Agriculture DLP Sample - 2018 PDF

Distance Learning Course
Sample Chapters
AGRICULTURE
Table of contents
A note about this sample content booklet .................................................... 3
How to subscribe for Evolution Distance Learning Programme ...................... 4
Sample Chapters ......................................................................................... 7
Chapter 1 [From Farm Management] Types and Systems of Farming ............................ 8
Chapter 2 [Weed Management] Weed Management..................................................13
Chapter 3 [Soil Science] Soil Forming Processes..........................................................23
Chapter 4 [From Plant Breeding] Use of Polyploidy in Plant Breeding ..........................30
Chapter 5 [From Entomology] Storage Pests & their Management ..............................34
Chapter 6 [From Horticulture] Propagation of Horticultural Crops ...............................46
Chapter 7 [From Cell Biology] Chromosome ...............................................................62
Chapter 8 [From Plant Physiology] Plant nutrition, absorption, translocation and
metabolism of nutrients ...........................................................................................81
Distance Learning Course Enrolment form .................................................. 91
Sample Chapters: AGRICULTURE DISTANCE LEARNING COURSE |Page 2

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booklet
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Sample Chapters

Chapter 1 [From Farm Management]
Types and Systems of Farming
The combinations of products on a given farm and the methods or practices that are used
in the production of these products based on the ways of economic and social functioning
in farming are known as system of farming. It is concerned with the organizational setup
under which the farm is being run. Systems mainly deal with questions like who is the
owner of the land, whether the resources are pooled or used individually, and who makes
the managerial decisions. It is classified into five groups, namely, peasant, co-operative,
state, capitalistic and collective farming.
Whereas, when farms in a group are quite similar in the kinds and productions of the
crops and livestock that are produced and the methods and practices followed in
production, the group is described as type of farming. It includes specialized, diversified,
mixed and ranching.
I. Types of Farming
A) Specialized Farming: Specialization means that the farmers specialize in one

enterprise such as crop, dairying, poultry or tea estate etc. The major enterprise
contributes more than 50% of the total farm income. Examples are sugarcane farm,
cotton farm, poultry farm, dairy farm, wheat farm etc.
Advantages:
i. Better use of land: It is more profitable to grow a crop on a land best suited to it.
E.g. Jute cultivation on a swampy land
ii. Better Marketing: Specialization allows for better assembling, grading, processing,
storing, transporting and financing of the produce.
iii. Better Management: As there are fewer enterprises, wastage can easily be detected
and they can be better managed.
iv. Less equipment and labour are needed.
v. Costly and efficient machinery can be used.
vi. Efficiency and skills are increased: Specialization allows a man to be more
efficient and expert at doing a few things.
Disadvantages:
i. There is a greater risk - failure of crop and market together may ruin the farmer.
ii. Productive resources– land, labour and capital are not fully utilized.
iii. Fertility of soil cannot be properly maintained due to lack of suitable rotations.

iv. By products of the farm cannot be fully utilized due to lack of sufficient livestock
on the farm.
v. Farm returns in cash are not generally received more than once in a year, i.e.,
there is no regular farm return.
B) Diversified Farming: A Farm on which no single or source of income yields 50 per cent
of the total receipt is called a diversified farm. More enterprises are taken up on the farm
and no single enterprise is relatively much more important.
 Advantages:
i. Better use of land, labour and capital: Better use of land through adoption
of crop rotations, steady employment generation and more efficient use of
equipment are obtained.
ii. Business risk is reduced due to a crop failure or unfavourable market

prices.
iii. Regular and quicker returns are obtained from various enterprises.
 Disadvantages:
i. Because of varied jobs in diversified farming, a farmer can efficiently

supervise only limited number of workers.
ii. Better equipping of the farm is not possible because it is not economical to
have extensive implements and machinery for each enterprise.
C) Mixed Farming: Mixed farming is a type of farming under which crop production is
combined with livestock rearing. The livestock enterprise is complementary to crop
production programme so as to provide a balanced and productive system of farming. In
mixed farming, the contribution of livestock activities to gross farm income should be a
minimum of 10 per cent and a maximum of 49 per cent.
 Advantage:
i. Mixed farming helps in the maintenance of soil fertility. Byproducts of crop

production, namely, green and dry fodder, rice bran, gram husk and so on are
better used for live stock. On the other hand, farm yard manure enriches the
soil with important nutrients.
ii. Draught animals are used in crop production and transport.
iii. It provides regular income and employment.
D) Ranching
A ranch differs from other type of crop and livestock farming in that the livestock graze
the natural vegetation. Ranch land is not utilized by tilling or raising crops. The ranchers

have no land of their own and make use of the public grazing land. Ranching is followed
in Australia, America, Tibet and certain parts of India (Bikaner in Rajasthan).
Determinants of Type of Farming

1) Physical Factors: The physical factors namely climate, soil and topography have
influence in determining the type of farming.
2) Economic Factors: Marketing cost, changes in input and output prices,

availability of resources like land, labour and capital, competition between
enterprises, miscellaneous factors such as personal likes and dislikes; prevalence
of pest and diseases, etc. influence type of farming.
3) Social Factors: The kind of people in the community and the provision of
protection of crops against the hazards of bird and animal ravages may influence
the farming community to change the pattern of cropping. The co-operative spirit
in providing security to crops, benefits resulting from low transport costs through
collective sale and better marketing facilities permit farmers to expand some
enterprises like fruit farming, dairying or poultry rearing.
System of Farming
The system of farming refers to the organizational set up under which farm is being run. It
involves questions like who is the owner of land, whether resources are used jointly or
individually and who makes managerial decisions. Systems of farming, which are based
on different organizational set up, may be classified into five broad categories:
1. Capitalist or Estate farming: In what is known as capitalistic or estate or

corporate farming, land is held in large areas by private capitalists, corporations
or syndicates. Capital is supplied by one or a few persons or by many, in which
case it runs like a joint stock company. In such farms, the unit of organization is
large and the work is carried on with hired labour; latest technical knowhow is
used and extensive use of machines is made and hence they are efficient.
Examples of this type of farming are frequently found in USA, Australia, Canada
and few in India too. Such types of farms have been organized in the states of
Maharashtra, Chennai and Karnataka for the plantation of coffee, tea and rubber
and sugarcane.
The advantages of such farming are good supervision, strong organizational set up,
sufficient resources etc. Their weaknesses are that it creates socio-economic
imbalances and the actual cultivator is not the owner of the farm.
2. State farming: State farming as the name indicates is managed by the

government. Here land is owned by the state. The operation and management is
done by government officials. The state performs the function of risk bearing and
decision making, which cultivation is carried on with help of hired labour. All the
labourers are hired on daily or monthly basis and they have no right in deciding
the farm policy.

Such farms are not very paying because of lack of incentive. There is no dearth of
resources at such farms but sometimes it so happens that they are not available in
time and utilized fully.
3. Collective farming: The name, collective farming implies the collective

management of land where in large number of families or villagers residing in the
same village pool their resources like land, livestock, and machinery. A general
body having the highest power is formed which manages the farms. The resources
do not belong to any family or farmer but to the society or collective.
Collective farming has come into much prominence and has been adopted by some
countries notably by the Russia and China. The worst thing with this system is
that the individual has no voice. Farming is done generally on large scale and
thereby is mostly mechanized. This system is not prevalent in our country.
4. Peasant farming: This system of farming refers to the type of organization in

which an individual cultivator is the owner, manager and organizer of the farm.
The peasant farming, on the one hand, places a greater importance on
management and the use of family labour which maximizes farm business income
and on the other hand allows the organization of the farm to be adjusted to the
capabilities of the various members of the family.
The biggest advantage of this system is that the farmers himself is the owner and
therefore free to take all types of decisions. A general weakness of this system is
that the resources with the individual are less. Another difficulty is because of the
law of inheritance. An individual holding goes on reducing as all the members in
the family have equal rights in that land.
5. Co-operative farming: Co-operative farming is a voluntary organization in

which small farmers and landless labourers increase their income by pooling land
resources. According to planning commission, Co-operative farming necessarily
implies pooling of land and joint management.
The working group on co-operative farming defines a co-operative farming

society as “a voluntary association of cultivators for better utilization of resources
including manpower and pooled land and in which majority of the members participate in
farm operation with a view to increasing agricultural production, employment and
income.”
A Co-operative farming society makes one of the following four forms:
i. Co-operative better farming: These societies are based on individual

ownership and individual operation. Farmers who have small holdings and
limited resources join to form a society for some specific purpose eg: use of
machinery, sale of product. They are organized with a view to introduce
improved methods of agriculture. Each farmer pays for the services which
he receives from the society. The earnings of the member from piece of
land, after deducting the expenses, his profit.

ii. Co-operative Joint farming: Under this type, the right of individual
ownership is recognized and respected but the small owners’ pool their
land for the purpose of joint cultivation. The ownership is individual but
the operations are collective. The management is democratic and is elected
by the members of the society. Each member working on the farm receives
daily wages for his daily work and profit is distributed according to his
share in land.
iii. Co-operative tenant farming: Such societies are usually organized by
landless farmers. In this system usually land belongs to the society. The
land is divided into plots which are leased out for cultivation to individual
members. The society arranges for agricultural requirements such as
credit, seeds, manures, marketing of the produce etc. Each member is
responsible to the society for the payment s of rent on his plot. He is at
liberty to dispose of his produce in such a manner as he likes.
iv. Co-operative collective farming: Both ownership and operations under
this system are collective. Members do not have any right on land and they
cannot take farming decisions independently but are guided by a supreme
general body. It undertakes joint cultivation for which all members pool
their resources. Profit is distributed according to the labour and capitals
invested by the members.

Chapter 2 [Weed Management]
Weed Management
Weed management is the application of certain principles and suitable methods that will
improve the vigor and uniform stand of the crop. At the same time ignore or discourage
the invasion and growth of weeds.
Weed Control Vs Weeds Management

Weed Control and Weed Management, are the two terms used in weed science. Weed
control is the process of limiting infestation of the weed plant so that crops can be grown
profitably, whereas weed management includes prevention, eradication and control by
regulated use, restricting invasion, suppression of growth, prevention of seed production and
complete destruction. Thus weed control is one of the aspects of weed management.
Principles of Weed Management

Prevention: Prevent the entry and establishment of weeds into uninfected area.
Important weed prevention practices are:
1. Use clean crop seeds/ weed free crop seed

2. Avoid feeding of screenings, grain or hey containing weed seeds to
live stock without destroying their viability by grinding, cooking
and ensiling.
3. Use well rotten\decomposed organic manure. Avoid reaching of
weed seeds into the compost pit
4. Prevent movement of weeds with other farm resources
5. Keep non crop area clean
6. Use vigilance
7. Follow legal & quarantine measures
Eradication: It is complete removal of all live plant parts and seeds of the weed from an area. It
may be a field/farm/village/geographical region depending upon the need.
In general eradication of common weed seeds is not practiced as these weeds harbor crop
pests or secretes soil nematodicides. They may be useful to hold the soil nutrients against
leaching losses during fallow period. However weed eradication is justified against weeds
like Striga, Cuscuta, and Lantana to prevent their dispersal to new areas of useful land and
water bodies. Weed eradication programme should begin when the weed growth is
limited. If the weed occupied large and continuous areas eradication is not economical. It
should be carried out more than one year. It requires intensive initial efforts to destroy all
plant parts and followed by many years of vigilance to prevent the new weed seedlings
from establishing into adult plants.
Control: Weed infestations are reduced but not necessarily eliminated. Weed control methods
includes:

I. Cultural
II. Physical/Mechanical
III. Biological.
IV. Chemical.
Cultural Methods of Weed Control

Principle behind this method is giving competitive advantage to the crop. Cultural methods,
alone cannot control weeds, but help in reducing weed population. They should,
therefore, be used in combination with other methods.
1. Proper crop stand and early seedling vigor: Lack of adequate plant
population is prone to heavy weed infestation, which becomes, difficult to
control later. Therefore practices like a). Selection of most adopted crops
and crop varieties b. Use of high viable seeds c. Pre plant seed and soil
treatment with pesticides, dormancy breaking chemicals and germination
boosters d. Adequate seed rates are very important to obtain proper and
uniform crop stand capable of offering competition to the weeds.
2. Selective crop simulation: In crop weed competition, competitive
advantage is in favor of can be achieved by selective simulation of crop
growth. Vigorous crop plants compete better with weeds as they close the
ground very quickly. Selective simulation can be achieved by a) application
of soil amendments like gypsum or lime may correct the soil conditions in
favour of crop growth) addition of FYM or synthetic soil conditioners to
very light or heavy soils may improve the soil structure and maintaining
better air water relationships and ultimately it improving the crop growth
c) manures and fertilizers application of proper kind in adequate quantities
improve the crop growth. D) Inoculation of crop seeds with suitable
nitrogen fixing and phosphorous solubilizing organisms may help in
selective simulation of some crops. Eg: Legume crop and non-legume
weed. Selective simulation in wide row crops like maize, sugarcane, cotton
can be achieved by foliar application of nutrients.
3. Proper planting method: Any planting method that leaves the soil surface
rough and dry will discourage early growth. Plough planting (minimum
tillage) methods proved to be very useful to reduce early weed growth. In
summer, furrow planting of crops reduce the weed problems. Because in
this method irrigation water restricted initially to the furrow only. In
transplanted crops farmers get opportunity to prepare weed free main
field.
4. Planting time: Peak period of germination of seasonal weeds coincides
with crop plants. So, little earlier or later than normal time of sowing is
beneficial by reducing early crop weed competition. Eg: Using photo
insensitive varieties we can make adjustments with regarding to time of
planting.
5. Crop rotation: Growing of different crops in recurrent succession on the
same land is called as crop rotation. Monocropping favors persistence and
association of some weeds. Crop rotation is effective in controlling of crop

associated and crop bound weeds such as Avena fatua in wheat and Cuscuta
in dodder. Wheat-pea and gram break the Avena in wheat, Lucern - grain
crop rotation control Cuscuta. The obnoxious weeds like Cyperus rotundus can
be controlled effectively by including low land rice in crop rotation.
6. Stale Seedbed: It is the one where one or two flushes of weeds are
destroyed before planting of any crop. This is achieved by soaking a well
prepared field with either irrigation or rain and allowing the weeds to
germinate. These weeds are controlled by using contact herbicides like
paraquat and by mechanical methods then sow the crop. Here the
advantage is the crop is germinated in weed free environment. In this way,
weed seed bank is exhausted.
7. Smother crop / Competitive crop: This crop germinates very quickly and
develop large canopy, capable of efficient photosynthesis within short
period. They possess both surface and deep roots. Competitive crop
smother the ground quickly than non-competitive crop. Eg; Cowpea,
lucern, berseem, millets
8. Growing of intercrops: Inter cropping suppresses weeds better than sole
cropping and thus provides an opportunity to utilize crops themselves as
tools of weed management. Many short duration pulses viz., green gram
and soybean effectively smother weeds without causing reduction in the
yield of main crop.
9. Minimum tillage & Zero tillage: Deep and frequent tillage may be useful
for some reasons but it serves to bring more of dominant weed seeds and
rhizomes to the soil surface, preserve the new weed seeds deep in the soil
for the future.
Zero tillage completely avoids burying of weed seeds and reduces persistence of
annual weeds but it induces vigorous growth of perennial weeds.
10. Summer fallowing: The practice of summer tillage or off-season tillage is
one of the effective cultural methods to check the growth of perennial weed
population in crop cultivation. In the month of April, May and June farmers
expose their lands to sun in order to control many soil born pests,
including weeds. roots, rhizomes and tubers of shallow rooted perennials
like Bermuda grass and nut sedge.
11. Lowering area under bunds: Bunds are made in field for the purpose of
irrigation is ideal places for the rapid growth of weeds. One way of tackle
the problem of weeds on bunds is to level the land well so that less no. of
bunds is needed to irrigate the field.
12. Flooding and drainage: Flooding is worldwide crop husbandry method
controlling weeds in rice fields. It controls terrestrial weeds: To ensure the
effectiveness of flooding the weeds should be submerged sufficiently for a
longer period (i.e. for 2 weeks or more). Excludes O2from environment and
kills the weed. In M.P. deep flooding of fallow land is followed in rainy
season and water is let out after 2-3 months. This practice locally called
Haveli. Drainage is used for controlling aquatic and semi aquatic weeds in
rice fields, channels, canals, and ponds.

