycologyLee.pdf
D. None of the above https://www.scribd.com/document/
17. The following is a parasitic alga, 463971679/A-Textbook-on-Algae-
PDFDrive-com-pdf
that causes rust disease. http://allaboutalgae.com/what-are-
A. Cephaleuros algae/
B. Chlorella https://pdfcoffee.com/linda-e-graham-
lee-warren-wilcox-algaepdf-pdf-
C. Tolypothrix
free.html
D. Gigartina Video Resources:
18. The following algal colloid is used https://www.youtube.com/
to make synthetic seeds. watch?v=CB2XlpD-Ld4
https://www.coursera.org/lecture/
A. Calcium alginate
algae/what-are-algae-GAiDG
B. Agar agar https://www.youtube.com/
C. Carrageenen watch?v=JIvOdfaLWNc
D. Diatomite https://www.youtube.com/
watch?v=HxhOTuz-MdI
19. Chemical nature of algal colloids https://www.youtube.com/
A. Polysaccharides watch?v=Dd8heneTj9I
B. Proteins https://www.youtube.com/
watch?v=ZRgeh7cN9PQ
C. Lipids
D. Terpenoids

APSCHE : SEMESTER-I BOTANY TEXT BOOK 309


References: Autospore: A non-flagellated spore
formed within the parent cell.
1. Fr itsch, F.E. (1945)The Auxospore: Specialized cells
Structure & Reproduction of formed in diatoms to restore cell
Algae (Vol. I & Vol. II) size.
Cambridge University Press Benthic:Inhabiting the bottom of
Cambridge, U.K.. an aquatic environment in a
2. Bold, H.C. & M. J. Wynne sedentary state.
(1984) Introduction to the Carpogonium: The female sex
Algae, Prentice-Hall Inc., New organ in red algae.
Jersey
Carposporophy te: A diploid
3. Robert Edward Lee (2008)
sporophyte in some red algae
Phycolog y. Cambr idge
University Press, New York produces haploid or diploid
4. Van Den Hoek, C., D.G.Mann carpospores.
& H.M.Jahns (1996) Algae : An Centric diatoms: Radially
Introduction to Phycolog y. symmetrical diatoms.
Cambridge University Press, Chemotaxis: Movement or
New York orientation of an organism or cell
5. Pandey, B.P. (2013) College in relation to a chemical substance.
Botany, Volume-I, S. Chand Chromatophore: A membrane
Publishing, New Delhi bound tubular structure containing
6. Hait,G., K .Bhattachar ya & pigments.
A.K.Ghosh (2011) A Text Book Coenobium: A colony having
of Botany, Volume-I, New definite number of cells with similar
Central Book Agency Pvt. Ltd., morphology and functions.
Kolkata Coenoc ytic condition: A
multinucleate condition due to
Glossary: absence of cross walls in a
multicellular/siphonous filament.
Accessory pigments: Light Colony: An aggregation of
absorbing pigments in indefinite number of similar cells
photosynthetic organisms, that with division of labour.
work in conjunction with Conjugation: Sexual reproduction
Chlorophyll-a. involving two non-f lagellated
Aero-taxis: Movement of an gametes (aplanogametes) of
organism or cell in relation to opposite sex.
Oxygen. Coralline algae:Red algae
Algal bloom: A nuisance algal associated with coral reefs and
grow th under nutrient rich contain calcareous deposits in cell
conditions. walls.
Anisogamy: The fusion of Dendroid: Tree like structure.
morphologically dissimilar Dioecious: A condition of having
gametes. separate male and female sexes of a
Antheridium: Male sex organ that species.
produces male gametes. Diploid: A condition of having two
Aplanogamete: A gamete without sets of chromosomes.
flagella and usually non-motile. Embryophytes: A plant group
comprises of br yophytes and

310 APSCHE : SEMESTER-I BOTANY TEXT BOOK


vascular plants, that produces Pennate diatoms: Diatoms having
embryo. bilateral symmetry.
Endosymbiont: An organism that Periphyton: Organisms that live by
lives inside another, benefitting attaching to a surface or substratum
both individuals. Phototaxis: Movement or
Epilithic: Growing on the surface orientation of an organism or cell in
of rock or stone. relation to light.
Epiphytic: Growing on the surface Phytoplankton: Free floating,
of plants or other algae. microscopic and unicellular plants.
Fragmentation: A condition in Pyrenoid: A proteinaceous body
which thallus breaks in to pieces, associated with reserve food in
meant for vegetative or asexual chloroplast of an alga.
reproduction. Rhizoid: Rootlike extensions
Frustule: The two valves and girdle usually used to attach a thallus to
bands of a diatom made of silica. the substrate.
Gametes: Single celled structures Spermatangium: The male sex
formed during sexual reproduction organ in red algae.
in algae. Spores: Single celled structures
Haploid: A condition of having one formed during asexual reproduction
set of chromosomes. in algae, if motile called zoospores,
Holdfast (hapteron): A structure if non-motile called aplanospores.
that attaches a thallus or filament Statospores: A siliceous thick-
to a substratum. walled resistant cyst formed within
Hypnospore: A thick-walled the frustules of marine centric
resting spore. diatoms.
Isogamy: The fusion of Sub-littoral: A zone just below the
morphologically similar gametes. shore of a sea.
Littoral: A zone on or near the Tetrasporophyte: A diploid
shore of a sea. sporophyte in some red algae that
Monoecious: A condition of having produces haploid tetraspores by
male and female sex organs on same meiosis.
plant, but in different places. Thallus: A plant body devoid of
Oogamy: The fusion of a motile root, stem and leaves.
male gamete with a non-motile Thylakoid: A flattened sac present
female gamete. in chloroplastthat contain pigments
Oogonium: A female sex organ in cell membrane, participate in
(usually single celled) without a both light and dark reactions of
sterile jacket, produces an egg cell. photosynthesis.
Parthenospore (azygospore): A Trichogyne: An elongated portion
spore/gamete that produces a new of the female reproductive organto
thallus without fertilization. receive the male gametes in red algae
Pelagic: A floating state of life in and charophycean algae.
surface water of sea or ocean. Trichome: A filament of a blue
Pellicle: The surface covering of green alga.
euglenoid cells made up of Zygospore: A thick-walled resting
proteinaceous strips in a helical spore formed from a zygote.
pattern.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 311


Unit - 5
Bryophytes

CONTENTS LEARNING OUTCOMES:


1. General characteristics At the end of this unit learners should be
of Br yophytes and able to:
classification 1. Understand the general
characteristics of parenchymatous
2. a. Marchantia
tiny plants of plant kingdom.
(Hepaticopsida) and
2. Develop a critical knowledge
b. Funaria (Bryopsida)
through a detailed inquiry of two
3. General account on Bryophytes.
evolution of 3. Recall and explain the
sprophytes in evolutionary trends among
Bryophyte amphibious plants for their shift
to land habitat.
4. Evaluate the ecological and
economic value of Bryophytes.

Bryophytes are tiny plants of thalloid nature to leafy, that grow


like a green carpet, and are inconspicuous to the naked eye in moist
shady places. Due to the absence of typical roots, vascular tissues,
mechanical tissues and cuticle they can’t attain great heights.
Though they are terrestrial in establishing, require external water
to complete their life cycle. Hence, they are called “Amphibians of
the plant kingdom.” Compared to seed plants, Bryophytes are
widely distributed throughout the world. They are less
independent of their environment, unlike other land plants. They
are conspicuously absent in the seas, but some present in salt

(charophyte), indicate the non-monophyletic development into


the vascular land plants, Pteridophytes. However, Bryophyte
lineage, i.e., the liverworts, hornworts, and mosses is

312 APSCHE : SEMESTER-I BOTANY TEXT BOOK


monophyletic from a single common ancestor. Due to the soft
parenchyma tissue, fossilization is very rare. So, the fossil record
of bryophytes has a poor history. Their report on fossils is from
the upper Carboniferous period was by Walton in 1925. Hence,
the existing bryophytes preserving ancestral characteristics can
give us some insight into the origin of land plants. Bryophytes
show an alternation of generations between the independent
gametophytic phase which produces the sex organs, and the
dependent sporophytic phase which produces the spores after
reduction division. The multicellular embryo develops inside the

is differentiated into foot, seta and capsule. In the capsule, haploid


meiospores of similar types are produced (homosporous). The
spore germinates directly into gametophyte in Marchantia,
Anthoceros and into juvenile gametophyte called protonema,
which is produced in buds that gives rise to adult gametophores.

Chapter -1 General Characteristics of Bryophytes and their


Classification.

Learning outcomes: At the end of the chapter you should be


able to:
1. Describe the general characteristics and classification of the
amphibians of the plant kingdom.
2. Compare and contrast the characteristics of Bryophytes with
that of Thallophytes and Pteridophytes.