Mechanical/Physical Methods of Weed Control
These methods are distinguished into a) mechanical b) manual methods. Physical method
of weed control utilizes manual energy, animal power or fuel to run the implements that
dug out the weeds. These methods are as old as agriculture. The hand hoe first animal
drawn implement invented by Jethro Tull in 1731.This methods include under non
chemical method of weed control. Implements used vary from simple to multiple tractor
drawn implements.
 Advantages
1. These methods are efficient, cheaper, safer, to crop and no harmful effect to crop
and user.
2. Oldest, effective and economical method
3. No special skill is required in adopting physical methods.
 Disadvantages
1. More labour is required, and tire some.
2. Its success depends on its timely operations when the weeds still young
3. Usually operations limited by too wet or too dry conditions
 Mechanical/Cultural Methods
1. Hand weeding: Removal of weeds either manually or by using tools like khurpi or
sickle, when weeds grown upto some extent. It’s effective against annuals and
biennials and controls only upper portion of the perennial. Higher labour is
required and is tire some.
2. Hand hoeing: Hoe has been the most appropriate and widely used weeding tool for
centuries. The weeds are taken out with the help of khurpi or hand hoes. Hoeing
by cutting the crown part gives proper control. Annuals and biennials can be
effectively controlled. Convolvulus arvensis which has shallow root system can be
controlled.
3. Spudding: Hand weeding, hand hoeing added by a sharp edged sickle.
4. Sickling: Sickling is also done by hand with the help of sickle to remove the top
growth of weeds to prevent seed production and to starve the underground parts.
These methods are useful for control of tall growing grasses. Especially sickling is
useful in irrigation channels, drainage channels and where undulating
topography is present.
5. Digging: Digging is useful for patch or spot control of obnoxious / perennial

weeds. Digging is very useful in the case of perennial weeds to remove the
underground propagating parts of weeds from the deeper layer of the soil. They

can be eliminated by digging with crowbar or Pick axe etc. For large areas, it is not
desirable because it is costly and labour oriented
6. Mowing: It is cutting of uniform growth from the entire area up to the ground
level. It is useful more in non-cropped areas than cropped areas. Mowing
improves aesthetic value of an area. It is effective against erect and herbaceous
weeds.
7. Cutting: Cutting is the topping/cutting of the weeds little above ground level. It is
done with help of axes and saws. It is mostly practiced against brushes and trees.
In aquatics under water weed cutters are used.
8. Dredging: This is used to control aquatic weeds growing in shallow ditches. It’s a
mechanical method of pulling of aquatic weeds along with their roots & rhizomes
from the mud.
9. Chaining: Very big & heavy chain is pulled over the bottom of a ditch with tractors
along with embankments of ditch. With rubbing action of chain weeds can be
fragmented & collected by nets and hooks.
10. Burning: It is cheapest method to eliminate the mature unwanted vegetation in

non-cropped areas and range lands. Coagulation of protoplasm occurs with
which plant dies.
11. Flaming: It is the momentary exposure of green weeds to as high as 1000oC from
flame throwers to control in row weeds. Eg. Flaming is used in western countries
for selective weed control in crops like cotton, onion, soybean and fruit orchards.
Dodder is also controlled by flaming in lucern.
12. Searing: Repeated application of flame to above ground parts destroyed the root
system and plant dies.
13. Soil Solarization: It is also called solar soil heating. It is effective against weeds
which are produced from seeds. It doesn’t involve any tillage of the field. It can be
achieved by covering the soil with transparent, very thin plastic sheets of 20-
25mm polyethylene (PE) film during hottest part of summer months for 2-4
weeks. This increases the temperature by 10-12 0 C over the unfilmed control
fields. Then weeds seeds are desiccated which are present at top 5 cm soil depth.
Eg: Phaliris minor, Avena and broad leaved weeds controlled by Solarization.
Whereas Melilotus sp. possess hard seed coat is resistant to Solarization treatment.
14. Chiseling: An implement called chisel (spade like implement with very long
handle) with which weeds & soil can be racked up. Generally, it is practiced in tea
plantations.
15. Tillage: Tillage is done for preparing good seedbed, conservation of soil moisture
& weed control. Tillage removes weeds from the soil resulting in their death. It
may weaken plants through injury of root and stem pruning, reducing their
competitiveness or regenerative capacity: Pre-plant tillage helps in burying the

existing weeds. Bring the seeds to the soil surface for germination and their
subsequent destruction by suitable secondary tillage implements. Incorporation of
pre-plant herbicides. Post plant tillage (row cultivation) helps in mixing of
manures and fertilizers & control of weeds, soil and water conservation.
16. Mulching: Principle is exclusion of sunlight from environment. Polythene Sheets,

natural materials like paddy husk, ground nut shells, saw dust etc. are used as
mulching material. The thickness should be enough to cut off light (i.e. 10-15
cm).The efficiency of polythene sheet is more (more polythene) if it is applied in
continuous sheet rather than in particle farm. It is effective against annual weeds
and perennial weeds like cynodon dactylon and sorghum halopense. Mulching is used
in high value crops like coffee tea plantations by using guatemala grass (
Tripsacum laxum ) and citronella grass ( Cymbopogan spp )
17. Flooding: Flood kills weeds by excluding oxygen from their environment. Flooding
is a worldwide crop husbandry method of controlling weeds in rice fields.
Biological Control of Weeds

In this approach of weed management, utilization of natural living organism, such as
insects, herbivorous fish, other animals, disease organisms and competitive plants are
employed to limit their growth. In biological control method, it is not possible to
eradicate weeds but weed population can be reduced. This method is not useful to control
all types of weeds.
Introduced weeds are best targets for biological control. Bio-control started in the year
1900.The control of Opuntia spp (prickly pear) in Australia and Lantana in Hawaii with
certain insect bioagents are two spectacular examples of early period biological control of
weeds.
Merits
1) Least harm to the environment

2) No residual effect
3) Relatively cheaper and comparatively long lasting effect
4) Will not affect non-targeted plants and safer in usage
5) It is very effective in control of weeds in non-cropped areas
6) Besides this some of the fish, snails and other animals convert weed vegetation
into seafood
Demerits
1) Multiplication is costlier
2) Control is very slow
3) Success of control is very limited
4) Very few host specific bio-agents are available at present
5)

Approaches in Biological Weed Control
Mainly, there are 2 approaches:
1) Classical biological control
Main objective of classical biological weed control is restoring balance between target
alien weed and its natural enemies in the ecosystem by introduction of suitable, exotic
bio-agent. Successful bio-agent reduce the weed population first then the Bio-agent
population due to starvation of food. After some time the bio-agent population may
recover. This process continues in cyclic fashion till the bio-agent and weed population
gets established at a low level. This method is a slow operating and currently used in
non-cropped areas. In crop fields, the bio-agent will not get opportunity to work on host
weed due to frequent use of insecticides and fungicides in modern agriculture. Otherwise
Cyperus rotundus can be controlled in crop fields with moth “Bactra verutana” and selective
bio control of Ludwigia parviflora (water purslane) by Haltica cyanea (steel blue beetle) in
rice fields.
Criteria / Characteristics of successful bio-agent
i. Host-specific: Bio-agents should be host specific and they should not attack other
economic plant spp. They should pass starvation test i.e. they prefer to starve to death
rather feed upon other than host weeds. Lantana was controlled by “Teleonemia
scrupulosa” insect bio-agent. But in India it is likely to damage teak (Tectona grandis) and
sesame (Sesamum indicum). Zygogramma bicolarata is an effective leaf eating bio-agent
against Parthenium (carrot grass). But it is found to attack sunflower in India.
ii. Bio-agent hardiness: Bio-agent should free from its own parasites and predators. Bio-
agent should withstand starvation for short or long periods of food shortage when the
target weed population is brought to low level. But carp can’t survive even a short period
of starvation.
iii. Feeding habit: Bio-agents are more efficient in controlling weeds if they attack either
flowers or seeds of the weed or bore into the stems than root and leaf feeders. But root-
feeding insects are more effective in controlling perennial weeds.
iv. Ease of multiplication: Bio-agent should have high rate and ease of natural
reproduction. It is very important for insects, pathogens, snails and competitive plants.
But it is not desirable with carp as its increased population compete with natural fish.
Kinds of Classical Bio-agents
Bio agent may be either specific or nonspecific. Specific bio agent attack only one or two
specific weeds, while nonspecific bio agent feed upon a variety of vegetation. Specific bio
agents are insects, plant pathogens and competitive plants. Nonspecific bio agents are
Carp fish, snails, and mites.
Six kinds of Bio-agents were used to control weeds. They are Insects, Carp fish, Fungi,
Competitive plants, Snails and mites

1. Insects: These are largely host specific i.e. one insect spp is employed to destroy
the only one weed sp. First successful example reported from Hawaii in 1902
“Lantana camara” controlled by Moth “Crocidosema lantana”. Insects that were
found effective belong to Lepidoptera, Hemiptera, Coleoptera, Diptera.
2. Carp fish: Certain fresh water Carp fish consume large quantities of aquatic weeds.
Whiteamur (Chines grass carp) “Ctenopharyngodon idella” is promising spp for
aquatic weed control. This can grow more than its body weight i.e. 5 kg/year and
attaining up to 50kg at its full size. Herbivorous fish are not food specific. Whereas
the common carp (Cyprinus carpio) a non-herbivorous fish used to control
submerged aquatic weeds.
3. Plant pathogen: Many fungi attack specific weed spp. For instance, “Acacia glauca
“controlled by spore suspension of “Cephalosporium zonatum”. Skeleton weed
(Chondrilla juncia) controlled by rust causing fungi “Puccinia chondrillana.”
4. Competitive plants: Certain plants sp are very competitive in suppressing specific

weeds. Slender spike rush (“Eleocharis acicularis”) aquatic plant can cover the canal
bottom and it is not allowing to establish destructive tall weeds. Typha sp can be
controlled by “Panicum purpurascens” or “Brachiaria mutica” (Para grass). Marigold
has potential to displacing Parthenium spp. Cassia sericea also suppressed the
Parthenium.
5. Snails: The large tropical fresh water snail “Marisa cornuarietis” feed on aquatic
weeds. Marisa feed on roots of water hyacinth, water lettuce and leaves of
“Salvinia”.
6. Mite: The mite “Tetranychus desertorum” controls prickly pear “Opuntia dellini”
Outstanding examples of Classical Bio-control
 Lantana Camera: Lantana was the first weed controlled successfully with certain
insect bioagents in Hawaii. Of this Crocidosema lantana, a moth was found to be
promising in destroying flowers and seeds of lantana. In Australia, three
successful insect biocontrol agents are hispine beetles (Octotoma scabripennis and
Uroplata girardi) and tingid /lantana bug (Teleonemia scrupulosa).
 Prickly pear (Opuntia sp): In Austrlia biocontrol of “Opuntia inermis”.with a moth

“Cactoblastis cactorum “.In Tamilnadu and Maharashtra 40,000 ha land was
recovered from the weed Opuntia delini by releasing “Dactyloplius tomentosus”. a
Cochineal scale insect.
 Water hyclinth: (Eichornea crassipes) it is world wide aquatic weed infested

transplanted paddy fields including India. Hyacinth moth Sameodes albiguttalis
feed up on young leaves and apical buds. Besides this beetles Neochetina bruchii and
N. eichorniae are also damaging the water hyacinth.
Some examples of Classical Bio-control

Weed Bio-agent Kind of bioagent
Chondrilla juncea Puccina chondrillina Plant pathogen
Cirsium arvense Septoria cirsii Plant pathogen
Cyperus rotundus Bactra verutana Shoot boring moth
Echinochloa spp. i) Emmalocera sp. i)Stem boring moth
(In rice fields) ii) Tripos spp. ii) Shrimp
Eupatorium Entyloma compositarum Plant pathogen

riparium
Hydrilla verticillata Hydrellia pakistanae Shoot fly
Orobanche cernua Sclerotinia sp. Plant pathogen
Parthenium i)Zygogramma bicolorata Leaf eating beetle

hysterophorus
ii)Epiblema strenuana Stem galling insect
iii) Conotrachelus sp. Stem galling insect
2) Bio-herbicide Philosophy of Weed Control
Bio-herbicides are pathogens cultured artificially and made available in sprayable

formulations; just like a chemical herbicide. The pathogen selected for the purpose is
usually from the native place of the weed, but it could also be from other places. The bio-
herbicides are also sometimes called mycoherbicides. A mycoherbicides can be both
specific and non-specific.
The bio-herbicide philosophy differs from the classical bio-control philosophy referred to earlier,
in certain ways as follows:
Bio herbicide remains active only on the current weed population, without any chance of
cyclic perpetuation of the weed (or of the bio gent); each new flush of the weed thus
requiring retreatment with it. Bio herbicide can be developed for selective control of
weeds in a crop just like any other selective herbicide, which is not the case with the
classical philosophy bio agents. The development of bio herbicides is of great interest to
industrialists since it involves every season requirement of the product for field use. In
variance with it, the classical biological control approach has no incentive to the private,
profit-oriented organizations; it must depend solely upon public sector support.

Some examples of Commercial Mycoherbicides
Product Content Weed controlled
De-Vine A liquid suspension of fungal spores of Strangler-vine.

Phytophthora palmivora It causes root rot in (Morrentia odorata) in
the weed. citrus orchards.
Collego Wettable powder containing fungal spores of Jointvetch (Aeschynomone

Colletotrichum gloesporiodes. The bio-herbicide sp). In rice fields.
causes stem and leaf blight in the weed.
Bipolaris A suspension of fungal spores of Bipolaris Johnsongrass (Sorghum

sorghicola halepense)

Chapter 3 [Soil Science] Soil
Forming Processes
The geological weathering produces weathered rock material i.e. the parent material and
when the genetic factors set the stage for soil development. The pedogenic processes
change the parent material in to soil with varying horizonations.
The Pedogenic/Soil forming processes are extremely complex and dynamic involving
many chemical and biological reactions, and usually operate simultaneously in a given
area. One process may counteract another, or two other processes may work
simultaneously to achieve the same result.
The relationship between pedogenic processes and genetic factors contributes in some
manner or another, to the pedogenic development of each mature soil. All the genetic
factors contribute to the development of each soil, but no single soil is influenced by all
pedogenic processes.
The sequence of processes in the formation of soils is:
Rock ->Weathering ->Regolith ->Soil forming factors and processes ->True soil
The basic pedogenic processes involved in soil formation:
 Gains or additions of water (mostly as rainfall) organic and mineral matter to the
soil.
 Losses of above material from soil.
 Transformation of mineral and organic substances with in the soil.
 Translocation or movement of soil materials from one point to another with in the
soil. It is usually divided into
i) Movement in solution (leaching) and
ii) Movement in suspension (eluviation) of clay, organic matter and hydrous oxides.
In contrast, the major changes that retard horizon differentiation are due to:
 Mixing of materials by burrowing animals
 Removal of surface soil by erosion (water or wind)
 Creep (by shifting old and its replacement by new materials); and
 Accretion of sediments in cultivated flood – plain areas, for instance silting of

irrigated areas

Basic / Fundamental Pedogenic Processes
I. Humification
Humification is the process of transformation (decomposition) of raw organic matter in

to ‘HUMUS’. It is an extremely complex process involving various organisms such as
bacteria, fungi, actinomycetes, earth worms and termites.
The decomposition of organic matter takes place in two phases: mineralization and
humification. Mineralization is a biochemical breakdown of dead plant tissues by soil
microorganisms to produce simple structured soluble organic substances, mineral
compounds, metal cations and gases (CO2). During the second phase, that is
humification; soluble organic substances regroup themselves in to large molecules by
polymerization and become poorly soluble. They form major part of soil humus and
provide site for retention of cations. The other part of humus is the polysaccharides –
gummy products of microbial excretions, which help in soil aggregation.
The activities of microorganisms in soil formation are as under:
 Mor: It refers to surface soil horizon developed under acid litter and humus from
coniferous and healthy vegetation, where fungi activity predominates.
 Mull: Designated as forest soil horizon (A1) is of intimately mixed mineral matter
and amorphous humus. It is slightly acid and is best developed under base rich
litter, where bacterial activity predominates.
 Sward: Is a dominantly rhizogenous A1 horizon in grasslands as contrasted with

zoogenous mull horizon of forest soils. This includes mollic epipedon or Ap
horizon formed by cultivation of forest soils, in general.
 Orterde: Is a humus rich B horizon in podzols.
II. Eluviation
Eluviation means “Washing out”. It is the process of removal of constituents in

suspension or solution by the percolating water from the upper to lower layers. The
eluviation encompasses mobilization and translocation of mobile constituents resulting
in textural differences.
Mechanical movement of clay and iron oxides from ‘A’ horizon without undergoing
chemical alteration is called Lessivage. Leaching refers to the movement and removal of
material in solution from the soil. It connotes the removal of the dissolved material from
the entire solum.
Elemental mobility
Ca2+ Na+ > K+, Mg2+ >>>> Fe2+ >> Si4+ >> Al3+
(Most mobile) (Least mobile)

The leaching of an element depends not only on its relative mobility but also on the rate
of water percolation through the soil. The effect of leaching is well illustrated with the
depth of accumulation of CaCO3 in soils (Jenny, 1941).
Hot arid zone - < 50 cm
Less hot semi-arid zone - 100 – 150 cm
Sub humid zone - >150 cm
III. Illuvation
The process of deposition of soil materials (removed from the eluvial horizon “E”) in the
lower layer (or horizon of gains having the property of stabilizing translocated clay
materials) is termed as “illuviation”. The horizons formed by this process are termed as
illuvial horizons (B-horizon especially Bt).
All these basic pedogenic processes, combine to result in a number of wide ranging soils
that are observed on surface of the earth.
Hot semi-arid - Calcisol
Cool humid - Podsol
Hot humid – Ferralsol
Specific Pedogenic Processes
The basic pedogenic processes provide a frame work for later operation of more specific
processes.
1. Calcification: The process of precipitation and accumulation of calcium carbonate in

some part of the profile is called calcification. This is a common process in arid and
semi-arid regions, which are low in rainfall (Rainfall<PET).
The illuviated horizon of CaCO3 is designated as ‘calcic horizon’. Whenever high carbon
dioxide is produced in soils, it combines with water and forms into carbonic acid. This
dissolves the calcium carbonate in soils into soluble calcium bicarbonate, which moves
along the percolating water. Again wherever a situation of high temperature and low
carbon dioxide prevails, there calcium carbonate precipitates.
CO2 + H 2O → H2CO3; H2CO3 + Ca→ Ca (HCO3)2
Temp
Ca (HCO3)2 2CaCO 3 + 2H2O + 2CO2
CO2
The calcium compounds are in solution as long as the CO 2 concentration or supply is

maintained. The depth of “calcic horizon” depends on percolating rain water, ground

water depth, amount of rainfall and the texture of the soil. The depth of calcareous layer
In hot arid zone- <50cm, in semi-arid zone 100-150cm and in sub-humid zone the
calcium carbonate accumulates at a depth of >150cm
2. Decalcification: It is the reverse of calcification that is the process of removal of CaCO3

or calcium ions from the soil by leaching. This occurs mostly in high rainfall or humid
regions.
CaCO3 + CO2 + H2O→Ca (HCO 3)2 [Soluble]
3. Podzolization [Pod = under and zola = ashlike (Russian)]: It is a process of soil

formation resulting in the formation of podzols and podzolic soils. It is the process of
accumulation of silica and eluviation of sesquioxides. It is almost a reverse of calcification
process due to leaching of all bases including calcium. The favourable conditions for
podzolization are:
 A cool and humid climate (Invariably found at high altitudes)
 Siliceous (sandy) or acidic parent material, having poor reserves of weatherable

minerals, favor the operation of podsolization, as it helps in easy percolation of
water.
 Acid loving vegetation, such as coniferous pines (Pinus roxburghii), hemlock (Tsuga
Canadensis) and heath ( Calluna vulgaris) are essential for this process.
 Under calcium free, acidic environment (pH <5.0), fungi plays active role in
organic matter decomposition
 Less microbial activity declines the polysaccharide production and keeps the
soluble organic products in soluble form.
 The soluble organic acids react with sesquioxides and the remaining clay minerals,
forming organic - sesquioxide and organic - clay complexes, which are soluble
and move with the percolating water to the lower horizons.
 Alluminium ions in solution hydrolyse and make the soil solution very acidic.
2Al + 6H2O→ 2Al (OH)3 + 6H+
As the materials move out, it gives a bleached appearance (E-horizon) below the surface.
The eluviated materials deposit in B horizon as dark coloured Bh (precipitated humus),
reddish brown Bs (deposition of sesquioxides) and a yellowishish brown (silicate clay)
layer which gradually merges with parent material. Hence a mature podzol has well
developed horizonation. Podzols are highly acidic, low in fertility and used for forestry or
pastures. Rarely crops like oats. Potato and clover can be cultivated.
4. Laterization: The term laterite is derived from the word “later” means “brick” or
“tile”. In tropics, certain soils are massively impregnated with sesquioxides to the extent
of 70 to 80% of the total mass, and forms a cemented horizon, which when dried

becomes very hard like a brick. This soil forming process is called “laterization” or
“Lotozation” Eg: Soils of Malabar hills of Kerala.
In laterization, unlike podzolisation, silica is removed leaving sesquioxides to remain in

solum. The favorable conditions are:
 Warm and humid (tropical) climate with 2000 to 2500 mm rainfall and continuous
high temperature (+25°C) throughout the year. Rapid decomposition of parent
material and organic matter, and intensive leaching are very likely in this climate.
 The rain forests of tropical areas are the suitable vegetation for this process. Under
this vegetation organic additions are low but organic matter decomposition is at
very high rate.
 Basic parent materials, having sufficient ferro-mangnesion minerals (Pyroxenes,

amphiboles, biotite and chlorite) are congenial for the development of laterites.
The iron released during weathering is oxidized to form FeO, Fe2O3 and coats clay, silt or
sand particles imparting characteristic red color to soils. The Al-oxides /hydroxides
impart grey coatings to the soil particles.
The high temperature, intense leaching and basic kind of parent material all favor the
removal of silica (de-silication) and accumulation of sesquioxides. The soluble basic
cations are quickly released during weathering, moves freely in the soil profile and shoots
up the pH to neutrality. Under this basic environment silica liberated from parent
material is solubilized and leached. The solubility of quartz and amorphous silica
increases with increased temperature.
The sesquioxides are left behind as these are more stable under these conditions. As the
alkaline bases are removed from the seat of their formation, the residual soil is acidic in
reaction. Though considerable eluviation takes place, there is no marked horizonsiation
as the eluviated materials are not re-deposited in the lower layers.
Laterite soils are non-plastic, non-cohesive and have granular structure. They are low in
cation exchange capacity and fertility. Phosphorus fixation is high in these soils.
Plantation crops are usually grown on these soils.
5. Gleization: “Glei” means blue, grey or green clay. The gleization is a process of soil
formation resulting in the development of a glei (or gley) horizon in the lower part of the
profile above the parent material due to poor drainage conditions or water logged
conditions. Such soils are called “hydromorphic soils”. This process is not particularly
dependant on climate (high rainfall as in humid regions) but often on drainage
conditions. Poor drainage may be due to lower topographic position, impervious soil
parent material and lack of aeration.
Under anaerobic conditions, iron compounds are reduced to soluble ferrous forms. The
reduction of iron is primarily biological and requires both organic matter and the
microorganisms capable of respiring anaerobically. Iron exists as Fe 2+ organo-compexes
in solution or as a mixed precipitate of ferric and ferrous hydroxides, which is responsible

for the production of typical bluish to grayish horizon. Due to seasonal fluctuations of
ground waters, the gley shows distinct mottling of yellow and rusty brown colors caused
by the alternate oxidation and reduction phenomena.
6. Salinization: It is the process of accumulation of salts such as sulphates, chlorides of

calcium, magnesium, sodium and potassium in soils in the form of salty (salic) horizon.
As a result of the accumulation of salts, solonchalks or saline soils develop with an
electrical conductivity of > 4 dSm-1. The soils are called saline soils, which have ESP less
than15 per cent and pH between 7 and 8.5.
The responsible factors:
 Arid or semi-arid climatic conditions, associated with shallow and brackish (high
amounts of sulphates and chlorides) ground waters.
 Lower topographic positions / depression land forms.
 Imperfect or poor drainage conditions
 Old lake bottoms
 Alluvial deposits along the sea coasts
 Use of saline irrigation waters
The ground water containing high salts moves in an upward direction by capillary action.
The water on evaporation leaves the salts behind, which accumulate at the surface or at
some depth depending upon the capillary fringe. Surface accumulation of salts gives
white appearance to soils .Hence the soils are called as white alkali soils. These soils can
be managed by leaching of salts followed by provision of sub-surface drainage.
7. Solonization or Alkalization: The process involves the accumulation of sodium ions on

the exchange complex of the clay to an extent of >15%, resulting in the formation of sodic
soils (solonetz) under arid and semi-arid conditions. This occurs when anions like
carbonates and bicarbonates predominate in soil.
The calcium and magnesium in soil solution will precipitate as corresponding carbonates
or bicarbonate whenever the ionic product of solution exceeds the solubility products of
respective carbonates. This reduces the concentration of Ca and Mg in soil solution, there
by releasing them from exchange complex. As this process continues the sodium
concentration on exchangeable complex increases. When the ESP in soils exceeds 15 %,
the soil is designated as alkali soil with a high pH of >8.5, which results in less nutrient
availability.
The high pH in soils results in dissolution of humus, which moves upward along the
capillary water giving black colour to soils. Hence the soils are called black alkali soils.
High sodium on clay minerals results in dispersion of soil aggregates leading to physical
problems like poor aeration, low infiltration and percolation of water.

This process results in a very thin, friable horizon followed by a dark horizon of hard and
impermeable heavy soils generally with illuviated clay and having a typical columnar
structure, which is characteristic of “solonetz”.
8. Solodization or Dealkalization: This process refers to the removal of Na+ from

exchange sites. The Na+ can be eliminated by increasing the concentration of Ca2+ or
Mg2+ in the water, followed by improved drainage facilities.
2NaX + CaSO 4→ Na2SO4 + CaX
9. Pedoturbation: It is the process of mixing of the soils. Mixing to some extent takes
place in all soils.
 Faunal pedoturbation: Mixing by animals such as ants, earthworms, moles,

rodents and man himself.
 Floral pedoturbation: Mixing by plants, as in tree tipping that forms pits and
mounds.
 Agrillopedoturbation: Mixing of materials in solum by churning process caused by

swell - shrink clays as is observed in deep black cotton soil.
 Calcification, podzolization and laterization are zonal soil forming processes,

where in the profile characteristics are influenced by prevailing conditions of
climate and vegetation.
 Gleization, salinization, solonization and solodization are the Intra-zonal soil

forming processes, wherein, the profile characteristics are more influenced by
certain local conditions, such as relief (topography) and / or parent material than
the climate and vegetation.
 Salinization, alkalization and dealkalization processes operate in sequence and

advance the soil to a certain point in transition until the zonal soils are formed.

Chapter 4 [From Plant Breeding]
Use of Polyploidy in Plant Breeding
The somatic chromosome number of any species, whether diploid or polyploid, is

designated as 2n, and the chromosome number of gametes is denoted as n. An individual
carrying the gametic chromosome number, n, is known as haploid.
A monoploid, on the other hand, has the basic chromosome number, x. In a diploid
species, n=x; one x constitutes a genome or chromosome complement. The different
chromosomes of a single genome are distinct from each other in morphology and or gene
content and homology; members of a single genome do not show a tendency of pairing
with each other.
Thus a diploid species has two, a triploid has 3 and a tetraploid has 4 genomes and so on.
In euploids, the chromosome number is an exact multiple of the basic or genomic

number. Euploidy is more commonly known as polyploidy.
When all the genomes present in a polyploidy species are identical, it is known as
autopolyploid and the situation is termed as auto polyploidy.
In the case of allopolyploids, two or more distinct genomes arepresent. Euploids may
have 3 (triploid), 4 (tetraploid), 5 (pentaploid), or more genomes making up their somatic
chromosome number.
In case of autopolyploidy, they are known asautotriploid, autotertaploid, autopentaploid,

and so on, while in the case of allopolyploidy they are termed as allotriploid,
allotetraploid, allopentaploid, etc. Amphidiploid is an allopolyploid that has two copies of
each genome present in itand, as a consequence, behaves as a diploid during meiosis. A
segmental allopolyploid contains two or more genomes, which are identical with each
other, except for some minor differences.
Breeding Autopolyploids
Origin and production of doubled chromosome numbers:
1. Spontaneous: chromosome doubling occurs occasionally in somatic tissues and

unreduced gametes are produced in low frequencies.
2. Production of adventitious buds: decapitation in some plants leads to callus

development at the cut ends of the stem. Such a callus has some polyploid cells
and some of the shoot buds regenerated from the callus may be polyploid.
Insolanaceae, 6-36% of adventitious buds are tetraplods. The frequency of
ployploid buds may be increased by the application of 1% IAA at the cut ends as it
promotes callus development.