5.1.1 General Characteristics of Bryophytes


Occurrence :
rocks, bases of trees or similar wet places, with the exception of few
water forms (Water moss -Fontinalis). Thalloid plants vary in size
from a length of 20 cm and a breadth of 5 cm. The smallest liverwort

APSCHE : SEMESTER-I BOTANY TEXT BOOK 313




 
   
 
        



 
     
 



Do you know: Bryophytes are the connecting links from
aquatic Algae to terrestrial vascular Pteridophytes.

: The division Br yophyta (Gr. br yon=moss) includes


nonvascular embryophytes such as mosses, liverworts and hornworts over
25000 species. Bryophytes -a traditional name for any nonvascular,
seedless plants. The study of bryophytes is called bryology. Hedwig is called
the ‘Father of Bryology’. Shiv Ram Kashyap is the ‘Father of Indian
Bryology’

314 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Sphaerocarpos Fontinalis, if reclining, can
reach lengths of more than 1 metre. Some, however, are less than 1
Ephemerum). The plant body is compact and
better protected against desiccation. The greatest diversity is at
tropical and subtropical latitudes. Bryophytes are almost
cosmopolitan. Thus they represent an extremely ancient group of
plants due to the distribution patterns.
Splachnaceae
decomposing animal waste, and in somewhat shaded cavern mouths
the liverwort Cyathodium Mittenia, on leaf surfaces
Ephemeropsis Metzgeria,
Carrpos Aschisma bases of quartz
Scopelophila on copper-rich substrata. Despite
being so adaptable, they have one limitation, they require free water,
not just soil moisture in their environment. Without it, they cannot
reproduce sexually.
Fact: Dawsonia superba, a bryophyte that resembles a pine tree

Activity: Observe and collect bryophytes during rainy season on the


old walls, tree barks, rocks and record them with your cell camera.

Alternation of generations: The gametophyte is the dominant


plant body on which the sporophyte is semi-parasitic for its
nutrition. The gametophyte form shows several developmental
stages: the spore, the protonema, and the gametophore, which
produces the sex organs. The gametophyte is conspicuous, usually
perennial and photosynthetically independent of the sporophyte.
The sporophyte forms an intimate interconnection with the
gametophytic tissue (Fig: 5.1.1).
Plant Body: Gametophyte : The gametophyte is thalloid in habit
and is not differentiated into roots (but of rhizoids), axis (stem)

APSCHE : SEMESTER-I BOTANY TEXT BOOK 315



  

       
       
 
       


   
  

       



5.1.1 : Alternation of generation


https://cnx.org/contents/eXo5eCdH@7/Bryophytes

316 APSCHE : SEMESTER-I BOTANY TEXT BOOK


and leaves. It is relatively simple in the lower forms, like the thallus
of Alga, prostrate on the ground, linked to the substrates with
unicellular rhizoids. The leafy liverworts are dorsiventral; in mosses,
it is erect. In Jungermanniales, neither thalloid nor perfectly foliose.
The leafy moss gametophyte has a stem-like central axis bearing
leaf-like appendages, fixed to the substratum by branched,
multicellular rhizoids. Branching of the thallus may be forked,
regularly frond-like, digitate, or completely irregular. The margin
of the thallus is often smooth (Liverworts) but toothed (Pellia). It
may be ruffled, flat, or curved inward or downward (Fig: 5.1.2;
5.1.3 & 5.1.4).

5.1.2 Liverwort
https://nhpbs.org/natureworks/nwep14c.htm

5.1.3 Hornworts
https://upload.wikimedia.org/wikipedia/ 5.1.4 Moss
commons/d/d1/ https://www.wikiwand.com/en/Moss
Hornwort_%283144429129%29.jpg

Porella, the stem is covered by three rows of leaves, two on the


dorsal side and one on each side of the stem. The third row is on

APSCHE : SEMESTER-I BOTANY TEXT BOOK 317


5.1.5 Spagnum
https://commons.wikimedia.org/wiki/File:Sphagnum_moos.jpg#/media/
File:Sphagnum_moos.jpg

5.1.6 porella
https://upload.wikimedia.org/wikipedia/commons/e/e6/Porella_platyphylla.jpg

     



 
      


   


  

318 APSCHE : SEMESTER-I BOTANY TEXT BOOK
the ventral side with small leaves called amphogastria. Leaves are
without a midrib. Leaves show great variation in form. In moss,
leaves are spirally arranged on the stem, with a single midrib.
(Fig:5.1.5 & 5.1.6).
Internal structure : Bryophytes lack complex tissue, and they
show considerable diversity in form. The thallus is simple without
tissue differentiation. In Anthoceros, each cell has a chloroplast with
a pyrenoid (Fig:5.1.7 & 5.1.8). Rhizoids are typically found on
the ventral surface of the plant, developing from either specialized
cells of the under leaves or the stem epidermis. They are generally
hyaline. Liverworts are unique in having membrane-bound oil
bodies, which synthesize an array of terpenoids and other aromatic
compounds. However, they will disappear in dry samples due to
their volatile character. These oil bodies prevent grazing, protect
from cold and UV radiation. In addition, the sequester terpenoids
and other secondary aromatics are like the secretory glands of
vascular plants. Vascular tissues xylem and phloem are absent in
lower groups and in Mosses, some possessing conducting tissue
hydroids leptoids,
stems and leaves of vascular plants.

Fig 5.1.7 Thallus of Riccia


https://extraclass.com/e/neet-plant-kingdom-3_Riccia

APSCHE : SEMESTER-I BOTANY TEXT BOOK 319


Fig 5.1.8 Anthocerosthallus T S
https://www.biologydiscussion.com/wp-content/uploads/2016/02/image-168.png

      


    

      



 



 

     

   

320 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Cellular Cryptogams: Bryophytes along with Thalophytes

As a special relationship with algae, the growth of the thallus will


be by an apical cell. Four types of apical cells occur in hepatics,
namely, tetrahedral (or pyramidal), cuneate (or wedge-shaped),
lenticular (orlens-shaped) and hemidiscoid type.

Activity: Find out the shapes of the apical cells and their cut
surfaces by which the new cells can be developed.

Reproduction : A) Extensive Vegetative propagation takes place


during the growing season (Fig: 5.1.9).
a. By death and decay of older parts- Porella
b. Persistent apices - Riccia
c. Adventitious branches - Anthoceros
d. Gemma- Marchantia
e. Tubers- Anthoceros
f. Primary Protonema- Funaria
B) The gametophyte with the sexual reproductive system bears
multicellular and jacketed male and female sex organs called
antheridia and archegonia. Sexual reprovduction is an oogamous
type. The male gametes (antherozoid) are motile, whereas the
female gametes (Egg) are large & non-motile. They are produced
within the sex organs. (Fig: 5.1.10).
The male sex organ, antheridium, has a single-layered outer
sterile jacket, like in Algae with a mass of antherozoid mother cells
called androcytes, each of which gives rise to a single ciliated motile
antherozoid. (Fig: 5.1.11). The female sex organ is a multicellular,
more or less flask-shaped organ. It has a swollen basal portion called
the venter and a slender, elongated upper portion called the neck.
The archegonium consists of an axial row of cells surrounded by a

APSCHE : SEMESTER-I BOTANY TEXT BOOK 321


   


       





Fig 5.1.9 structure of reproduction


https://www.brainkart.com/media/article/article-Bryophytes-iZj.png

       





     
  


322 APSCHE : SEMESTER-I BOTANY TEXT BOOK


sterile jacket. An axial row consists of a variable number of neck
canal cells and ventral canal cells, which is an identification feature
of the plants. The archegonium provides nourishment and
protection to the egg and to the later developing embryo.

Fig 5.1.10 sex organs


https://files.askiitians.com/cdn1/cms-content/biologyplant-
kingdombryophytes_5.jpg

5.1.11 antheridia& Archegonia


https://extraclass.com/e/neet-plant-kingdom-3_Riccia

Archegoniatae : Archegonium becomes a common feature along


with of other higher plants such as Pteridiphytes and Gymnosperms.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 323


   

      

Sporophyte: Water is essential for fertilization, as the ciliated
antherozoids swim to the neck of the archegonium, pass through
the neck canal and fertilize the egg. This diploid cell begins to grow
immediately and develops into a multi cellar sporophyte, which is
retained within the archegonium and protected by the ‘calyptra’,
the basal part of the archegonium as a cap. The sporogonium is
differentiated into foot, seta and capsule. But foot, seta is absent
Riccia Marchantia In Mosses,
the seta elongates and forms a tough structure bearing the capsule.
(Fig: 5.1.12 & 5.1.13).
The zygote develops into a multi-layered embryo with
amphithecium and endothecium layers. The endothecium forms
into archesporium in Riccia and Moss, whereas in Anthocerotales,
it is formed by amphithecium. The spore mother cells, which are
originated from the archesporium, divide by reduction division to
form the spores and are released from the capsule. The spores are
released by a special mechanism such as elaters, pseudo elaters and
Riccia, sporogenous cells give rise to spores
Marchantiato, spores and elaters.
Anthoceros, alternative bands of sporocytes and sterile cells. In
Mosses, the sporogenous tissue is limited.
Spore is non-motile and cutinized. They germinate directly
into a gametophyte plant ( called
protonema. In Mosses, they germinate into a filamentous
protonema which gives rise to adult gametophores through buds
(Funaria, Polytrichum (Fig: 5.1.14).

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5.1.12 moss sporophyte
https://www.logicalclass.com/contant/view/115

5.1.13sporophyte comparision
https://plantscience247.blogspot.com/2020/07/comparison-of-sporophyte-of-
bryophytes.html

APSCHE : SEMESTER-I BOTANY TEXT BOOK 325


5.1.14 protonema
https://www.researchgate.net/
figure/The-protonema-of-
Physcomitrella-patens-consists-
of-chloronemal-and-caulonemal-
cells_fig1_51385913

Fact: The sporogenous tissue is reduced progressively from lower


groups to higher bryophyte groups. Those Bryophytes follow
heterologous haplo-diplobiontic type of life cycle.

Activity: Find out the function of the seta and foot in the life
cycle of Bryophytes.

5.1.2 Classification of Bryophytes : According to ICBN


(International Code of Botanical Nomenclature), bryophytes have
been divided into three classes.
1. Hepaticae( Hepaticopsida = Liverworts)
2. Anthocerotae (Anthocertopsida= Hornworts)
3. Musci (Bryopsida= Mosses)
Class 1. Hepaticae or Hepaticopsida:
1. Gametophytic plant body is either thalloid or foliose. If
foliose, the dorsiventral and lateral appendages without mid-
rib (leaves) are present.
2. Aseptate Rhizoids.
3. Thallus cell contains many chloroplasts without pyrenoids.
4. Sex organs are embedded in the dorsal surface.
5. Ricciawith only a capsule.
Marchantia, Pallia and Porellait, it is differentiated into

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foot, seta and capsule, capsule without columella and with
elaters. It has four orders:
(i) Calobryales (ii) Jungermanniales
(iii)Spherocarpales (iv) Marchantiales.
Class 2. Anthocerotae or Anthocerotopsida
1. Gametophytic plant body is simple, thalloid, without air
chambers, and internal differentiation of tissues.
2. Scales are absent in the lower epidermis.
3. Cells possess a single large chloroplast with a single pyrenoid.
4. Sporophyte is cylindrical and partially dependent on the
gametophyte for its nourishment. It is differentiated into a
bulbous foot and cylindrical capsule. Seta is meristematic
and shows continuous growth.
5. Endothecium forms the sterile central columella in the
capsule, Pseudo elaters. It has only one order -
Anthocerotales.
Class 3. Musci or Bryopsida
1. Gametophyte is differentiated into protonema and
gametophore.
2. Gametophore is foliose, differentiated into stem and lateral
appendages like leaves without a midrib.
3. Multicellularrhizoids with oblique septa.
4. Elaters are absent in the capsule of the sporangium.
5. The sex organs are produced in separate branches immersed
in a group of leaves. It has only three orders:
(i) Bryales (ii) Andriales and (iii) Sphagnales.

Activity: Identify the adaptations of bryophytes for these problems


in establishing as land plants

APSCHE : SEMESTER-I BOTANY TEXT BOOK 327


Know more : The establish on land the plants the problems of
life were preventing death by drying out, dispersing spores through
the air, and avoiding damage from weather and solar radiation.
However, these bryophytes have pioneered, and were subsequently
passed on to all their descendants.

5.1.3 Bryophytes Origin As there is no clear evidence of fossils,


the origin of the Bryophytes origin is something like a hypothesis.
Based upon the comparative studies of morphology, habitat and
ontogeny of other living organisms, Bryophytes might have
originated from algae. However, there are two views on their origin.
(A) Hypothesis of the Algal origin.
(B) Hypothesis of the Pteridophytes origin.
Algal Origin : Bryophytes share more features with Chlorophyta;
both have the undifferentiated thalloid structure without vascular
tissue. The predominant stage in the life cycle is the gametophyte.
The filamentous nature of the protonema in the moss is like the
algal filamentous nature. Further, the photosynthetic pigments,
cellulose cell walls, starch as reserve food material, and the nature
of the flagella in the antherozoids are also similar to Algae. Fritsch
considered that Chaetophorales among the Chlorophyceae give rise
to transitional organisms having heterotrichous habits leading to
the thalloid Liverworts. The development of the parenchymatous
nature as in Pheophyceae ( apical growth, dichotomous
growth also supports the origin from algae. Further, the reseeding
waters of the ocean led to the appearance of the land parts,

Pteridophytes Origin : Evolved from Pteridophytes by progressive


simplification or reduction. The characters are :

328 APSCHE : SEMESTER-I BOTANY TEXT BOOK


1. Presence of stomata and chloroplast in cells

2.
3. zygote development into a multicellular embryo,
4. Equisetum

5.
6. Sporangia of moss resemble the Psilophytales of
Pteridophytes.
Kashyap has supported this and explained the retrogressive
evolution of these plants from Equisetales of Pteridophytes in three
independent lines as Marchantiales, Jungermanniales, and

Origin of sporophyte Regarding the origin of sporophyte in


bryophytes, there are two theories – homologous and antithetic
theories.
(A) Homologous theory
modification of the gametophyte. Because of the presence of
isomorphic alternation of generation, both are morphologically
similar, like cladophora of algae. The presence of photosynthetic
tissue in the sporophyte emphasizes the similarity with
gametophyte.
(B) Antithetic theory
intercalated between two gametophytic generations. Simple
sporophyte (Riccia)
of the sporogenous tissue as in higher mosses, with the presence of
chlorophyll, stomata, formation of peristome for dissemination of
spores, and columella as advance features. The sterilized tissue is
thus diverted to carry assimilation and effective spore dispersal.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 329


5.1.4 Economic Importance of Bryophytes
A. In Ecology and pollution:
1. By growing on sandy soils and on steep mountain slopes, they
can prevent soil erosion. The rhizoids and the moss stems build the
soil particles together; even the protonema stage forms the mat
and prevents soil erosion.
2. They play a major role along with the Lichens in soil formation
too. In rocky areas, mosses play a vital role in plant succession. The
peat mosses as mats established on the banks of the lake or open
waters giving the appearance of the quacking bogs, these mats,
because of the moisture and the humus, form as a suitable
substratum for germination of various hydrophytic plants,
ultimately changes in the landscape takes place. Certain mosses
Bryum grow in association with other aquatic plants where
calcium carbonates are in large amounts. They decompose the
bicarbonic ions and precipitate with other insoluble minerals to form
hardened rocks around the plants.

Conocephalum Candida albicans was


Sphagnum
decease.
4. Both the live and dried mosses can absorb metal complexes enable
them to be used as atmospheric and aquatic pollution indicators.
Along with the Lichens, bryophytes can be employed in developing
an Index of Atmospheric Purity (IAP), which is based on the number,
frequency and resistance factors of the species. Some Bryophyte
will show some visible symptoms due to the minute quantities of
pollutants, whereas some can absorb and retain the pollutants. This
approach can give a fair idea of the nature of the pollution in a given
area.

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B. Sph sps are being used in making bread
and as poultry feed.
C. The Sphagnum moss forms as peat in the marshy areas and can
be used as fuel in the furnaces as they generate a high amount
of heat.
plants were sued as bandage
cotton.
E. Sphagnum
fresh flowers and plants as they can retain the moisture for a
long duration.

know more :Importance of the study of Bryophytes: Technological


advances in understanding the alternation of generations in plants
moreover from the gametophytic dominance to sporophyte
establishment in the higher plants from Pteridophytes, bryophytes
have proven to be invaluable organisms for this research. They
continue to play a central role in new developments as Bryophytes
are occurred on specific pH of the substrate and indicate the air
quality significant as the ecological indicator. Chemical
constituents of bryophytes attract much attention.

Campbell and Takhtajan classified the Bryophytes into three classes,


Hepaticopsida, Anthocerotopsida and Bryopsida. Schimper grouped
them as divisions. Engler classified into two divisions and each division
into three orders i.e., Hepaticae ( Marchantiales, Jungermanniales, and
Anthoceratales) and Musci (Sphagnales, Andrecles, and Bryales).
Rathmiller changed them as Hepaticopsida, Anthoceratopsida and
Bryopsida as per international code of Nomenclature.Due to the peculiar
characters Anthocerate was created on par with Hepaticae and Musci.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 331


Chapter -5.2 Marchantia and Funaria

5.2.1. Marchantia

Learning outcomes : At the end of the chapter you should be able


to:
1. Explain the morphology anatomy reproduction and life
cycles in the typical study of thalloid liverwort– Marchantia
and the evolved Moss (Funaria).
2. Discuss and debate the similarities and difference between
bryophyte of two different classes

Class : Hepaticopsida
Order : Marchantiales
Family : Marchantiaceae

Occurrence : Marchantia
Marchantia polymorpha
Himalayan hill range and southern hills have this sp., about 11 sp.
were reported in India. These plants grow in moist places, wet open
lands, banks of the streams and on the shaded cliffs abundantly.
They grow on the damp burnt soils by the forest fires due to the

FACTS: As the thallus of Marchantia looks like liver lobes it is


called, liverwort. This is an advanced taxon in that family.