3. Treatment with physical agents: Heat or cold treatment centrifugation, x-ray
orgamma ray irradiation may produce polyploids. Exposing the plants or ears of
maize to a temperature of 38-45oC at the time of the first division of zygote
produce 2-5 % tetraploid progeny.
4. Regeneration in vitro: polyploidy is a common feature of the cells cultured in-

vitro.
5. Colchicine treatment: Colchicine treatment is the most effective and the most
widely used treatment for chromosome doubling. Chromosome doubling reduces
the risk of meiotic complications, and thus increases chances of fertile
autopolyploids.
Morphological and cytological features of autopolyploids
1. Polyploids have larger cell size than diploids. Guard cells of stomata are larger the
number of stomata per unit area is less in polyploids than diploids.
2. Pollen grains of polyploids are generally larger than those of the corresponding
diploids.
3. Polyploids are generally slower in growth and later in flowering.
4. Polyploids usually have larger and thicker leaves, and larger flowers and fruits
which are usually less in number than in diploids.
5. Polyploids generally show reduced fertility due to irregularities during meiosis

and due to genotypic imbalance leading to physiological disturbances.
6. In many cases, autopolyploidy leads to increased vigour and vegetative growth.
7. Different species have different levels of optimum ploidy. For sugar beet the
optimum level is 3x, sweet potato 6x while for timothy grass it is between 8-10x.
8. Autopolyploids generally have a lower dry matter content than diploids.
Application of Autopolyploidy in Crop improvement
Triploids
Triploids are produced by hybridization between tetraploid and diploid strains. They are
generally highly sterile, except in a few cases. This feature is useful in the production of
seedless watermelons. In certain species, they may be more vigorous than the normal
diploids, e.g., in sugar beets.
Seedless watermelons are produced by crossing tetraploid (4x, used as female) and
diploid (2x, used as male) lines, since the reciprocal cross (2x x 4x) is not successful. The
triploid plants do not produce true seeds; almost all the seeds are small, white
rudimentary structures like cucumber (Cucumissativus) seeds. There are several problem

viz. genetic instability of 4x lines, irregular fruit shape, a tendency towards hollowness of
fruits, production of empty seeds and the labour involved in triploid seed production.
1. Triploid Sugar beets: Among root crops triploid sugar beets apparently represent the
optimum level of polyploidy because 3n plants have longer roots than diploid and also
yield more sugar per unit area.
2. Tetraploid Rye: The advantage of tetraploid over its diploid counterpart are large
kernel size, superior ability to emerge under adverse condition and higher protein
content. Tetraploid rye varieties have been released for cultivation e.g. Double steel, Tetra
petkus.
Limitations of autoployploidy
1. Larger size autopolyploids generally contain more water (gigas effect) and produce less
dry matter content than diploids.
2. High sterility with poor seed setting is observed
3. Due to complex segregation, progress through selection is slow.
4. Monoploids and triploids cannot be maintained except through clonal propagation
5. The varieties cannot be produced at will
6. Effects of autopolyploidy cannot be predicted.
Allopolyploidy
Allopolyploids have genomes from two or more species production of allopolyploids has
attracted considerable attention; the aim almost always was creation of new species.
Some success has been evident from the emergence of triticale, Raphanobrassica and
allopolyploids of forage grasses.
Morphological and cytological features of allopolyploids
1. Allopolyploids combine the morphological and physiological characteristics of

the parent species but it is very difficult to predict the precise combination of
characters that would appear in the new species.
2. Many allopolyploids are apomictic as in Tulips, Solanum.
3. The chromosome pairing in the new species depends upon the similarities
between the chromosomes of the parental species. Chromosomes with such
similarities are known as homeologous chromosomes. After chromosome
doubling, the allopolyploid would have two homeologous chromosomes for each
chromosome present in the F1 hybrid, comparable to the diploid species. Such
allopolyploid is referred as amphidiploid or Allotetraploid.
4. Fertility of allopolyploids can be improved by hybridization and selection.

Application of allopolyploidy in crop improvement
1. Utilization as a Bridge species: Amphidiploids serve as a bridge in transfer of

characters from one species to a related species, generally from a wild species to
cultivated species. Creation of new crop species as in Triticale, Raphanobrassica.
2. Widening the genetic base of existing Allopolyploids: The genetic base of some
natural allopolyploids may be narrow, and it may be useful to introduce variability
in such cases by producing the allopolyploids afresh from their parental species.
Limitations of Allopolyploidy
1. The effects of allopolyploidy cannot be predicted. The allopolyploids have some

features from both the parental species, but these features may be the undesirable
ones, e.g., Raphanobrassica, or the desirable ones, e.g., Triticale.
2. Newly synthesized allopolyploids have many defects, e.g., low fertility, cytogenetic
and genetic instability, other undesirable features etc.
3. The synthetic allopolyploids have to be improved through extensive breeding at

the polyploidy level. This involves considerable time, labour and other resources.
4. Only a small proportion of allopolyploids are promising; a vast majority of them

are valueless for agricultural purposes.
Aneuploidy
Aneuploidy is referred to individuals with an incomplete set of chromosome that is

equivalent to the euploid number plus or minus one or more specific chromosome. As in,
normal set if is 2n then 2n-1, 2n-2, 2n+1 are referred to as monosomy, nullisomy, and
trisomy respectively.

Chapter 5 [From Entomology]
Storage Pests & their Management
Types and Species
o Stored grain pests cause loss in quality and quantity of food grains. Nearly 5-10%
of grains are lost in storage due to insects and mites, especially beetles and moths.
o They also contaminate food, lower its nutritive value, and create conditions
favorable for mould growth.
o Primary pests are capable of initiating injury on undamaged grains. e. g. rice

weevil, granary weevil, lesser grain borer, and angoumois grain moth.
o Secondary pests tend to feed on the grains that have already been damaged. e. g.
red and confused flour beetles, Indian meal moth, Mediterranean flour moth, and
the saw-toothed grain beetle.
I. Storage Beetles
Khapra beetle, Trogoderma granarium (Dermestidae: Coleoptera)
o Larvae chew up the seed coat and grain into frass.
o Attacks wheat, maize, sorghum, pulses etc
o Adult: brown with or without red markings, 2-3 mm long.
o Egg: 125 eggs, singly or 2-5 eggs, period 6-16 days
o Larva: yellowish white later brownish, period 3-7 weeks
o Pupa: pupation in last larval skin, period 4-6 days
Lesser grain borer, Rhyzopertha dominica (Bostrychidae: Coleoptera)
o Damage: grubs and adults bore irregular holes
o Paddy, wheat, maize
o Life cycle 2 months
o Affect the grains in both larval and adult stages
o Adult: small, cylindrical, 3mm long, dark brown
o Egg: white elongate, oval, 300-500 eggs, incubation 5-6 days
o Larva: white, oligopodous, period 40 days

o Pupa: white, exarate, 7-8 days
Rice weevil, Sitophilus oryzae
Maize weevil, Sitophilus zeamais (Curculionidae: Coleoptera)
o Adults and larvae infest the grain and flour.
Saw toothed grain beetle, Oryzaephilus surinamensis (Cucujidae: Coleoptera)
o Damage: scaring of grain surface, burrowing holes.
o Attacks rice, wheat
o Life cycle: 3-4 weeks
o Adult: elongate (3 mm), brown, 3 longitudinal ridges, saw-like edges
o Egg: elongate, white, shiny, period 4 -12 days
o Larva: yellowish white, conical apex to abdomen, 4 mm, period 2 weeks
o Pupa: creamy white, exarate, period 1 week
Red flour beetle, Tribolium castaneum (Tenebrionidae: Coleoptera)
o Damage: grubs and adults infest stored foods, broken grains
o Attacks wheat flour, rice bran, flours of other grains
o Life cycle: 6 weeks
o Adult: flat, shiny, reddish brown, 2 mm
o Egg: elongate, oval, whitish, 400-450 eggs on grains, period 4-12 days
o Larva: whitish, 3 mm, period 27-30 days
o Pupa: yellowish white, hairs on dorsal surface, period 6-9 days
Pulse beetle, Callosobruchus chinensis (Bruchidae: Coleoptera)
o Damage: grub, cavity made, round exit holes, cigar shaped eggs
o Attacks stored cowpea, grams, lablab (pulses)
o Adult: reddish black, 3 mm, serrated antenna
o Egg: laid singly, several on one grain, translucent, smooth, shiny, later greyish
white, period 4-5 days
o Larva: white, 6 mm, cylindrical, fleshy, wrinkled
o Pupa: brown, period: 4 days (summer), 4 weeks (cooler month)

o Life cycle: 2 months
Cigarette or tobacco beetle, Lasioderma serricorne (Anobiidae: Coleoptera)
o Adult: light brown, humped appearance, elytra with minute hairs, antenna
uniform thickness
o Damage: grubs and adults, pin head sized bore holes
o Attacks tobacco, ginger, wheat, peanut, bean, coriander, chilli, turmeric, dried
fruits.
o Egg: creamy white, oval, period 9-14 days
o Larva: whitish hairy, oligopodous, period 17-29 days.
o Pupa: exarate, period 2-8 days
Drug store beetle, Stegobium paniceum (Anobiidae: Coleoptera)
Damage: circular pin head size hole made on turmeric, coriander, ginger, etc.
o Adult: reddish brown, striated elytra,3 mm, antenna clubbed
o Egg: batches (10-40 eggs)
o Larvae: not hairy, pale white, fleshy with abdomen terminating in 2 dark horny
points, period- 10-20 days
o Pupa: period 8-12 days
Longheaded flour beetle, Latheticus oryzae (Tenebrionidae: Coleoptera)
o Damage: grubs and adults feed on milled products, secondary infestation
o Attacks cereal flour, packaged food, rice and rice products
o Adult: yellowish brown, flat, antennae 10 segmented, eye with anterior and
posterior projections, 2.5-3mm
o Egg: 7-8 eggs, oval, white to opaque, period- 3 days.
o Larva: white, finely hairy, 5 mm, period 15 days
o Pupa: yellowish white, thorax concealing the head, period 3-7 days
o Life cycle: 45-55 days
Flat grain beetle, Cryptolestes minutus (Cucujidae: Coleoptera)
o Damage: grubs and adults feed on broken grains, heavy infestation
o Attacks rice, maize

o Adult: small, light to dark reddish brown, 1.5-2mm
o Egg: white eggs in flour, grain, period 5 days
o Larva: cigar like, yellowish white, 2 reddish brown spots at anal segment, period-
21 days
o Pupa: gelatinous cocoon
o Life cycle: 42 days
Cadelle, Tenebroides mauritaricus (Tenebrionidae: Cleoptera)
o Biggest beetle, 1/4” length, white fleshy body, 2 hard processes at posterior end,
prefers moist places
o Infest whole grains of cereals, millets, processed items like flour, maida
Copra beetle, Necrobia rufipes (Cleridae: Coleoptera)
o Infests stored copra and ground nut kernels
Dried fruit beetle, Carpophilus dimidiatus (Nitidulidae: Coleoptera)
o Attacks dry fruits, copra and groundnut kernels in storage
II. Storage Moths
Angoumois Grain Moth, Sitotroga cerealella (Gelechiidae: Lepidoptera)
o Major pest of stored husked rice
o Attacks rice, wheat, maize, sorghum, barley and oat
o Larvae feed on grain kernels.
o Grey or buff colored moth, nocturnal in habit, 5-10 mm, fore wings with darker
spots, apex of hind wings fringed with hairs, short lived (5-10 days)
o Life period : 4-6 weeks
o Whitish eggs are laid on the grain, hatch in a week
o Larva yellowish white in color with dark head; enters the grain through crack or
split in the husk and closes the hole with silken web
o Feeds on the grain kernels and the grain gets filled with refuse
o Pupates inside the grain spinning a silken cocoon; pupal period 7-10 days
Rice Moth, Corcyra cephalonica (Galleriidae: Lepidoptera)
o Serious pest of milled rice

o Attacks rice, rice bran and other stored products
o Larva webs the grains that adhere to each other (caking).
o When more than 16 larvae develop in 250g of rice, the grain color changes with an
unpleasant odour.
o Adults nocturnal, pale greyish moths, fore wings uniform in color, hind wings
creamy white with fringe
o Life cycle period: 7-12 weeks
o 10-320 eggs laid in crevices of stores; white in color, oval in out line; incubation
period: 5-7 days
o Larva varying in color from whitish to grey; live inside a tough silken tunnel;
larval period : 5-6 weeks
o Pupa dark brown; pupal period 7-10 days inside a silken coon
Indian meal moth, Plodia interpunctella (Phycitidae: Lepidoptera)
o Infests a variety of food: stored grains, flours, corns, nuts, powdered milk
o Larvae web together the milled products that become dirty silken masses
containing the excreta of larvae.
o Adults have peculiar markings on the forewings, i. e. reddish brown with a copper
luster on the outer two-thirds but whitish grey on the inner or body ends
o Whitish , ovate eggs are deposited on grain surface
o Grown up larvae (1-2 cm) are whitish, greenish or pinkish in color and they spin
the webs
o Pupate within a thin silken cocoon
o Life cycle complete within 6 weeks
Warehouse moth, Ephestia cautella (Phycitidae: Lepidoptera)
o It is also called Cocoa moth
o Attacks nuts, almond, cocoa beans and tobacco
o Serious problem in chocolate and tobacco industry
o Adult lays the eggs on the finished product
o The female may lay 200-300 eggs over or near the product
o Larvae are mobile over the products and they produce silk

o Adults are short lived (13-14 days)
o Life cycle takes 50-90 days
Grain Pest Life Cycle Remarks
Granary Weevil 4 weeks Universal feeder on whole grains.
Rice Weevil 4 weeks Universal feeder on whole grains.