Adult Gametophyte phase : The plant body is gametophyte,


dorsiventral, prostrate, with dichotomous branching. The thallus
has a central midrib n the dorsal side with a notch at its apex. This
is the growing point of the thallus. The upper surface is marked by
rhomboidal or polygonal areas known as areolae. They indicate the

332 APSCHE : SEMESTER-I BOTANY TEXT BOOK


outline of the air chamber below it. The air chambers have an
opening called an air pore. Along the midrib, cup-shaped structures
are present called gemmae. These gemmae cups have numerous
multicellular bodies called Gemmae useful for vegetative
reproduction. On the ventral side, numerous rhizoids and
multicellular scales are present. The rhizoids are of two types,
namely smooth-walled and pegged walled. Besides the rhizoids,
purplish flattened scales are arranged in two or four rows on either
side of the midrib. As the scales are standing close arrange together,
they retain water by capillarity and protecting growing tip. (Fig:
5.2.1).

different thalli. On attaining sexual maturity, the thallus develops


upright, stalked, umbrella-shaped structures bearing gametangia
from the apical notch of the vegetative thallus called
antheridiophore or archegoniophore. The stalked structures have
eight lobed discs representing the apex of a branch, which is massive
and bearing the antheridia or archegonia in eight rows. The
antheridia are in acropetal succession, the oldest being nearest the
centre, youngest towards the outer extremity of the lobe facing
upwards. Whereas the archegonia are also arranged in a similar
manner in the early stage and after fertilization due to the
overgrowth of the lobe in the central region more than the margin
of the lobes, the archegonia pushed to the lower surface. Thus, the
archegonia are hanging with their necks downwards. (Fig: 5.2.2).

Activity: Collect the liverworts in your area and record the


morphological characters and take a cross-section of them and
observe the internal characters. Identify to which class they
belong.

APSCHE : SEMESTER-I BOTANY TEXT BOOK 333




 
  

        



   
  
      





5.2.1 a) Marchantia thallus


Source: https://cutt.ly/6Q9zMcn

334 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig 5.2.2) Marchantia female and male plants
Source:https://cutt.ly/sQ9cUm5

Internal structures : Marchantia


three regions; these are the epidermal region, the Photosynthetic
region and the storage region. The upper epidermal region is single-
layered thin-walled cells with chloroplasts. This layer is interrupted
by barrel-shaped air pores. Each pore is surrounded by four or five
superimposed tires cells forming a raised pore above the surface of
the thallus. Each pore leads into the air chamber below it. They
resemble the stomata of higher plants without an opening and
closing mechanism. They are only analogous in nature. They are
useful for gas exchange into the thallus. The lowermost layer of the
storage region functions as the lower epidermis and is single-layered
without air pores. The cells produce unicellular rhizoids and
multicellular scales. The rhizoids are two types, smooth and
tuberculate (pegged) inner walls. The scales help to retain the
moisture below the thallus. (Fig. 5.2.3 a, b)

chambers are present. The chambers are bounded by one layered


partition. Each chamber opens out through the air pore. From the
floor of the chamber arise simple or branched green filaments with
abundant chloroplasts. This region constitutes the centre of
photosynthesis in the thallus. According to Barnes and Land, the

APSCHE : SEMESTER-I BOTANY TEXT BOOK 335



    
 
       
  



  
      
        
       


 

      
       



  


       


336 APSCHE : SEMESTER-I BOTANY TEXT BOOK


chambers are produced schizogenously. The storage region is below
the photosynthetic region on the ventral side of the thallus. It is
made up of large, colourless, thin-walled, compactly arranged
polygonal parenchymatous cells. They usually lack chloroplast. They
possess starch and protein grains. Some isolated cells contain large
oil bodies or are filled with mucilage.

growing point is present. They have the horizontal row of


meristematic cells giving cutting surfaces on dorsi ventral surfaces.
The dorsal cells differentiate into the upper region of the thallus,
whereas the lower cells form the storage region. By the longitudinal
division of the apical cell, dichotomous branching takes place.
Reproduction : Marchantia reproduces by vegetative, asexual and
sexual methods.
Vegetative Reproduction: Thallus show vegetative reproduction

A) Fragmentation of the thallus is by the rot and decay of the old


vegetative parts and separating the lobes at the dichotomy into
independent plants.
B) Formation of adventitious branches, particularly from the ventral
surface and sometimes at the archegoniophore stalk. When
detached by the decay of the connecting tissue, they become
independent thallus.
C) The third and specialized method of vegetative reproduction is
by a characteristic special asexual method in the Marchantia is the
They are produced in the small cups called
gemmae cups, which are present on the dorsal surface midrib region.
Gemmae are small, green, disc-like biconvex structures having a
small one-celled stalk. Intermingled with the gemmae are mucilage
hairs. These hairs will swell due to the absorption of rainwater and

APSCHE : SEMESTER-I BOTANY TEXT BOOK 337


Fig. 5.2.3 a) thallus internal structure. b) T.S of Thallus
Source: https://img.brainkart.com/extra2/JW1zoEY.jpg

Fig. 5.2.3 b) T.S of Thallus


Source: https://img.brainkart.com/extra2/JW1zoEY.jpg

      


   

        

   
        
 
338 APSCHE : SEMESTER-I BOTANY TEXT BOOK
release the gemmae from the cup. The detached gemma grows as a
new thallus on reaching the suitable substratum. (Fig 5.2.4 a,b,c).

Structure of the gemma: Gemma is a multicellular lens-shaped


discoid body with two growing points in the side of the margin with
the notches. All the cells are full of cytoplasm with abundant
chloroplasts. Isolated cells will have oil bodies. Some cells of both
surfaces are colourless and known as rhizoidal cells. There is no
dorsiventral differentiation in this disc. When the falls on the
suitable soil, the rhizoidal cells develop rhizoids, and the apical cell
at the growing points will simultaneously grow in opposite
directions. Rotting of the old parts ultimately develops into two
plants. Gemmae formation is rapid in the growing season. As
Marchantia is dioecious, female plants that produced gemmae on
germination will give rise to female plants only and similarly male
plants gemmae.
Sexual reproduction : Marchantia diecious, and sexual
an oogamous
the growing season. Sex Organs are born on the specialized vertical
branches called gametangiophores which are terminal in position
with limited growth. They develop on different thalli and are called
antheridiophore and archegoniophore; respectively,
Antheridiophore bears male sex organs e called antheridia, and
archegonia bear female sex organs archegonia. The internal
structure of these gametophores is similar to that of the vegetative
thallus, except that these are vertical in position. There are reports
of the branched gametophores in certain sp of Marchantia.
A. Antheridiophore and Antheridium : Antheridiophore has an

with eight lobes and teethed margins. In the cross-section, the stalk
is similar to the thallus, with air chambers on the dorsal side and

APSCHE : SEMESTER-I BOTANY TEXT BOOK 339



      

       

   
  
  


Fig 5.2.4 a) Marchantiagemmae cups


b) Cross section of thallus with gemma cup and the gemma
Source: https://cutt.ly/XQ9btXe
Source: https://cutt.ly/iQ9bLKL

Fig 5.2.4 a (c) Gemma.


Source: https://cutt.ly/BQ9QUdV

340 APSCHE : SEMESTER-I BOTANY TEXT BOOK


rhizoids on the ventral side. In the terminal receptacle, each lobe
has its growing points indicating the dichotomy branching in quick
succession. On the upper side of the receptacle, air chambers are
arranged and possessing photosynthetic filaments. The antheridia
are deeply sunken in the flask-shaped antheridia chambers
alternating with the air chambers, with a narrow opening called
the ostiole. These are arranged in eight radial rows in the acropetal
(Fig 5.2.5 a,b)
Each antheridium is ovoid and has one short stalk. The body
has a one-celled thick jacket layer of sterile cells covering
androcyte mother cells. Each androcyte mother cell divides
androcytes
antherozoids or sperm.
sperm is a small curved structure with a nucleus and thin cytoplasm.
The antheridium dehisces with water or dew. When water moves
into the antheridia cavity through the ostiole, the upper part of the
jacket cells will be disintegrated and ruptured. A slimy mass of
androcytes comes out of the chamber and spreads into the water
layer. Soon the antherozoids are liberated from them. Two flagella
are at one end and are pointed backwards, which allow the sperm

Similar to antheri-
diophore, the female thallus develops archegonio-
(carposcephalum) at the tips of the growing branch.
Archegoniophore has an upright stalk and an apical receptacle. The
receptacle has a conspicuous eight lobes. The growing apical regions
will bend downwards and inwards towards the stalk from the
margins of the disc. The internal structure is similar to that of the
vegetative thallus with air chambers and the air pores on the dorsal
side and rhizoids on the ventral side. Due to the three successive

APSCHE : SEMESTER-I BOTANY TEXT BOOK 341



 
  
   n


      

     
       

       
       
     
  
  
 

Fig 5.2.5
a)thallus with antheridiophores and b)Cross section of the antheridium.
Source: https://cutt.ly/yQ9Wh4g , Source: https://cutt.ly/2Q9WEmP

342 APSCHE : SEMESTER-I BOTANY TEXT BOOK


dichotomy division, eight lobes are developed on the apex of the
receptacle. And eight groups of archegonia develop on the surface
of the disc corresponding to the growing tips in acropetal succession.
In each row, there may be 10 - 15 archegonia. In this stage, the
archegonia are upwards and become mature, and fertilization can
take place at this stage. In this, the young archegonia are in the
margins. (Fig 5.2.6 a,b)
Know More: With the special structures of perichaetium and the
rays we can easily identify the male and female plants of
Marchantia.