Most common stored whole grain
pest.
Coffee Bean Weevil 4 weeks Lays eggs in corn in field; infestations

may continue for 3 months after
storage.
Lesser Grain Borer 4 weeks Universal feeder on whole grains.
Angoumois Grain Moth 5 weeks Most important in stored corn.
Indian Meal Moth 6-8 Prefers coarse grades of processed

weeks grain.
Mediterranean Flour 8-9 Prefers finer grades of processed

Moth weeks grain.
Saw-toothed Grain Beetle 4 weeks Prefers grain products
Red Flour Beetle 5 weeks Attacks grain and grain products.
III. Rodents
Two species of rodents, the Norway rat Rattus norvegicus and the house mouse Mus
musculus infest large storage godowns. They feed and contaminate even more of the
stored material with their droppings, urine, and hair.
IV. Storage Mites
1. Grain mite, Acarus siro
2. Cheese mite, Tyrophagus casei
3. Mold mite, Tyrophagus putrescentiae
4. Dried fruit mite, Carpoglyphus lactis
5. Itch mite, Glycyphagus domesticus
o The cheese mite, known to cause dermatitis, is larger than both the grain mite and
the mold mite.

o The mold mite is a pest of many foods, especially those having a high fat or
protein content. Infestations have been found on wheat flour, soy flour, wheat
germ, rye bread, and mixtures of oats, barley and wheat. Not serious on stored
products in tropical or subtropical regions.
o The dried fruit mite infests dried fruits, jams, and products containing lactic,
acetic, or succinic acids. It has been also found in honeycombs, fruit drink residue,
senna confections, rotting potatoes, flour, dried-milk powder, and caramel used
in manufacturing sweets.
o The grocer's itch mite is found in flour, wheat, linseed, tobacco, sugar, cheese,
and bee frames, in bees' and birds' nests.
Principles and Methods of Management
Bin Storage
• Make needed bin repairs
• Avoid mixing old and new grain
• Treat empty bins prior to filling. After cleaning bins thoroughly, spray walls,
floors and ceilings to the "point of runoff". Apply the insecticide 2-3 weeks before
harvest.
• Before storing the grain, sweep the bin to remove all dead insects.
Loading Grain for Storage
• Reduce the moisture content to a storable level of 13 percent or below.
• The grain bin should not be overfilled
• Space (headroom) must be left on top of the grain.
• After the bin is filled, the grain's surface should be leveled
• Application of insecticidal treatment, aeration, and fumigation cannot be effective

on uneven grain surfaces.
• Moisture condensation and subsequent mold and insect problems are more likely
to develop in mounded grain.
Surface Treatments
• Surface treatments are applied to prevent insects such as moths from entering
the grain from the outside.
• This should be done as soon as the bins are filled and the surface leveled.
Treatments can be repeated if necessary with 57 % malathion EC, 1% malathion
dust, or Bacillus thuringiensis (Dipel).

• The grain should be inspected at 2- to 4-week intervals to monitor these
conditions.
Fumigation
• Fumigation is needed when no other pesticide or control method can reach the
insect infestation.
• If the insects are already inside the grain mass, no spray or dust can reach them.
• Fumigants act on all insect life stages.
• Methyl bromide, aluminum phosphide
• In recent years aluminum phosphide (Celphos, Phostoxin) has been found to be an

effective fumigant for stored grain insects.
• The main advantage of aluminum phosphide is the ease of application.
Pest Management in Storage Go-downs
Maintain store house hygiene
o Eliminate various bio-stages of pests, hiding creeks and crevices.
o Clean the store room or go-down before stocking the grains fresh.
o Arrest rat entry by providing a metal sheet up to a height of 25 cm at the bottom

of wooden doors.
o Provide wire meshes on windows, ventilators, gutters, drains, etc. to prevent the
entry of rats.
o Fill crevices, cracks, rat holes, etc. found on the walls and floor.
o Follow an integrated approach by blending all known technology to minimize the

damage by storage pests.
Reduce the moisture content of the grains to prevent insect build up
o The seeds or grains have to be dried in sun thoroughly and repeatedly to bring
down the moisture content below 10 – 13 %.
o All the bags, bins, etc. which were used previously should be dried in the sun
repeatedly to kill the insect bio-stages harboured in them.
Eliminate conditions which favour storage pests
o Seeds should be sieved and all broken grains removed before bagging since broken
grains favour the pest build up.
o All–torn bags should best stitched before filling.

o Loose bags should be filled or stitched to make them tight.
Surface treatment of store house before storage
The walls, ceiling and floor or empty store rooms or go-downs should be treated with any one of
the following chemicals before stocking the bags:
o Malathion 4 % 25g / sq. m
o Malathion 50 % EC 10 ml/litre of water and 3 litres of spray solution per sq. m
o DDVP 76 % SC 7 ml/litre of water and 3 litres of spray solution per 100 sq. m
Good storage minimizes insect infestation
o Provide good dunnage by arranging wooden blocks or bamboo poles or spreading

thick polythene sheets on the floor.
o The dunnage materials have to be treated with one of the above chemicals
suggested for surface treatment.
o The bags should be arranged over the dunnage in a crisscross pattern, i.e. arrange
one layer of bags length-wise and the next layer of bags breath-wise.
o The bags have to be stored in such a way that they do not lean or touch the walls.
o A gangway or alley of 0.75 to 1.0 meter has to be left around and at convenient
intervals for inspection, aeration, prevention of moisture seepage and for
fumigation and chemical spraying, if necessary.
o Bags should be stored only up to 15 bags high or as specified.
o Adequate space has to be left between the roof and the top layer of the bags.
o Bags containing different seeds or grains should be stacked separately and they
should not be mixed.
Prophylactic treatment of seeds/grains and bag surface
o If the produce is meant for seed purpose, mix 1 kg or activated kaolin or malathion
5% D with every 100 kg of seed and packed in gunny or polythene lined bags.
o If the produce of cereals is meant for grain purpose, mix 1 kg of activated kaolin to
every 100kg of grain and store. To protect the pulses grains, activated kaolin is
mixed at the above dosage or any one of the edible oils at 1 kg for every 100 kg or 1
kg of neem seed kernel for every 100 kg of cereal or pulse grain and stored.
o One of the following pesticides has to be applied at the specified dosage over the bags:
Malathion 5% 25 g per sq. m
Malathion 50% EC 10 ml/ litre of water (3 litres of spray solution per 100 sq. m)

DDVP 76 % SC 7 ml per litre of water (3 litres of spray solution per 100 sq. m)
o No synthetic insecticide should be mixed with the grains which are meant for
consumption.
o Alleys or gang ways have to be treated with one of the following chemicals:
Malathion 50% EC 10 ml/litre of water
DDVP 76% SC 7 ml/litre of water
Apply one litre of spray solution for every 270 cu.m or 10,000 cu. ft.
o The chemicals have to be sprayed on the walls and floors and the treatment
repeated based on the extent of flying and crawling insects.
For continued healthy insect free conditions of seeds or grains
o Draw samples of seeds or grains at fortnightly intervals and classify the

infestation as follows based on the number of insects found per 1 kg of sample:
When there is no pest Free
Up to 2 insects Mild
More than 2 insects Severe
o Decide the need for shed fumigation (entire store house or go-down) or cover
fumigation (only selected blocks of bags).
o The store houses/go-downs and the black polythene sheets or rubberized

aluminum covers should be checked for holes and got them ready for fumigation.
o Choose the fumigant and work out the requirement on the following guidelines:
Aluminium phosphide
1. For cover fumigation: 3 tablets of 3 grams each per ton of grain
2. For shed fumigation: 21 tablets of 3 grams each for 28 cu. metres
3. Duration of fumigation: 5 days
o In case of cover fumigation clay or red earth with water is mixed and made
into a paste and kept ready for plastering all around the fumigation cover,
or sand snakes are kept ready.
o The required numbers of aluminum phosphide tablets are inserted in

between the bags in different layers or methyl bromide is applied as
required.

o The bags should be covered immediately with fumigation cover.
o The edges of cover all around are to be plastered with wet red earth or clay
plaster or weighed down with sand snakes to make leak proof.
o The bags are kept airtight for a period of 5 or 7 days under fumigation
based on the fumigant chosen.
o The mud plaster is to be removed after specified fumigation period and the
cover lifted in a corner to allow the residual gas to escape.
o Aeration has to be allowed and the cover lifted after a few hours.
o Similar steps should be followed to ensure leak proof condition, fumigation

period, aeration etc. in case of shed fumigation.
o Fumigants are used for curative treatments and they have no residual
action on new immigrant insects which can infest grains.
o Sampling has to be done periodically and the stored materials fumigated

based on the need.
o Fumigants should be handled with utmost care as per specifications.
Rodent management in store houses
o Trim the branches of trees or plants hanging over the store house or go-
down.
o Provide metallic mesh to windows, ventilators, drains, etc.
o Man holes or covers of the drainage systems have to be covered.
o The doors should closed tightly without gaps. Do not give clearance of
more than 0.5 cm between floor and door.
o Remove the wooden stairs in front of go-downs as soon as the work is over
and they should never be allowed to remain at nights.
o All rat holes should be plugged with cement after filling with glass pieces.
o If possible, automatic door closers may be provided to prevent entry of

rats.
o Check for the rodent infestation periodically. If rodents are noticed, they
may be baited with multi-dose or chronic anticoagulant rodenticides. In
case of dry concentrated food, the bait may be prepared as follows:
Cereal or flour 450 g
Any edible oil 10 g

Powdered jaggery or brown sugar 15 g
Anticoagulant rodenticides like coumarin compounds 25 g
o Mix the ingredients thoroughly and keep it in small cups on the rat runs, dark
places, etc. where the rats frequently move. Replace the consumed bait daily. The
rats which begin dying should be collected after 5 or 6 days and buried.
o Water soluble bait may be prepared my mixing 25 grams of water soluble

anticoagulant in 475 ml of water. Keep the shallow cups or plates at a number of
places inside the go-down for the rats to drink the poisoned liquid before death.
o If needed, the single dose or acute poison bait may be prepared as follows:
Food materials 97 grams
Any edible oil 1 gram
Zinc phosphide 2 grams
The ingredients are mixed thoroughly and kept in cups in the places frequented by rats.
o Before providing the poisoned zinc phosphide bait the “plain or non– poisoned
bait” are provided for two or three days to make the rats to accept the bait.
o The same material stored in the vicinity has to be used for preparing baits to make
the rats accept and eat them without suspicion.
o Once the rat population has been contained, discontinue the baiting by removing
all the baited food and destroyed.

Chapter 6 [From Horticulture]
Propagation of Horticultural Crops
I. Asexual Propagation
Asexual propagation is the method of multiplication of a plant from a tissue other than
zygote which is formed by the combination of male and female gametes. The cellular
basis for this method of multiplication is mitosis viz., regeneration of a daughter plant
from the somatic tissue. The different methods of asexual propagation are:
A) Cuttings
1. Root cutting – Red raspberry, Bread fruit etc.,
2. Stem cuttings
a. Hardwood –fig, grape, gooseberry, rose etc.,
b. Semi hard wood – coleus, geranium, sweet potato etc.,
c. Softwood-lilac, jasmine etc.,
d. Herbaceous – coleus, geranium, sweet potato etc.,
3. Leaf cutting - Begonica, Bryophyllum, Sansevieria etc.,
4. Leaf bud cuttings – Hydrangea
B) Layering
a) Ground layering
1. Tip layering : Black berry
2. Simple layering: Guava, Pomegranate, crotons etc.,
3. Mound layering : Goose berry, apple etc.
4. Compound layering : Grape, Honey suckle etc.,
5. Trench layering : Etiolation method eg. Cherry
b) Air layering (Gootee (or) marcotage) : Litchi, guava, crotons etc.
C) Grafting
1) Root grafting: Whip graft-apple and pear
2) Crown grafting: a) Whip and tongue graft – Persian walnut, apple

b) Cleft graft – camellia, plums
c) Side graft - Narrow leaved evergreen, mango
3) Top grafting:
a) Cleft – various fruit trees
b) Notch graft
c) Bark graft
d) Side graft
e) Whip and tongue graft
f) Veneer grafting
D) Budding
a) T budding (Shield budding) – Pomefruits, rose, ber etc.
b) Patch budding – Citrus
c) Ring budding – Walnut and pecan
d) Flute budding – Walnut and pecan
e) Chip budding – citrus
E) Tissue culture - Micropropagation
F) Other special parts
Advantages of Asexual Propagation
1. In most horticultural plants, the genetic make up (genotype) is highly

heterozygous. The unique characters of such plants are immediately lost if they
are propagated though seed
2. It is necessary to grow cultivars that produce non viable seeds, eg. Bananas, fig
and grape
3. Propagation of some species may not be easier through seeds . For eg.
Cotoneaster seed – it has complex dormancy condition but it is easily propagated
through cuttings
4. To reduce pre-bearing period/or to reduce long juvenile stage.
5. To induce dwarfness eg. in apple
6. To induce disease and pest resistance. ‘Troyer citrange’ is used as a rootstock for
citrus. It is resistant to tristeza virus.

7. To induce hardiness in cultivars eg. ‘Alnarp’ apple used for its winter hardy
properties
Disadvantages of Asexual Propagation
1. Longevity is not high when compared to the seedling progeny.
2. Asexual method is uneconomical and impractical in the case of vegetable crop

propagation and grains (eg.tomato, brinjal, amaranthus etc.) since cost of
cultivation is high when compared to sexual method
3. Most of the virus disease are not seed borne. When propagated vegetatively the
virus are carried to the next generation eg. ‘Katte’ disease of cardamom.
Genetic variation in sexual propagation
 Gene/chromosome change
 By mitosis, it becomes permanent
 It is found in a part of the plant only
 The plants with normal and mutated cells are called ‘Chimeras’
 Eg. Coleus, crotons, Bougain villeas.
Kinds of Chimeras
1) Sectorial Chimeras- Growing point of the stem is found with two types of tissues.
- The leaves & lateral buds are also mutated
2) Periclinal - The mutated tissue occurs as a thin skin with several cell
layers
- The most common type of chimeras
- Relatively stable
- This type will revert back if propagated by seed or root cuttings
3) Mericlinal - Similar to periclinal
- The outer of mutated cells does not surround fully
- It occupies as a segment of the whole part only
Budsport
 Budsport is one where a branch of a tree alone is found with genetic change from the
rest of the part
 The characters of budsport are inheritable

 They can be vegetatively propagated
 Eg. Apple – varieties –‘star kind’ and Richa Red ar ebudsports from ‘Delicious apple’
Apomixis
It is the occurrence of an asexual reproductive process in place of normal sexual

reproductive process of reduction division and fertilization. Simply, it is an asexual
seedling developed from a seed viz., a seedling that arises from tissue of the seed other
than embryo
Plants that produce only apomictic embryos are known as obligate apomicts, (Eg.
Mangosteen) those that produce both apomictic and sexual embryos are facultative
apomicts eg. Acid lime
Type of apomixes
Recurrent apomixes: Here, embryo develops from the egg mother cell which doesn’t under
go any meiosis. So., egg has normal diploid number of chromosome, the same as in the
mother plant. The embryo subsequently develops directly from the egg nucleus without
fertilization. In some cases, the embryo develops with stimulus of pollination (eg Allium)
and in some cases, without stimulus of pollination (eg. Malus)
Adventitious or Nucellar embryony: Here, embryo will rise from a cell or group of cells
either in the nucellus or in integuments. Here, embryo develops outside the embryo sac
in addition to the regular embryos. Eg. Citrus
Non-recurrent apomixes: Here embryo arises from the egg nucleus without
fertilization. Since the egg is haploid, the resulting embryo will also he haploid. The case
is very rare.
Vegetative apomixes: In some cases, vegetative buds or bulbils are produced in the
inflorescence in place of flowers eg. Agave and grass species
Polyembryony: The phenomenon in which two or more embryos are present within a
single seed is called polyembryony (Nucellar embryony)
Significance of apomixes
1. Apomictic seedlings are true to its mother and apomictic cultivar can be
considered as a clone
2. They are uniform and vigorous
3. Virus diseases are not seed borne. So, it is the best method to rejuvenate virus
affected plant crops.