After fertilization, due to the growth of the stalk and in the


centre of the disc more than the margins, the archegonia pushed to
the lower side of the disc. Now the young archegonia will be in the
centre near the stalk, and the mature archegonia are in the
periphery. The neck is directed downwards. Due to these bending
archegonia will be inversed. After the curvature, one layered plate
of tissue will be developed on either side of the archegonia. Thus,
each archegonia group is enclosed by a two-lipped pendent
perichaetium
hanging down vertically below the lobe. In some species, there is
an intercalary development of long, stout, green, cylindrical
processes from the periphery that takes place and is known as rays.
This structure looks like an umbrella.

Fact: The presence of six rows of jacket cells in the neck is the
characteristic feature of the Marchantiales. The Number of neck canal
cells are specific and used in species identification.

The mature archegonia are flask-shaped structures with a


short stalk. It has a venter and a long neck. Venter is surrounded by
a single-layered wall that encloses a single large egg cell and a venter
canal cell. Neck has six vertical rows of jacket cells enclosing 4-8

APSCHE : SEMESTER-I BOTANY TEXT BOOK 343



   
    
       

     
 n n 

  


 
  
 
        

   
  

    

   
 

   
 


344 APSCHE : SEMESTER-I BOTANY TEXT BOOK


axial neck canal cells. The tip of the neck is covered by a rosette of

neck canal cells and the ventral canal cell shows signs of degeneration
with the presence of moisture. On absorption of the moisture, this
cellular material will change into mucilage and exert pressure on
the cover cells to apart. This will make a passage for the flagellated
sperm to reach the egg and fertilize it. Attracted by the mucilage,
the sperms will move towards the archegonia.
C. Fertilization :
fertilization has to be very efficient. The presence of water is crucial
for the process. The liberated sperms reach the archegonia in various
ways.
i. The male disc upper surface is a little concave. By the splash of
raindrops on this facilitates the sperms to disperse and fall on
the neighbouring female receptacle.
ii. The sperm fallen off the ground can swim in the water medium
to reach the archegonia.
iii. In the water submerged male and female plants the sperms
can swim and reach the archegonia
iv. Small insects like mites and springtails will be attracted by
the mucilage secreted by the paraphyses can facilitate
fertilization.

Zygote. Following the


fertilization, the stalk of the archegoniophore elongate and produces
rays. Many archegonia can be
fertilized in each receptacle, but all of them will not develop. The
calyptra. From
the base of the venter a cylindrical sheath known as
perigynium

APSCHE : SEMESTER-I BOTANY TEXT BOOK 345



       







Fig 5.2.6 a) Marchantia archegonia thallus and


b) Cross section of archegoniophore with archegonia
Source:https://www.flickr.com/photos/gjshepherd/2883149186
Source; https://cutt.ly/lQ9Pma6

* * * * * * * *
archegonium and gives protection to the embryo against drought.
At this stage, the zygote is called young sporangium. The young
sporangium is surrounded by three protective sheaths of
gametophyte- calyptra, pseudoperianth and perichaetium.
The zygote(2n) will be the first cell of the sporophyte.
The zygote undergoes one transverse division and one vertical
quadrant. The upper two
epibasal cells, and the lower two cells are
hypobasal cells. From the epibasal cell, the capsule is formed,
while from the hypobasal cell, the foot and seta of the sporophyte

346 APSCHE : SEMESTER-I BOTANY TEXT BOOK


are formed. Further irregular divisions take place, and a globular

parenchymatous tissue that develops in the food and seta. The lower
cells of this tissue ultimately form an anchor-shaped or bulbous
mass of cells that presses into gametophytic tissue for anchoring
and absorption of food and water from it. The upper cells of the
tissue form the seta. In which the cells are isodiametric and remain
short until spore tetrad formation. Then by many transverse
divisions take place, and cells elongate rapidly to increase the seta
length. This ruptures the calyptra and pushes the capsule out.

Fig 5.2.7: a).Marchantiaarchegoniophore with sporophyte and


b). Sporophyte structure
.Source: https://cutt.ly/QQ9AHL4
Source: https://cutt.ly/WQ9A1p4

amphithecium
endothecium. The amphithecium by anticlinal divisions
forms a single-layered jacket of the capsule. Endothecium cells are

sporogenous tissue. About half of the sporogenous tissue cells


divide transversely, forming vertical rows of spore mother cells.
Spore mother cell will divide meiotically to form 4 haploid
tetrahedral spores. The remaining sporogenous cells become sterile
and form long slender cells with tapering ends. The protoplasm of

APSCHE : SEMESTER-I BOTANY TEXT BOOK 347


this will partly disappear and form inner spiral thickenings. These
spindle-shaped, long, slender, spirally thickened cells are
(Fig.5.2.7 a,b.). These are hygroscopic and perform
squirming movements to help in loosening the spore mass and for
dispersal.
The mature sporogonium is differentiated into foot, seta,
and capsule. The foot is a bulbous structure anchoring into
archegoniophore and absorbs water and nutrient from the
gametophyte. The immature foot cells, seta cells, capsule wall cells
and elaters contain active chloroplasts, so they are not fully
dependent on the gametophyte for their food material. At maturity,
the seta elongates and rupturing the calyptra, the pseudoperianth.
The capsule gets dry and splits irregularly into 6-8 valves along with
the apical cap and assisted by the elaters spores are released and

Germination of the spore :


gametophyte generation. It is a haploid and spherical structure. The
spore wall is differentiated into exospore and inner endospore. The
spore germinates immediately under favourable conditions. The
spores are viable for one year. Two spores of the tetrad will develop
into a male gametophyte, and the other two develop into a female
gametophyte. By the dehiscence of the exospore, the endospore

two lateral cutting faces. Later the apical cell itself giving rise to the
group of meristematic cells at the tips, and the fully grown thallus

Life cycle : Marchantia diplohaplontic


distinct vegetative phases;a green thalloid plant and a sporogonium.
The former is a conspicuous, prominent and independent
gametophyte. This bears the sex organs on localized receptacles.

348 APSCHE : SEMESTER-I BOTANY TEXT BOOK


As Marchantia is dioecious, male and female organs grow on separate
thalli. These structures are haplophase (n), and by the fertilization,
the diploid (2n) sporophyte phase starts. In this phase, zygote,
sporogonium with the foot, seta, and capsule are present. By the
meiosis of the spore mother cells, the future gametophyte phase
will start. Thus, the gametophyte and sporophyte phases of
Marchantia are alternating with each other. (Fig: 5.2.8).

Do you know: In M. polymorpha, nearly 3,00,000 spores are


liberated from a single capsule. The unequal expansion of the
cellulose walls and the spiral thickenings are responsible for
hygroscopic movements of elaters.

Fig 5.2.8: Marchantia life cycle


Source: https://commons.wikimedia.org/wiki/File:Marchantia_Life_Cycle.pdf

APSCHE : SEMESTER-I BOTANY TEXT BOOK 349


5.2.2. Funaria Class : Bryopsida
Sub Class : Bryidae
Order : Funariales
Family : Funariaceae

Occurrence : is a very common moss, widely distributed.

is cosmopolitan and grows in moist, shady and,


recently burnt damp soils as it will have more nitrites besides other
elements. It also occurs on rocks, crevices and, on the slopes of the
mountain ranges during the rainy seasons. It is mostly grown in
tropical and temperate conditions They grow as dense green patches
as mat-like structures. The name is derived from the Latin word
“Funis”, (=a rope).

FACTS: Funariais also called cord moss or leaf moss,as the twisted
seta is very hygroscopic and untwists when moist.

The Gametophyte : The gametophyte shows two distinct phases;


1) prostrate, filamentous algae-like structure called protonema.
2) upright-leafy persistent moss plant the gametophore. The adult
gametophore has a small, slender, erect, radial stem covered with
simple small leaves. The stem shows monopodial branching, and
lateral branches are extra axillary in development. At the base of
the stem, there are numerous branched, slender, colourless,
multicellular rhizoids with oblique septa. At maturity, these rhizoids
are brown or red. Their function is similar to the roots of the higher
plants in holding and absorption of water and minerals from the

Leaves are spirally arranged on the stem. In the early stage,


leaves are arranged in three vertical rows corresponding to the three
cutting faces of the apical cell in their formation. However, at

350 APSCHE : SEMESTER-I BOTANY TEXT BOOK


maturity, they will be displaced and arranged spirally with 3/8
divergence. In contrast, the lower leaves are smaller and scattered,
while the upper leaves are larger and crowded. Leaves are sessile,
ovate, with pointed apex and entire margin. Each leaf has a distinct
midrib and is inserted on the stem with an abroad base. (Fig: 5.2.9
a, b & c).

Know more: The stem is called as cauloid and the leaves phylloids.

Fig 5.2.9 Funaria gametophyte plants


Source: https://cutt.ly/KWxS7xc
Source: https://cutt.ly/kWxDqEf
Source: https://cutt.ly/OWxDrzi

Growth : Growth takes place by the activity of a single pyramidal


(Tetrahedral) apical cell located at the tip of the stem. Each segment
divides by a periclinal wall into inner and outer cells. The inner
segment will develop the inner tissue of the stem. At the same time,
the outer segments give rise to three leaves. In leaves, the apical
cell is with two cutting faces and cuts off segments to the right and
left alternately.

Internal structures : Though it is gametophyte in nature, the moss


stem has internal differentiation comparable to that of the stem of
the sporophyte of vascular plants. Stemshows three distinct zones

APSCHE : SEMESTER-I BOTANY TEXT BOOK 351



       
     
       




 
  

   
  
      
  
   
  
  


       
 
       
  
      


352 APSCHE : SEMESTER-I BOTANY TEXT BOOK
1) Epidermis: It consists of a single layer of cells external to the
cortex. Cells are small and possess chloroplast in the early stage.
On maturity, the wall becomes thick and devoid of chloroplasts.
True stomata are absent.

2) Cortex: It is made up of undifferentiated large, thin-walled


parenchymatous cells. Young stem possesses chloroplast, and on
maturity, they lack them. The fully mature cortex usually grades
off from thick-walled cells to thin-walled cells next to a central
cylinder. In the periphery of the cortex, small isolated patches of
cells constituting the leaf traces but do not join the central cylinder.
Cortex cells store the food.
3) Central cylinder: It forms the core of the stem consists of long
narrow, thin-walled, colourless cells lacking protoplasm with
nonlignified walls commonly called hydroids. Which are functional
counterparts of the tracheids. This is useful for the conduction and
storage of food.
The leaf has several cells thick midrib region and single-
layered wings on either side of the midrib. The midrib has central
thick-walled cells surrounded by thin-walled cells; the wings have

Fig 5.2.10 a. Cross section of the Moss stem b. cross section of the leaf
Source: https://cutt.ly/qWxDfEz
Source: https://cutt.ly/1WxDj5w

APSCHE : SEMESTER-I BOTANY TEXT BOOK 353




    
  



      
   
       
      


 


       

  
 
   

       



354 APSCHE : SEMESTER-I BOTANY TEXT BOOK


polygonal parenchymatous cells with numerous chloroplasts (Fig:
5.2.10 a, b).
Reproduction :
methods.
Vegetative propagation: Thallus vegetatively reproduces in
favourable habitats in the growing season by several methods.
A) Progressive growth and death of the gametophores (which is a
characteristic feature of Hepaticae, is present in prostrate rhizome
bearing erect branches) after the death of prostrate parts, the erect

B) When growing in light, rhizoids of the leafy shoots may become


protonemal stage and produce buds similar to the buds on the
protonema. These are called resting buds. Secondary protonema
may also develop from wounded parts of the leaf or stems.
C)From the terminal cells of the protonema, developed small bud-
like structures are called gemmae. They are thin-walled and possess
chloroplasts. The detached gemmae germinate to form the
protonema. The gemmae also develop in unfavourable conditions
or during the injury on the stem and leaves.
D) Wounding of the sporophytic tissue into green protonemal
filaments can develop the leafy gametophores. The production of
the leafy gametophores directly from the vegetative cells of the
sporangium without the inter vention of the spores is
apospory

Find: Identify the difference between apospory gametophores


and normal gametophores.

Sexual reproduction : sex organs are borne terminally


in clusters on leafy gametophores as homothallic, in which the
antheridia and archegonia are born on the same plant (monoecious),

APSCHE : SEMESTER-I BOTANY TEXT BOOK 355


  
       



n
n

* * * * * *

on separate branches (autoicous moss). Thus the development limits


the growth of the axis. The antheridia are produced at the tip of the
main stem and develop first (protandry), ensuring the cross
fertilization. Later the archegonia develop on the lateral branch. A
cooler temperature is needed for the maturation of the sex organs.
Antheridia :
branch. The vegetative leaves at the top are crowded together to
form a rosette of leaves called perigonial leaves. Thus, appear to be
a flower. At the convex tip, large numbers of antheridia are formed
without any order. They are projected from the surface of the
receptacle and at different stages of development. Numerous green,
sterile, hair-like filaments are associated with antheridia. They are
multicellular and upright with capitates heads and are known as
paraphyses. Each has a single row of 4-6 cells with chloroplasts. As
the top cells are meeting over the antheridia, they afford protection.
They help to conserve moisture as capillarity and secrete mucilage
in protecting from the antheridia due to water loss and also in the
efficient release of the sperms.

356 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Mature antheridium :
colour body on a short massive stalk. The body has a single-layered
outer jacket of polyhedral f lattened cells containing many
chloroplasts during an early stage, and later they turn an orange
colour. The apex of the jacket is closed by one or two large hyaline
opercular cells with thicker walls. Within the jacket, a dense central
mass of androcyte mother cells is present. Each androcyte mother
cell divides into two androcytes. Metamorphosis of androcytes into
biflagellate antherozoids takes place. At maturity, the sperm is an
elongated, spirally coiled, anterior biflagellate structure (Fig: 5.2.11
a, b). Dehiscence of antheridia takes place by absorbing
considerable water. By intake of water, the jacket cells become
mucilaginous and swell. The antheridial contents rupture the
operculum and force out the mass of antheridia. The presence of
the lipid material in the matrix possessing the sperms mass permits
the rapid spread of sperms as a thin film into the surroundings.

Fig .5.2.11 Funaria: Antheridium

Archegonia: Archegonia are present on the apex of the lateral


branch, surrounded by the vegetative leaves as a cluster. These leaves
are known as perichaetial leaves. Paraphyses are present
intermingled with archegonia, similar to that of antheridia. This
branch is convex in shape, looks like a regular branch (Fig: 5.2.12
a,b).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 357


Fig. 5.2.12. a. b. Funaria: Archegoniaum

Mature Archegonium :
with venter and neck parts. It has a long massive multicellular stalk.
Venter has double layered sterile cells enclosing venter cavity; in
the cavity, an egg cell and above it a ventral canal cell is present. A
long twisted neck has six rows of neck cells enclosing an axial row
of ten or more neck canal cells. At the tip of the neck, four cap cells
are present. In the mature archegonia, the axial row of neck canal
cells and ventral canal cells degenerate to produce slimy fluid. With
the absorption of water by this, the terminal cells of the neck are

Fertilization :
Funaria.
The water possessing sperms may trickle down from antheridia
branch to a lower archegonia branch. The perigonial leaves form a
cup-like structure, the splashing of rainwater on this may disperse
the thin film of sperms. Air current also adds to the splash. Small
insects like mites, which are attracted to the mucilaginous material,
may behave as agents in sperms dispersal. After reaching the
archegonia neck, the sugar substance (emitted by the disintegration

358 APSCHE : SEMESTER-I BOTANY TEXT BOOK


of the neck canal cells and venter canal cell) guiding sperms to reach
the egg in the venter. Only one sperm will fuses with the egg in the
venter. The resultant zygote develops into a sporophyte.