I. Principles and methods of vegetative propagation by cuttings
Cuttings are vegetative plant portions such as stems, leaves and roots taken from plants
to produce new independent plant which, in most cases, will be identical with the parent
plant. This is one of the least expensive and easiest methods of vegetative propagation.
Cuttings are taken from 1) stem 2) leaf 3) leaf bud and 4) root
In the case of stem cuttings, it has four groups
1. Hard wood cutting-spring – February, March
2. Semi-hard wood cutting-summer – April, may
3. Soft wood cutting -fall (or) autumn – June, july & August
4. Herbaceous cuttings-Winter–September, October, November, December & January
Hardwood cutting
Deciduous Hardwood
The cuttings are fully matured with more reserve food and anatomically, the maximum of
sclerenchyma can be seen. The cuttings are prepared during dormant season (late fall,
winter or early spring) from wood of previous season’s growth. In some species, such as
fig, olive and certain plum varieties, two year old wood can be used. Fruits propagated
through hard wood cuttings are fig, olive, mulberry, grape, gooseberry, pomegranate,
some plums and rose
Cuttings should be taken from healthy plant grown in full sunlight. Length may vary
from 4 to 30 inches (Common 6-8”) the diameter of cuttings may range from ¼ inch to
even 2” depending upon the species.
At least, two nodes are included in the cutting. The basal cut is usually just below a node
and a top cut ½ to 1” above a node. After preparing cuttings, bundles of cuttings may be
buried out of doors in sandy soil or stored in a refrigerated room before planting in
spring.
While planting, cuttings should be planted 3 or 4” apart and deeply enough (1/3 of its
length placed inside the soil)
Evergreen hardwood
Grapes, pomegranate and some

citrus fruits are propagated
through hard wood cuttings.
Length of cuttings range from 4-7”
with 5 to 6 nodes. Cuttings are
taken during late winter. Spring

season is good for planting.
Semi-hard wood cuttings
Stem cuttings of trees and shrubs that are taken from current season shoots, which are
partly matured are known as semi-hard wood. They have lesser reserve food compared
to hard wood and similarly, the formation of sclerenchyma in the anatomical
development is also comparatively less. Length of cuttings ranges from 3 to 6 inches.
Here we can retain one or two terminal leaves.
Soft wood cutting
Cuttings of 3-6” length prepared from soft, succulent and new growth may be called as
soft wood cuttings eg. Vernonia
Herbaceous cuttings
This type of cuttings made from succulent herbaceous plants just near the terminal buds
is called herbaceous cuttings (Geranium, Coleus, Alternanthra and Sweet potato) Length
of cuttings is 3-5” with leaves
Leaf cuttings
Leaf blades are utilized in starting a new plant. Adventitious roots and an adventitious
shoot form at the base of leaf. (eg.) Sansevieria, Begonia & Bryophyllum
Leaf bud cuttings
They consist of a leaf blade, petiole and a very short piece of stem with attached axillary
bud.
This type of cuttings will be very useful in species which have a tendency to produce root
from the leaf, stem or petiole but do not produce a shoot system out of any one of the
three parts. In this case, the axillary bud serves as a source for new shoot system.
Eg.Lemon, Rhododendron.
Root cuttings
Root piece of 2-4” length are planted horizontally at 1” to 2” depth. Eg: Bread fruit, Crab
apple, Black berry, Rasp berry
Anatomical and physiological basis of rooting
The formation of adventitious roots in cuttings or layering can be divided into two
phases. One is initiation which is characterized by cell division and the differentiation of
certain cells into root initials and then into recognizable root primordia. The second
phase is the growth and emergence of new roots, by a combination of cell division and
cell elongation including rupturing of other stem tissues and formation of vascular
connections with the conducting tissue of the cutting

These root initials are formed adjacent to vascular tissue. In herbaceous plants which
lack a cambium, the root initials are formed near the vascular bundle close to the phloem.
In woody perennials, the adventitious roots in stem cuttings usually originate in the
young, secondary phloem although they may also arise from other tissues such as
vascular rays, cambium or piths.
In some plants, adventitious root initials form during early stage of intact stem
development and are already present at the time of preparation of cuttings. These are
termed ‘preformed’ of latent root initials. These generally lie dormant until the stems
are made into cuttings and placed under environmental conditions favourable for further
development and emergence of the primordial as adventitious roots. Willow, Hydrangea,
Poplar, jasmines, Citrons are some of the species which produce preformed root initials.
The position of origin of these preformed root initials is same as that of other
adventitious roots. After elaborate studies with easy and difficult to root plants, some
insight into the physiological basis of rooting has been established. The important aspects
are summarized below:
1. Auxin level is closely associated with adventitious rooting of stem cuttings.
2. Nutritional status of plants especially high carbohydrate levels with optimum N is

associated with vigourous root growth.
3. Few organic compounds interact with auxin to affect rooting and they are called
rooting co-factors
II. Principles and methods of propagation by layering
It is a propagation method by which adventitious roots are caused to form on a stem

while it is still attached to the parent plant. The rooted stems are then detached and
established in a medium to become a new plant growing on its own roots.
Types of layering
a) Air layering or Gootee
b) Ground layering
1) Simple layering
2) Compound layering
3) Mound (Stool) layering
Air layering
In air layering, roots form on aerial part of plants where the

stem has been girdled and covered with rooting medium. It
should be done during humid months because root initiation
will be high under high humid conditions

Steps:
1. The branch selected should be of pencil thickness
2. The stem should be girdled for about a length of 1cm to 1” to induce adventitious
root formation above the cut. It should be given at 12-15” from the tip of the
branch
3. A ball of slightly damp sphagnum moss is placed around the girdled stem.
4. A wrap of polythene film is placed around the sphagnum moss and tied airtight on
both ends.
Time of removal
It is better determined by observing root formation through the transparent film. In

some plants, rooting occurs in two or three months. Layering made in spring or early
summer is the best and it will give high percentage of success.
The rooted layers should be potted in a suitable container and placed under cool humid
conditions as a hardening process before it is used for planting.
Ground layering
1. Simple layering: Branches that have formed roots in one area only are called simple
layers. Such layers are made by bending the branches to the ground and covering the
portion with soil.
This should be done in early spring for temperate species before growth has started. For
other tropical species an actively growing period is selected. The tip of the shoot is left
exposed to carry out normal process of the plant.
Procedure
1. A healthy shoot of pencil thickness from a lower branch near the ground level has
to be selected.
2. The common practice is to injure the portion

to be covered, by notching, girdling, cutting
or twisting. This practice destroys the
phloem tissue partially or completely and
retards the downward movement of food
material as well as hormones manufactured
by leaves. Injury is given 6-12” back from
the tip
3. The bent part of shoot is inserted into the soil
4. The usual time for layering depends on species eg. for temperate species, it is
done in early spring and for this, dormant, one year old shoots are used.

5. The rooted layers may be removed from the parent plant and kept under cool
humid conditions for curing.
2. Compound or serpentine layering: It is essentially the

same as simple layering, except that the branch is
alternately covered and exposed along its length. So
that, the roots strike wherever the plant is covered by
soil
3. Mound layering (or) stooling: Here, the plant is

pruned close to the ground level and all the branches are
covered with soil. Striking of roots takes place at a number of
places and the plant also produce new shoot system which
come out of the mound. Each shoot with part of roots formed
will be separated and planted in pots for further establishment.
Apple rootstocks are propagated by this method.
The anatomical development of roots
Stem cuttings: Propagation through cutting/layering is common

in dicotyledonous plants. However, cuttings of some
monocots, such as asparagus can be rooted under proper
conditions
Process of root initiation in stem: It is divided into three stages
1. Cellular differentiation of cambium leading to initiation of meristematic cells.

Proliferation of certain cells to form root initials near vascular bundle.
2. These differentiated cells group into recognizable root primoridia
3. The growth and emergence of new roots.
Initiation of root primordia in herbaceous plants
1. Origin is usually just outside and between the vascular bundles (from cmbium)
2. Small group of cells, the root initials, continue dividing, forming groups of many
small cells which develop into root primordial (it looks like root tip)
3. A vascular system develops in the new root and becomes connected with adjacent
vascular bundle
Initiation of roots in woody plants: Origin is in the young secondary phloem, sometimes
from vascular rays or cambium. The time at which root initials develop after cuttings are
placed in the propagating bed varies widely
Callus: After stem cuttings have been made and placed under favourable environmental
conditions, callus will usually develop at the basal end of cuttings. This is an irregular
mass of undifferentiated parenchyma cells. It was believed that callus formation would

be essential for rooting. In most cases, formation of callus and formation of roots are
independent of each other and if they occur simultaneous it is due to their dependence
upon similar internal and environmental condition.
III. Principles of grafting and budding
It is the process of operation of inserting a part of one plant into another or placing it
upon another in such a way that an union will be formed and the combination will
continue to grow as one plant. The part of graft combination which is to become the
upper portion if termed as the ‘scion’ (ion) and the part which is to become the lower
portion or root is termed as ‘root stock’ or ‘understock’ or the ‘stock’ Rootstocks are
commonly grown from seeds, cuttings or layers. All methods of joining plants are
popularly termed as ‘grafting’ but when the scion part is only a small piece of bark (and
sometimes wood) containing a single bud, the operation is termed as ‘budding’.
Reasons for grafting and budding
1. When other methods of asexual propagation are not successful in perpetuating a

clone, eg: Mango and Sapota can be successfully propagated on commercial scale
by grafting only.
2. Plants propagated on their own roots may be weak, susceptible to pests and
diseases, or to any adverse environmental conditions may not adaptable to a
particular soil or climate. For many plant species, rootstocks are available which
tolerate all the above cases and hence they may be exploited as a rootstock
through grafting or budding.
3. For converting poor trees into more desirable one by top-working
4. For overcoming pollination problems, self-fertile varieties may be grafted over

self-sterile trees
5. For fancy purposes, different types of scion may be grafted in the same plant
6. To modify the growth of the plant as dwarf one by employing suitable dwarfing
rootstocks
7. Occasionally the roots, truck or large limbs of trees are severely damaged by
winter injury, cultivation implements, certain diseases or rodent. But use of
bridge grafting or in arching such damage can be repaired and the tree saved.
Limitation of grafting or budding
One of the requirements for a successful graft union is the close matching of the callus-
producing tissues near the cambial layers. Grafting is generally confined to dicotyledons.
These plants have a vascular cambial layer existing as a continuous tissue between the
xylem and phloem. For grafting, it should be borne in mind that the plants to be
combined are capable of uniting. Generally, the more closely the plants to be grafted are

related botanically, the more favourable is the chances of the graft union being
successful.
1. Intra-varietal grafting: When a scion can be grafted back on the same plant or a
scion from a plant of a given clone can be grafted to any other plant of the same
clone eg. Elberta peach on Elberta peach
2. Inter-varital grafting: when different varieties of a species are employed as graft

parents eg. mango
3. Inter-specific grafting: In this case, grafting between the species of the same
genus is done. But this is usually difficult but widely used between species in the
genus citrus. Japanese plum (Prunus salicina) is grafted commercially on peach
(Prunus persica)
4. Intergeneric grafting: when the plants to be grafted together are in different

genera but in the same family the chanes of union are more remote. But successful
union has been reported in the following cases:
Citrus spp. on trifoliate orange (Poncirus trifoliatat)
Sathugud (Citrus sinensis) on wood apple (Ferronia elephantum)
Sapota (Achras sapota) on pala (Manilkara hexandra)
Stock-scion relationships
A grafted or budded plant can produce unusual growth patterns which may be different
from what would have occurred if each component part of a graftage viz., rootstock and
scion was grown separately or when it is grafted or budded in other types of rootstocks.
Some of these have major horticultural value. this varying aspect of rootstocks in the
performance of a scion cultivar or vice versa is known as stock-scion relationship
A) Effect of stocks on scion cultivars
1. Size and growth habit: In apple, rootstocks, can be classified as dwarf, semi-
dwarf, vigorous and very vigorous rootstocks based on their effect on a scion
cultivar. If a scion is grafted on dwarf rootstocks eg. Malling IX, the scion grows
less vigorously and remain dwarf only. On the other hand if the same scion is
grafted on a very vigorous rootstock eg. Malling II the scion grows very
vigorously,. In citrus, trifoliate orange is considered to be the most dwarfing
rootstock for grapefruit and sweet oranges. On the other hand, in mango, all
plants of a given variety are known to have the same characteristic canopy shape
of the variety despite the rootstocks being of seedling origin. But recently,
rootstocks of Kalapade, Olour have been found to impart dwarfness in the scion
cultivars of mango. Guava cultivars grafted on Psidium pumilum are found to be
dwarf in stature.

2. Precocity in flowering and fruiting : The time taken from planting to fruiting i.e.,
precocity is influenced by rootstocks. Generally fruiting precocity is associated
with dwarfing rootstocks and slowness to start rootstocks are precocious than
those grafted on sweet orange or sour orange or acid lime rootstocks
3. Fruitset and yield: The rootstocks directly influence on the production of flower
and setting fruits in oriental Persimmon (Diospyrous kaki cv. Hichiya). When it is
grafted on D.lotus, it produces more flowers but few only mature but when D.kaki
is used as the rootstock, the fruitset is more. the influence of rootstock on the
yield performance of cultivar has been well documented in many fruit crops. Acid
limes budded on rough lemon register nearly 70 percent increased yield than
those budded on troyer citrange, Rangpur lime or its own rootstock. Sweet
orange var. Sathugudi budded on Kichili rootstock gave higher yield that on
Jambhari or on its own seedling.
4. Fruit size quality: Sathugudi sweet oranges grafted on Gajanimma rootstocks

produced large but poor quality fruits while on its own roots they produced fruits
with high juice content and quality. The physiological disorder ‘granulation’ in
sweet orange is very low if grafted on Cleopartra mandarin seedlings, on the
other hand rough lemon seedlings stocks induced maximum granulation. the
physiological disorder black end in Bartlett Pear did not appear if Pyrus communis
was used as the rootstock. When P.pyrifolia was used as the rootstock, this
disorder appeared, affecting fruit quality.
5. Nutrient status of scion: Roost stocks do influence the nutrient status of scion
also. Sathugudi orange trees have a better nutrient status of alnutrients in the
leaves when it is budded on C.volkarimariana rootstock than on its own rootstock
or Cleopatra mandarin stocks
6. Winter hardiness: Young grape fruit trees on Rangpur lime withstand winter
injury better than on rough lemon or sour orange. Sweet oranges and mandarins
on trifoliate stocks were more cold hardy.
7. Disease resistance: In citrus considerable variability exists among the rootstocks

in their response to diseases and nematodes. For instance, rough lemon rootstock
is tolerant to tristesa, xyloporosis and execortis but is susceptible to gummosis
and nematode. On the other hand, treyer citrange is tolerant to gummosis but
susceptible to execortis virus disease. Similarly, guava varieties grafted on
Chinese guava (Psidium friedrichsthalianum) resist wilt diseases and nematodes
8. Ability to resist soil adverse conditions: Among the citrus rootstocks, trifoliate
orange exhibits poor ability, while sweet oranges, sour orange, Rangpur lime
rootstocks exhibit moderate ability to resist excess salts in the soil. Im pome
fruits, similarly, ariation exists among rootstocks to resist excess soil moisture or
excess boron in the soilyrobolan plum rootstocks generally tolerate excess boron
and moisture than Mananna plum root or other rootstocks viz., peach, apricot or
almond.