Sporophyte :
and is the first cell of the sporophyte, which is useful for the
formation of meiospores and effective dispersal. It increases in size
and, by active segmentation, develops two active apical growing
points at opposite ends (Biapical). Mass of cells will be produced by
the embryo within the venter, and it will become a long slender
structure. This is also accompanied by the growth in the venter for
protective covering around the developing embryo. The enlarged
venter is called calyptra.

stalk of the archegonia, and to the female branch, forms the foot.
The upper part of the embryo undergoes a well-developed capsule.
The middle part of the embryo elongates into seta. The continuous
growth and elongation of seta rupture the calyptra at the base, and
it will be carried aloft as a hood on the developing capsule (Fig:
5.2.13 a, b). Funaria
at the distal end of the female branch. Gradually the lateral female
branch replaces the main branch. The sporangium is differentiated
into foot, seta and capsule.

tissue of the female branch and absorbs food and water for the
growing sporophyte. Seta is a long slender, tough stalk-like structure
bearing the capsule at its top end. The seta has a simple structure,
with more tissue differentiation as central strand, cortex and
epidermis. The central strand is a primitive conducting system. The
thick-walled cortex gives strength to the seta. The capsule is a highly
organized structure. It is pear-shaped and green in colour. Its upper

APSCHE : SEMESTER-I BOTANY TEXT BOOK 359


part is covered by a hood called calyptra. This has three well-marked
regions called- i. apophysis, ii. theca, and iii. operculum (Fig:
5.2.14).

Fig. 5.2.13 a.b. Funaria: Sporophyte on the gametophyte.

i.Apophysis: The extended region of the seta into the basal region
of the capsule to form the local assimilatory zone is called the
apophysis. In the centre, it has a conducting tissue that is continuous
with a central strand of seta. Surrounding this, spongy green tissue
with intercellular spaces is seen. They are with abundant
chloroplasts. The epidermis in this region has true stomata with
the guard cells in the initial stages; later, it develops as a single
annular guard cell with two nuclei.

ii. Theca: It is the fertile region of the sporophyte. In the centre of


this is a sterile column of thin-walled parenchymatous cells known
as the columella. The upper portion of this is cone-shaped and
extends upwards into the concave inner portion of the operculum.
At the base is connected to the apophysis. The columella is
surrounded by a barrel-shaped spore sac, in the longitudinal section

360 APSCHE : SEMESTER-I BOTANY TEXT BOOK


Fig: 5.2.14 a. Funaria capsule and b. Funaia capsule peristoem opening
Source: https://cutt.ly/lWxDoBr
Source: https://cutt.ly/qWxDg8S

layer of inner wall layer without chloroplasts. In between the walls


of the spore sac, spore mother cells are present. The spore mother
cell undergoes meiotic division and produces the spore tetrad.
Outside the spore sac, there is cylindrical air space, traversed by
green filamentous cells connecting the capsule wall. The capsule
wall in this region has a well-defined epidermis. Below, the epidermis
consists of 2-3 layers of compact, parenchymatous cells forming
hypodermis. The inner layers of this are composed of loosely
arranged cells with chloroplasts. Near the fertile region, these layers
are thinned. Whereas near the inner tissue, they are green, thick,

iii. Operculum: The terminal, obliquely placed conical cap-like


portion is operculum. The upper region of the capsule has a highly
modified spore dispersal mechanism. This operculum and the
peristome are marked off by a constriction from theca. Below this,
there is a diaphragm (Rim) composed of 2-3 layers of radiately
elongated, pitted cells. The rim has the form of a circular ledge

APSCHE : SEMESTER-I BOTANY TEXT BOOK 361


perforated by the thin-walled tissue in continuation with the
columella. The rim stretches inwards from the epidermis of the
capsule wall and joins the peristome. When the operculum separates

superimposed layers of epidermal cells that occupy the periphery


of the operculum. The two lower layers of the annulus cells have
thinner walls and are useful in dehiscence. Below the edge of the
diaphragm, two rows of the curved, narrow, triangular plate-like
teeth, sixteen teeth in each row, are present as peristome. Both the
sets are on the same radii and thus opposite to each other. The
peristome teeth close the opening of the spore sac. These teeth are
sculptured. The outer set is hygroscopic and responds to the
environment. By the destruction of thin-walled annulus cells, the
operculum is loosened and later dropped off. In Funaria, the
archesporium or fertile tissue is less when compared to other moss
plants as it has more cells in columella, operculum, peristome,
apophysis, and multi-layered capsule wall.
Find:

Activity: with the photosynthetic cells and the sporangium this


may resemble the Pteridophyte capsule – Justify with reason

Dehiscence of capsule : On maturity of the capsule, the thin walled


operculum and annulus proper cells shrink and shrivel. As a result,
loosened from the underlying tissue, it drops off, exposing the
peristome. This is also assisted by the hygroscopic cells of the
peristome teeth. By this time, columella and other adjoining tissue

Dispersal of spores :
movements assist in a gradual release of the spores. Slight rotation

362 APSCHE : SEMESTER-I BOTANY TEXT BOOK


of the teeth with changes in the moisture content of the air can
permit spores to shift out gradually. The teeth bend inward and
outward with a change in the humidity. In wet weather, spores are
prevented from escaping; in dry weather, the peristome will lose
the water, and the slit opens out to release the spores.
Germination of the spore :
falling on the moist soil and under the suitable condition of light
and temperature. By imbibition of water through the aperture
region of the spore, germination talks place. Initially, swelling at
the unthickened part of the intine ruptures the exine. The disc-
shaped thickened intine protrudes. The tubular outgrowth takes
place and develops rhizoids, and other green plastid cells hence
formed the protonema. Gradually this forms the net-like prostrate
and erect branches of green chloronemal branches. The nongreen
rhizoids will penetrate soil and function as absorbing and anchoring
parts. The chloronemal stage will continue for many days. Few apical
cells produce another type of filaments called caulonema, on which
buds will develop and produce the leafy gametophores. They grow
rapidly along the surface of the substratum by regular branching.
The bud arises as a lateral protrusion from a chloronemal branch
cell just below a septum. By the activity of its apical cell, the bud
grows into a leafy gametophyte (Fig: 5.2.15).
Life cycle :
haploid leafy moss plant, the other diploid leafless sporogonium,
this is partially dependent on the leafy gametophyte for nutrition.
Haploid gametophyte plants produce the gametes in the antheridia
and archegonia, and the diploid sporophyte produces haploid spores
after meiosis division. Gametophyte represents the sexual
generation and alternates with the sporophyte, an asexual
generation. This phenomenon is called the alternation of
generations (Fig: 5.2.16).

APSCHE : SEMESTER-I BOTANY TEXT BOOK 363


Fig. 5.2.15 a. Funaria spore germination
Fig. 5.2.15 b. FumariaProtonema
and protonema
Source: https://cutt.ly/sWxDu7o
Source: https://cutt.ly/VWxDs6a

Fig.5.2.16. Funaria Life cycle


Source: https://cutt.ly/5WvIU3I

364 APSCHE : SEMESTER-I BOTANY TEXT BOOK


General account on evoluation of sporophyte in
Chapter -5.3 Bryophyte

Learning Outcomes: At the end of the chapter you should be


able to:
1. Describe the evolutionary trends in shifting of Sporophytic
phase from complete parasite to partial parasite.
2. Explain the progressive and retrogressive theories on
evolution of sporophyte bryophyte

5.3.1: Evolution of sporophyte in Bryophyte

developed from the diploid zygote and represents asexual

gametophyte. They produce meiotic spores, which on germination,


develop the haploid gametophyte. Thus the bryophytes express the
alternation of generation in their life cycle. In form, it varies from
Riccia
object differentiated into foot, seta and capsule.

evolution of the sporophyte -


1. Theory of progressive sterilization of sporogenous tissue
2. Theory of progressive simplification or Reduction theory
Theory of progressive sterilization of sporogenous tissue:

of sporophyte plants. This is also known as interpolation or


intercalation theory, as the gametophyte generation is original while
the sporophyte generation is an entirely new phase derived from
the progressive elaboration of the zygote into the life cycle. Bower
opined that simple sporophyte to more complex sporophytes
evolved by progressive sterilization of the potentially sporogenous

APSCHE : SEMESTER-I BOTANY TEXT BOOK 365



     



       



       

       

       

    

   

        

 
    
   



366 APSCHE : SEMESTER-I BOTANY TEXT BOOK


tissue instead of forming spores; they remain sterile. These sterile
cells take up the function of nutrition, dehiscence and dispersal.
Cavers, Campbell, Strasburger and Smith also supported this theory.

Bryophyte sporophyte did not make its appearance until the


development of archegonium. The zygote in the archegonium did
not divide meiotically but divided mitotically to develop diploid cells.
Then they divide meiotically to form spores. All the cells were
sporogenous. The sporophyte evolved with some superficial cells
as sterile and only a few cells in the formation of the sporogenous.
Further evolution of the sporophyte came about as a result of
additional sterilization of sporogenous tissue, leading to the
differentiation of sporophyte with the foot, seta and capsule
development. There was a progression from a simple to a more
complex sporophyte by progressive sterilization of sporogenous
Riccia,
Funaria.

the zygote, after fertilization, increases in size and


undergoes a single division forming a two-celled embryo. By
repeated divisions forms undifferentiated cells, later on, get
differentiated into sterile outer cells of capsule wall and an inner
mass of fertile sporogenous cells. It has a spherical capsule without
a seta and foot. This is totally dependent on the gametophyte, and
until the disintegration of the gametophyte thallus, spores will not
be liberated. There will not be any special features in the sporophyte
R.crystallina
nutritive cells. These sterile cells may be the forerunners of the
of marchantiales, the inner
endothecium cells differentiate into sporocytes and elaters like
Sphaerocarpos, the sporophyte has a sterile

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      
       
        




  

    
      

* * * * * * *

bulbous foot and a narrow seta along with some sterile nurse cells
developed by endothecium.

the sporophyte is differentiated into bulbous


foot, massive seta and a capsule with half of the endothecium giving
rise to fertile sporogenous tissue and remaining sterile elaters with
spiral thickenings. These are hygroscopic in nature and useful in
spore dispersal. This is the beginning of the spore dispersal

Marchantia, the sporophyte is much evolved, showing the


differentiation of foot, seta and capsule. The foot is embedded into
the gametophyte tissue absorbing the nutrient material. On
maturation, the short seta in the initial stage will elongate on
maturity to facilitate the spore dispersal.

368 APSCHE : SEMESTER-I BOTANY TEXT BOOK


In P of Jungermanniales, due to the further
development, the jacket has two or more layered. A bunch of
elatophores are present at the proximal end at the base of the
capsule. These are developed from sporogenous cells with spiral
thickening. They occupy more than half of the sporophyte. The
remaining sporogenous tissue will produce spore mother cells and
elaters. Ultimately, only 10% of the sporophyte will be sporogenous
tissue. At maturity, the capsule wall is split into four valves and
spores are liberated.

Anthoceros,
of four cells each. From the lower tier foot and from the upper part,
the capsule is developed. In the capsule, the inner endothecium is
sterile and forms central columella. Surrounding this sheet of
sporogenous tissue is present. The archesporium is much reduced
and produces spore mother cells and pseudo elaters. The covering
layers and the archesporium are derived from the amphithecium.
The jacket layers will have chloroplasts and stomata, showing a
photosynthetic mechanism. Further, with the presence of the
intercalary meristem at the base of the capsule, the continuous

Funaria, the capsule is developed from epibasal cell and


foot and seta from hypobasal cell. The epibasalcell divide and
produce outer amphithecium and inner endothecium.
Amphithecium differentiates into the epidermis and photosynthetic
tissue, outer spore sac of the capsule. At the apex, it produces the
operculum, the inner parts of the peristome. The outermost layer
of the endothecium forms the archesporium of one cell thickness
of fertile cells. There are large air spaces traversed by delicate
Funaria, the sporophyte is highly evolved

APSCHE : SEMESTER-I BOTANY TEXT BOOK 369


among bryophytes as it is partially dependent on gametophyte only
for water and minerals. Spore dispersal in this is unique with

Thus, Bower’s theory of sterilization is well supported from


the simple structure of sporophyte to the well-differentiated
structure of capsule (Fig: 5.3.1).

Fig 5.3.1 EVOLUTION OF SPOROPHYTE

Theory of progressive simplification of Reduction theory

and others. This theory holds that evolution is in a downwards


direction- retrogressive evolution. The reduction or simplification
Riccia

From the moss plants, the sterile parts of the sporophyte have
Riccia, in
which only the fertile cells are present. The descending evolution

370 APSCHE : SEMESTER-I BOTANY TEXT BOOK


• The leaves of the erect sporophyte were lost due to intense
insolation and desiccation.
• The erect leafy sporophyte becomes attached to
gametophyte.
• The air spaces in the photosynthetic system disappeared due
to more dependence on gametophyte.
• The functionless stomata in the Sphagnum disappeared in
marchantiales.
• Simplification in the dehiscence mechanism
• Decrease in the capsule wall, and elimination of the foot and
seta.
All these led to the progressive increase in the fertility of the
sporogenous cells.
Schofield, W. B., (2001)Introduction
Video source: to Bryology, The Blackburn Press,
h t tp s : //w w w.yo utub e .c o m/ Caldwell, New Jersey
watch?v=1UKaOzYZeTc
h t tp s : //w w w.yo utub e .c o m/ Self-Analysis:
watch?v=ZK2V8lOUlsM
h t tp s : //w w w.yo utub e .c o m/ 1. Observe the presence of either
watch?v=_uAaw5By5ro thalloid or Moss bryophytes in
the field. Take cross section of
the specimen, record the
References : findings.
Jonathan Shaw, A., (2010) 2. Analyze the reason for the
Bryophyte Biology, Cambridge diecious state of the
Marchantiathallus.
University Press, London
3. Why inversion of the
Parihar, N.S., (2019) An archegonia has taken place
Introduction to Embryophyta after fertilization. Discuss.
:Bryophyta Vol. I, Surjeet 4. Give the reason for the
Publications, Delhi development of the three-level
protection for the capsule.
Vashishta, B. R., A K Sinha and
5. What are the differences
Adarsh Kumar (2019) Botany for
among Marchantia and
Degree Students –Bryophyta, S.
Funaria in the following
Chand Publishing, New Delhi
points

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 Thallus differentiation (b) antherozoids
 The organization in the sex (c)
organs (d) Conidia
7. The basal swollen portion of
 Spore dispersal mechanism
the archegonium is:
 Spore germination process.
(a)
Learning tips:
(b)
 Identify and label the (c)
figures with reference to (d) Oospere
the text. 8. Meristematic tissues are
Self-Assessment present in:
(a)
1. Which plant does belong to (b)
(c)
Hepaticopsida?
(d) Anthoceros
(a)
9. Protonema is found in:
(b) Polytrichum
(c) (a)
(d) Marchantia (b)
(c)
2. Which plant does belong to
(d) Anthoceros
Anthoceropsida?
10. The structure not involved
(a)
in asexual reproduction is:
(b) Polytrichum
(c) (a)
(d) Anthoceros (b)
(c)
3. Which plant is a moss?
(d) (d)Bract
(a)
11. A structure is present in the
(b)
(c) centre of the capsule called:
(d) Anthoceros (a)
4. Which of the followings is (b)
(c)
absent in bryophytes?
(d) pseudoelaters
(a) Archegonia
12.
(b) Oosphere
(c) Zoospore the columella. This region
(d) Antheridia contains spores and:
5. Androcytes give rise to: (a)
(a) (b)
(b) (c)
(c) (d) pseudoelaters
(d) Sporocyte 13. Endothecium divides to
6. Antherozoid mother cells form:
are: (a)
(a) (b)
(c)

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(d) pseudoeaters Glossary
14. Acrocarpous: relative to a moss,
form:
having sporophyte produced at apex of
(a)
(b) spores stem or main branch.
(c) Acropetal: Development of organs
(d) pseudoelaters successively towards the apex. The
15. The leaves adjacent to the oldest is at the base and the youngest
sex organs are at the apex.
(a)
Acrogynous: In many leafy liverworts,
(b)
(c) sporophyte growing at top of stem
(d) (lemma (from apical cell).
16. The superficial cells of the Androcyte: Antherozoid mother cell,
stem give rise to: the cell later develop in to antherozoid
(a) Antheridium: The male sex organ of
(b)
(c) involucre the cryptogams
(d) Gemma Archegonium: The female sex organ
17. The archegonia and bracts in Br yophyta, pteridophyte and
forms structure called: g ymnosperms, containing the egg
(a) inside a cellular jacket.
(b) paraphylls
(c) involucre Calyptra: Covering developed from
(d) Gemma the venter of the archegonium in
18. Pseudofoot is present in: bryophytes and pteridophytes, which
(a) Funaria, surrounds the young sporophyte.
(b)
Capsule: Part of the sporangium
(c)
(d) Anthoceros containing the spores
19. The hypobasal cell does not Caulonema: second stage of the
divide further and forms: protonema (chloronema is the first
(a) stage),stage giving rise to buds.
(b) Chersophilous:growing on poor and dry
(c)
(d) haustorium habitats
20. Rhizome is present in Columella: sterile column located in
(a) the center of the capsule
(b) Potytrichum Cosmopolitan: present in almost all
(c)
(d) Anthoceros parts of the world.
Deoperculate: relative to a capsule,
lacking an operculum, the lid has fallen.

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Dioecious: Having unisexual male Hydroid : conductive cell, elongated and
and female sexual reproductive with thin and concave walls, pre-sent in
organs bor ne on different the centra strand .
individuals Hypophysis: referring to a capsule :
Distichous: leaves arranged in two sterile neck located at the base of the
opposite rows on the stem. capsule, between seta and urn.
Egg: A female gamete Leptoid: food conductive cell with thin
Elater : in liverworts, specialized walls and large lumen in some mosses.
(elongated and with spiral Perichaetialleaves:specialized leaves
thickenings) cell present among surrounding archegonia.
spores in the capsule.

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APSCHE : SEMESTER-I BOTANY TEXT BOOK 375
376 APSCHE : SEMESTER-I BOTANY TEXT BOOK

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