B)Effect of scion on rootstock
1. Vigour of the rootstocks: In apple, it has been found that if apple seedlings were
budded with the ‘Red Astrochan’ apple. The rootstock produced a very fibrous
root system with few tap roots. On the other hand, if scion ‘Golden burg’ was
budded on the seedlings, they produced two or three pronged deep roots without
fibrous root system. In citrus, if the scion cultivar is less vigorous than the
rootstock cultivar the rate of growth and the ultimate size of the tree is more
determined by the scion rather than the rootstocks.
2. Cold hardiness of the rootstock: Cold hardiness of citrus roots is affected by the
scion cultivar. Sour orange seedlings budded to ‘Eureka’ lemon suffered much
more from winter injury than the unbudded seedlings.
3. Precocity in flowering: Young mango rootstock seedlings (6 months to one year

old) were found to put forth inflorescence when the branches from old trees are
inarched which can be attributed to the influence of scion on the rootstock.
IV. Propagation by specialised plant parts
Certain plants posses’ specialized vegetative structure whose primary functions are
storage of food and vegetative reproduction. If such structures are naturally detachable
for propagation, this procedure is termed as ‘separation’ On the other hand such
structure are to be can into sections for ht purpose of propagation, then this process is
called as ‘division’ the following specialized vegetative structure are used in propagation
1. Bulb: A bulb is a specialized underground organ consisting of a short, fleshy, usually

vertical stem axis bearing at its apex a growing point or a flower primondium enclosed by
thick flesh scales. The outer bulb scales are generally fleshy and contain reserve food
materials whereas the scales towards the inner contain relatively less food materials an
dare more leaf-like. Bulbs possessing dry and membranous outer scales are tunicate
bulbs and bulbs which lack this cover is non-tunicate
2. Corm: A corm is the swollen base of a stem axis enclosed by the dry, scale-like leaves.
It is solid stem structure with distinct nodes and internodes. the propagation of cormous
plants is principally by the natural increase of new corms. the development of miniature
corms between the old and the new corms is termed cormels.
3. Tuber: A tuber is modified stem structure which develops below ground as a result of th
swelling of the subapical portion of a stolon and sub-sequent accumulation of reserve
materials. A tuber has all the parts of a typical stem. Certain plants produce aerial tubers
in the axils of leaves which are known as tubercles.
4. Tuberous roots and stem: In certain plants, the adventitious roots become thickened and
they do have external and internal structures of roots nodes and internodes. These are
known as tuberous roots. In other plants such as tuberous Begonia, Cyclamen or
Gloxinia, they have thickened structures which have arisen from enlarged hypocotyls

tissue. They have a vertical arrangement and may show features of stems. Propagation
of platn with such tuberous roots or stem consists of division of such materials.
5. Rhizome: It refers to a specialized stem structure in which the main axis of the plant
grows horizontally at or just below the ground surface. A rhizome consists of nodes and
internodes having leaf scars on the node. In determinate types of rhizomes each clump
ends in a flowering stalk and growth continues only from lateral branches.
6. Runner: Runner is a specialized stem which develops from the axis of a leaf at the
crown of a plant and grows horizontally along the ground and forms a new plant at one
of the nodes
7. Offset: It refers to special types of lateral shoot or branch which develops from the main
stem in certain plants and is characterized by shortened, thickened stem of rosette-like
appearance. Offsets which produce sufficient roots can be removed by cutting them close
to the main stem with a sharp knife and used for propagation
8. Suckers: A sucker is a shoot which arises on a plant from below ground usually from an
adventitious bud on a root. Suckers are further known as root suckers, ground suckers
and shoot suckers if they arise respectively from root, near the ground and stem of the
plant
9. Crown: The term crown designates that part of a plant at the surface of the ground from
which new shoots are produced. This kind of crown is observed in herbaceous perennials
like strawberry, pyrethrum, Gerbera or African violet wherein the stem is a short and
thickened structure from which the leaves are produced in a rosette like arrangement
Certain plants do have one or more of the above mentioned specialized structures useful
for propagation. But particular structure is preferred for commercial propagation for
obvious reasons. Strawberry can be propagated both by runners and splits from crown.
V. Principles of Micropropagation and Its Advantages
Micro propagation or in vitro propagation refers to the development of new plant in an

artificial medium under aseptic conditions from very small pieces of plant, such as
embryos, seeds, stems, shoot tips, root tips, callus, single cells and pollen grains. This
technique has been put into various applications in the discipline of agriculture,
horticulture and forestry ever. The various applications of micro propagation are as follows
1. Rapid rate of multiplication of a plant clonally.
2. Production of disease-free and disease resistant plants.
3. Induction of mutant and selection of mutants.
4. Production of haploids through anther culture
5. Wide hybridization through excised embryo and ovule culture
6. Somatic hybrids and cybrids through protoplast fusion

7. Transformation through uptake of foreign genome
8. Nitrogen fixation
9. Cryopreservation of germplasm types
Requirements for micro propagation
1. Laminar air flower chamber – It is useful to perform all operation in aseptic

culture
2. Auto-clave or pressure cooker – It is used to sterilize the media, containers,

petridishes and the various accessories required in the transfer operation.
3. Alcohol lamps, disinfectant and sterile water are also required
4. Culture medium – A medium consists of mineral salts, carbon and energy source,
vitamins, plant growth regulators and other organic components
Procedure for micro propagation
1. Collection of explant: The small piece of plant used to begin a culture is referred
to as an explant. The size, age and type of explant affect the success of in vitro
propagation.
2. Surface sterilization: Explants so collection from field grown plants harbor

numberous fungi and bacteria, which when inoculated into a nutrient medium
contaminates the entire in vitro system. Hence, surface sterilization is resorted to
prior inoculation of explants. The efficacy of the sterilants used are found to vary
depending upon the type of chemical, concentration used, time of exposure etc., A
few drops of teepol are also added to facilitate better contact between the explants
and the sterilant.
3. Inoculation: Transfer of the explant into the culture medium is known as

inoculation. This must be done in an aseptic condition. This is achieved by
surface sterilization of the working table of the laminar air flower chamber with
absolute alcohol followed by UV light for 30 minutes.
4. Sub-culturing: After inoculation, the explant increases in volume or it proliferate.

At this stage, it is divided into different components or parts and transferred into
a fresh medium under above mentioned aseptic sterile condition. This process is
known under above mentioned aseptic sterile condition. This sub cultured mass
should produce a shoot and root system which is dependent upon the type of
growth regulator and its concentration used in the medium. It is generally
observed that if the concentration of cytokinins is high relative to auxin in a
medium, shoots are induced and on the hand, when the concentration of cytokines
is low to auxin, roots are induced and at intermediate concentration, the tissue
grows as undifferentiated callus.

Various methods of culturing plant tissues and organs: There are five classes of plant tissues
culture:
1. Callus culture: A piece of sterile plant tissue with living cells is transferred to a
culture medium to induce callus proliferation. Subculturing is then done onto a
medium with or without altered growth regulator concentration, ultimately
resulting in the induction of adventitious organs or embryos.
2. Cell culture: Cells are maintained in suspension cultures so as to produce free cells
and are then subcultured to regenerate complete plants from single cells. This
technique is now useful to induce variability in plant cells and slowly exposed to
select desirable cell variants and regenerate complete plants from these variants.
3. Meristem culture: This technique involves aseptic culture of shoot meristems on

nutrient medium so as to produce complete plants. Most important application of
meristem culture is the production of virus free plant from these variants.
4. Embryo culture: involves aseptic excision of the embryo and its transfer to a
suitable medium for development under optimum culture conditions. After the
embryo has grown into a plantlet in vitro, it is transferred to sterile soild or
vermiculite and grown to maturity in a green house
5. Protoplast culture: From different sources, protoplasts, the plant cells without any
rigid cellulose wall but with plasma membrane only, is allowed to fuse to form a
somatic hybrid. These are cultured in suitable media to regenerate the cell wall
and are again cultured in suitable medium for differentiation and morphogenesis.
Hardening
The plant lets developed in the culture tubes are acclimatized to a specific environment
having a high humidity, a low light level and a constant temperature. Besides, the roots
developed in vitro are hairless and hence delicate, requiring care during transfer from
culture medium. To have better survival rate, the plantlets may be transferred to
container kept in mist chamber where relative humidity is maintained at higher order.
Once new growth is seen, the plants may be slowly transferred to outside by exposing to
increased light intensity in stages.

Chapter-3: [From Cell biology):
Chromosome
Introduction
The chromosomes are the bearers of hereditary material and vehicles of transmission
from cell to cell and generation to generation. The genes which are the unit basis of
heredity are located inside the chromosomes. Each chromosome is thus a giant complex
molecule made up of smaller but less complex molecules - the genes which are arranged
in linear sequence in the chromosome thread, inside the nucleus.
The presence of chromosome was first demonstrated by Strasburger (1875) and coined as
chromosome (chroma–colour; soma–body) by Waldeyer (1888) due to their marked affinity
for basic dyes. The “chromosome” was observed by Baranetsky as the principal
component in the nucleus and responsible for maintenance of hereditary stability.
In the lower organisms, in the prokaryotes with no distinct nuclear membrane, the
chromosome is represented in the DNA alone. This structure is referred to as genophore,
(sensu stricto), as compared to the complex organized structure – the chromosome, in
higher forms of life or eukaryotes. The structure of genophore being at the level of
molecular dimension is beyond the resolution of light microscope and can be analyzed at
the ultra-structural level under electron microscope. The chromosomes of higher
organisms on the other hand, can be observed under light microscope and are made up of
nucleic acid and proteins – both basic and non–basic.
Chromosome research has been greatly enriched through physical, chemical and
molecular studies, coupled with basic cytological and genetic analysis. Such a synthetic
approach has given a deeper understanding of structure and behaviour of chromosomes
and modality of the genetic control on every aspect of cell metabolism.
Number and Size
The number and size of chromosomes remain constant for a particular species. Each and
every cell of an organism contains the same number and types of chromsomes which
enables each gene to exert the influence. The constancy in the number throughout the
body is maintained by a type of cell division otherwise termed as mitosis which involves
equational separation of chromsomes. Each daughter cell contains chromosome
complements of both the parents and is designated as diploid. The halving of
chromosome number is achieved through another type of cell division, otherwise termed
as meiosis or reduction division which leads to the formation of cells with half the
chromosome number in resulting gametes–both male and female and are designated as
haploid. The original diploid number is restored following sexual re-production, which
involves the union of both male and female gametes, leading to fertilization. The
reduction division or halving of chromosome number leading from diploidy to haploidy is
accompanied by pairing and inter-change of segments between paternal and maternal
chromosomes (homologous chromosome pair).

Though the number of chromosomes remains constant for a particular species, it varies
from species to species. The lowest and highest chromosome number recorded in plants
are Haplopappus gracilis (2n-4) of Compositae and Ophioglossum (2n=>1200) of
Pteriodophyta respectively.
The size of chromosomes as studied from mitotic metaphase may be as short as 0.25 μ in
fungi to as long as 30 μ in Trillium; the breadth ranging from 0.2 μ to 2 μ.
Structure and Morphology of Chromosome
Chromosome is the principal component of the nucleus of higher plants. It contains

genes in linear order, responsible for maintaining hereditary stability. In the cell, the
chromosomes appear as coiled threads in the early part of nuclear division, which become
condensed at later stage.
Chromonema and Chromomere: The basic structure of chromosome sensu lato, is a

thread like structure termed “chromonema” by earlier authors. The chromonema thread,
on the other hand, is beaded in appearance, the beads being termed as chromomeres. The
understanding of such beaded thread like structure of chromosomes as visualized by
earlier authors, has undergone much refinement in later years.
The present concept of chromosome structure is regarded as fibriller, made up of fibrils,

folded several times to give a diameter of 10 mμ thickness. The structure appears a coiled
at the beginning of cell division which undergoes spiralization and condensation upto a
stage, followed by successive de-spiralization and de-condensation (Fig. 3.1). It is a
continuous DNA–protein fibre in which condensed and de-condensed segments
alternate. The single condensed segment of a chromosome of higher organism is
comparable to the entire DNA thread of a microbe. The chromonema and chromonemic
component of the chromosome can, however, be resolved under the light microscope at
certain stages of division, such as prophase, where the structure is not fully spiralized.
The chromonema represents coiled thread and is the permanent component of
chromosome. The chromomere represents a higher order of compaction and contain
submicroscopic beads, the nucelosome which can be resolved only at the ultra structural
level.

Chromatid: At the light microscopic level, observation of chromosome is dependent on
the use of basic dyes. The chromosome structure is analyzed principally from the
metaphase i.e. the mid phase of cell division where the chromosomes are highly
condensed and spiralized and as such responds strongly to basic dyes. Each chromosome
at metaphase is made up of two symmetrical chromatids which are held together by
centromere and become separated at anaphase.
Constriction-primary and secondary: By the usual methods of fixation and staining, the
chromosomes at the dividing stages appear as somewhat cylindrical bodies constricted at
one region or more along their lengths (Fig. 3.2). The position of these constrictions are
constant characteristics of any particular chromosome of the complement. They are
frequently referred to as primary and secondary constrictions, the former representing a
region of the chromosome involved in movement during division, while the latter occurs
only in certain chromosomes being associated with the production of nucleoli (nucleolar
zone). The primary constriction referred to as “centromere” or “kinetochore” is obviously
an essential part, since chromosomes which lack the centromere cannot move to the
poles during mitosis.

Centromere: The centromere or the kinetochore is a specialized region of the chromosome
associated with its movement along the spindle to the two poles during anaphase. It may
be localized on a particular part of the chromosome – the primary constriction region or
it may be diffused in which the entire chromosome expresses the property of spindle
attachment. The centromere forms: (i) final point of adhesion for the sister chromatids
during metaphase to anaphase transition, (ii) attachment point for mitotic spindle fibres,
as well as, (iii) site for motor protein to mediate chromosome movement.
The localized centromere appears basically as small distained gap during metaphase. This
appearance is due to the presence of two blocks of heterochromatin at the two sides of
the centromeric apparatus, which exhibit allocycly. They are distained at metaphase and
stained at inter-phase to form brightly stained bodies or pro-chromosomes in the inter-
phase nucleus.
The position of the centromere has been taken as the criterion for designating
chromosomes in different categories. Based on the relative position of the centromere,
chromosomes are described as metacentric, submetacentric, acro-centric, and telocentric
(Fig. 3.3). If the centromere is towards the middle, dividing the chromosome in two equal
arms, the chromosome is known as metacentric. Similarly submetacentric and acro-
centric are terms used to designate chromosome having constrictions at the sub-median
position or towards the end respectively. If the centromere is located at the end of the
arm, it is known as telocentric. On the basis of position of centromere, the chromosomes
look ‘V’ shaped, ‘L’ shaped, ‘J’ shaped, ‘I’ shaped respectively during anaphasic
movement (Fig. 3.4).

During a normal division, centromeric chromomeres and threads divide lengthwise so
that an identical replica is transmitted to each daughter chromosome. However, under
certain conditions, transverse division has been observed, referred to as mis-division or
bursting of the centromere by Darlington (1939) in species of Nicandra. The resultant
daughter chromsomes show terminal centromeres.
On the basis of the number of centromere, chromosome may be mono-centric (one

centromere-usual), dicentric (two centromeres –as in wheat), polycentric (many
centromeres–as in Ascaris) or acentric (without centromere–does not survive).
The diffuse centromeres do not show any single attachment point and show parallel
movement of two chromatids throughout the length on the spindle. The diffuse nature of
the centromeres of Luzula has also been confirmed through irradiation of chromosomes,
where the broken fragments behave as chromosomes with centromeres. As such, in this
genus, increase in chromosome number arises through fragmentation.
Electron microscopic studies have given further details of the structure of the localized
centromere. Microtubules are attached to the kinetochores, which are trilaminar plate
structures on either side of centromere constriction. Ultra-structural analysis further
shows that the two chromatids of a metaphase chromosome are held together at the
centromere. Various types of connections are seen between microtubules and chromatin
in plants and animals.
In certain higher plants, such as species of Lilium and Tradescantia, long blocks or
repetitive sequences have been located around the centromeres of several species. In
general the centromeric DNA protein complex is resistant to nuclease digestion in vitro.
Centromeres are shown to suppress meiotic recombination in some systems. Many plants
have clusters of highly repetitive sequences at or near their centromere. In a few plant
species, tandem repeats are being specifically located at or around the centromere.
Different types of proteins too have been localized in the centromere.
Secondary Constriction and Satellite: In addition to the centromere, otherwise known as

primary constriction, which has a role in chromosome movement, some of the
chromosomes possess another constriction, termed secondary constriction. This
constriction, if is located almost at the end of the chromosome arm so that the terminal
segment assumes a dot like appearance, the term satellite is used and the chromosome is
called SAT chromosome. The word satellite, though often widely used, is in a strict sense
is the end segment of the chromosome arm. The satellite thread i.e. continuous thread of
the chromosome representing the connecting link between the so called satellite and the
main chromosome arm is often visible. The secondary constriction or the satellited
segment is responsible for the organization of the nucleolus, hence called Nucleolus
Organizing Region or NOR. As this segment contains genes coding for 18 S and 28 S
ribosomal RNA, it is necessary for ribonucleoprotein metabolism. In addition, the
chromosome in certain cases, may have more than one secondary constriction–known as
supernumerary constrictions.

Telomere: Telomeres are nucleoprotein structures and occur at the ends of the
chromosomes. They are enlarged terminal ends of chromosomes and are heterochromatic
as observed by a different staining cycle. They perform several vital functions including
end protection. Telomeres enable the chromosomes to distinguish between normal
chromosome ends and breaks, so that the cycle can be delayed to repair chromosome
breaks. The protection of the chromosome ends is mediated through a special mechanism
involving telomerase–a reverse transcriptase. This compensates for the inability of DNA
polymerase to replicate chromosome completely. Normal DNA replication mechanism
cannot lead to complete telomere replication as they yield a blunt DNA end and the other
end having 3’ overhang.
The stability and complete replication of the telomeres of higher organisms indicate that
they have certain common features.
Telomeric DNA, comprising the extreme molecular ends of chromosomes, consists of

simple tendemly repeated sequences, characterized by clusters of G residues in one
strand. An overall asymmetric strand composition results in G-rich and complementary
C-rich strands. The 3’ end of each strand of the duplex chromosomal DNA molecule is the
G-rich telomeric strand and it forms a 3’ terminal overhang, 12 to 16 nucleotides in
length, protruding from the duplex (Fig. 3.5). Each species has a characteristic telomeric
repeat sequence. Limited sequence variations are found in some species. However, widely
divergent species can have the same telomeric repeat unit as well.
Polymerization of TTGGGG is achieved by adding into the telomeric oligonucleotide

primer at the 3’ end of a G-rich strand, independent of an exogenously added nucleic acid
template. Each telomerase synthesizes its species –specific G-rich strand sequence (Fig.
3.6).
The model of telomere serves as a molecular clock controlling the life span, The telomeric
DNA in eukaryotic cell is progressively lost in successive generations on account of both
incomplete replication and strand –specific exonuclease activity. In case the loci is not
made up, the telomere continuous to shorten with generation doubling. Thus in absence
of telomeric activity, chromosome shorten at each division.

Chromatin–Heterochromatin and Euchromatin: Chromosomes are principally composed
of two components–Heterochromatin and Euchromatin. Euchromatin constitutes the
principal functional regions of chromosomes. Heterochromatin on the other hand, is
present on both sides of the centromere, at the telomeres, secondary constrictions and as
well as at the intercalary segments.
The term ‘heterochromatin’ was originally derived from heterochromosome or sex

chromosome, differentiating it from autosomes by Heitz. Since sex chromosomes differ
from rest of the chromosomes in their staining behaviour, the chromosome segments
showing staining cycle, similar to sex chromosomes, were called heterochromatic.
However, Darlington attributed to heterochromatin, the property of allocycly. It implies,
positive staining in chromosomes or heteropycnosis during inter-phase and reverse
behaviour in metaphase. However in a number of cases, the chromosomes may be
apparently condensed or de-condensed throughout, as in the secondary constriction
region. As such, all these segments were clubbed under the category of heterochromatin
the common property being, staining behaviour different from rest of the chromosome
segments.
According to Vanderlyn (1949), any segment displaying properties different from those of
euchromatin should be termed heterochromatin. This statement does not necessarily
imply that all heterochromatic regions, including those of primary and secondary regions
are fundamentally alike,
Heterochromatin thus covers a rather heterogeneous assemblage of chromatin

represented by diverse characteristics in different species. The terms applied to
heterochromatic regions earlier were constitutive or inherited and facultative or appearing
during development.
The best example of Facultative heterochromatin manifested during development is

provided by mealy bug, where one set of chromosomes becomes heterochromatinized
during development and other remains enchromatic. In sex chromosomes of mammals

due to dosage compensation, a single X remains active, the other becoming in active or
heterochromatinized. In plants, following cold treatment certain regions of chromosomes
often show negative staining as in species of Fritillaria, Trillium and Paris. Such areas
termed by Darlington as nucleic acid starved areas arising out of environmental changes,
come also under the category of facultative heterochromatin.
The constitutive heterochromatin, on the other hand, remains stable in the chromosomes
and appears condensed during inter phase as noted in a large number of plant species
including such as Vicia faba. It is also characterized by late or early replication of DNA,
different from that of euchromatin, as in case of sex chromosomes. The best example of
constitutive heterochromatin is provided by segments on both sides of the centromere
which remains as single condensed block during metaphase being designated as
prochromosome.
In addition to centromere as discussed above, constitutive heterochromatin is also located

at the telomeric regions, nucleolar organizing regions as well as intercalary positions in
certain organisms. The entire chromosome may also be heterochromatic, such as sex
chromosome in some plants and accessory chromosomes.
It was also presumed that genetically heterochromatin, though representing condensed

state of chromatin, may be made up of genes having small, similar and supplementary
effects, as embodied in the concept of polygenes of Mather. In later years, the discovery
of repeated DNA sequences and their distribution in chromosomes indicated that
heterochromatic segments are made up of repeated DNA sequences, both GC/AT – rich,
the number and condensed state varying in different segments of chromosomes. Some
segments like pro-chromosomes show high homogenous repeats.
No specific qualitative functions have been attributed so far to heterochromatin. However,

such segments may be involved in regulating chromosome metabolism, cytoplasmic
synthesis and non-specific functions at the chromosomal levels which are all categorized
as nucleotypic. Its variations at different taxonomic level make it suitable for the study of
bio-diversity. Genotypes may differ in the nature of heterochromatin.
Regarding the functions, constitutive heterochromatin helps in pairing of homologous

also in Triticale. It also influences the shortening of the duration of cell cycle as noted in
several species growing in stress situations like desert and alpine regions. During
evolution, moreover, there is often a change in total DNA amount to great extent, which
principally confined to the repeated DNA sequences of heterochromatin.
Karyotype and Idiogram
The term Karyotype is used for a group of characteristics that allow the identification of a
particular chromosomal set i.e. the number of chromosomes, relative size, the length of
the chromosome arms, position of centromere, presence of secondary constriction and
size of satellite. The Karyotype when represented diagrammatically showing all the
morphological features of the chromosomes called ideogram.

Chemical Constituents of Chromosome
The knowledge regarding the chemistry of the chromosome goes as far as 1874 when
Miescher, on chemical analysis of nuclei of various animals, reported the presence of a
protein component and another substance-nuclein in Salmon sperm, now known as
nucleic acid. Later, the works of different authors led to the understanding of chemical
structure of chromosomes having the continuous framework of protein and nucleic acid,
latter being responsible for hereditary stability.
Table 3.1 Differences between euchromatin and heterochromatin
Euchromatin Heterochromatin
1. Euchromatin is found in less 1. Heterochromatin is found in more

condensed region. condensed region.
2. The chromatin fibres in this region 2. The chromatin fibres in this region are
are loosely coiled as compared with more tightly folded than euchromatic
heterochromatic regions. regions.
3. It is deeply stained in divisional 3. It is deeply stained in inter phase but

cycle but less stained in inter-phase. less stained in divisional cycle.
4. The region is genetically active and 4. The region is genetically less active,
contains unique DNA. contains repetitive DNA.
5. Euchromatic regions are able to 5. Heterochromatic regions are unable to

synthesise mRNA in vitro. synthesize mRNA in vitro.
6. Euchromatic regions are seen to 6. Heterochromatic regions are seen to

replicate their DNA earlier than replicate their DNA later than euchromatic
heterochromatin. They replicate regions. They replicate during the onset of
during ‘S’ phase. divisional cycle.
7. Due to addition or loss of this region 7. Addition or loss of this region does not
spontaneous mutation is affected. affect spontaneous mutation.
8. This region is not sticky. 8. This region is sticky.
9. The crossover frequency is more in 9. Crossover frequency is less than

euchromatin. euchormatin.
10. It does not show heteropycnosis. 10. It shows heteropycnosis.
11. Euchromatin is less labile than 11. Heterochromatin is more labile than
heterochromatin. euchromatin and is affected by
temperature, sex, age of parents, proximity
to the centro mere.

Deoxyribonucleic acid, the principal component of chromosomes of nucleus, in
combination with protein, forms the nucleoprotein component of the chromosomes. The
initial concept of a chromosome being composed of a skeleton of basic protein
surrounded by deoxyribonucleic acid, has undergone significant changes. It is now
known, that both basic and acidic protein along with deoxyribonucleic acid enter into the
chromosome constitution. The ribonucleic acid too, according to some authors, appears at
certain stages of the cycle, principally in the synaptonemal complex. The presence of both
types of protein as well as DNA has been confirmed, through enzyme digestion studies
utilizing deoxyribonuclease for DNA and trypsin and pepsin for digesting basic and non
basic protein respectively. However, from the studies on animal organs, the term residual
chromosome was adopted which is supposed to be the fundamental skeleton of the
chromosome. This fundamental skeleton which undergoes spiralization during divisional
cycle reveals the presence of acidic proteins and RNA in addition to DNA. It is supposed
that the nucleohistone fraction forms the matrix over the residual chromosomes. The
former can be removed by sodium chloride treatment and fractionation, keeping the
fundamental structure intact. However, such residual structure, claimed to be the
fundamental skeleton, has not been located in all organisms.
Molecular Organization of Chromosome
The chromosomes of higher organism have been shown to be made up of fibrils,

approximately 20-30 A in diameter, folded several times to yield a diameter of 10 μm
thickness. It represents a continuous DNA-protein fibre. Regarding the number of fibrils
along the length of chromosome, multi-stranded concept was proposed by different
scientists (Steffensen, Ris) earlier. Later it has been claimed with experimental evidences
that the single molecule runs along the entire length of the chromosome from telomere
to telomere uninterrupted through the centromeric region. Thus the chromosome
structure has been demonstrated to represent a uninemic skeleton (single stranded
concept). Several models have further explained the manner of association and folding of
DNA with protein in chromosome.
Folded Fibre Model: According to this model proposed by DuPraw, each chromatid of
chromosome consists of a single fibre of 200-500 A in diameter folded both
longitudinally and transversely (Fig. 3.7).
Nucleosome Model: This is the most modern and acceptable model proposed by Roger
Kronberg in 1974.
Nucleosome: Electron microscopy and bio chemical evidences have revealed that the
chromosome (chromatin) structure is beaded in appearance (Fig. 3.8), each bead is
termed nucleosome or Nu-body. Nucleosome is a complex of DNA and histone protein.
There are five histone fractions in the nucleosome assembly, of which four i.e., H2A, H2B,
H3, and H4 enter into the composition of nucleosome and fifth one H1 protein, serves as a
linker between the nucleosome beads. The octamer is surrounded by a DNA molecule
having almost 200 b.p. of which approximately 140 b.p. enter into the coiling of octamers
and the rest base pairs remain associated with the linker H1 molecule.

The structure of nucleosome reveals evolutionary stability as noted even in yeast and
human.
Experimental evidences for Nucleosome: The evidences in support of nucleosome are as

follows:
X-ray diffraction pattern studies: X-ray diffraction of chromatin indicated the presence of a
structure repeating every 10 nm.
Electron microscopic studies: Electron microscopy of ruptured nuclei showed the presence
of a series of spherical particles connected by a final filament – the so called beads on a
string. The beads have a diameter of 7-10 nm but the length of the filaments is variable.
Digestion of Chromatin with Micrococcal Nuclease: It is already stated that the nucleosomes
grossly appear as beads on string. It is obvious that beads are connected by non-beaded
string or linker DNA which holds the nucleosomes. When a small fragment of DNA
containing 4-5 nucleosomes are treated with Micrococcal nuclease. It gradually digests
the linker DNA but nucleosomes remain partially resistant to nuclease action. An analysis
of the size of DNA showed that spacing between the successive nucleosomes was about
200 b.p. DNA is reduced first to 166 b.p. and finally to 146 b.p. and H1 is lost.
Association of Histones: There are only five histones in each repeating unit. The four of
them are H2A, H2B, H3 and H4 and the fifth one is H1.

It was observed that H3 and H4 were present as a tetramer (a pair of dimmers) i.e, (H3)2,
(H4)2. It was shown that the chromosomes of most organisms had equal number of
molecules of H2A, H2B, H3 and H4 and 25 nucleotides of DNA were present per histone
molecule. Since H3 and H4 are present as tetramers, each repeating unit may have one
such tetramer and 2 molecules each of H2A and H2B. Histones will thus form an octamer
(2H2A + 2H2B + 2H3 +2H4) associated with 200 base pairs in each repeating unit. One
molecule of H1 is associated with each repeating unit but absent in the core.
It was also suggested that tetramers make the core of the unit and oligomers determine
the spacing thus giving a flexible structure. H2A and H2B were added as 2 dimers on each
face of the tetramer (H3)2. (H4)2. The (H3)2, (H4)2 tetramer makes the central kernel
associated with 2 independent dimmers H2A–H2B (Fig. 3.9). DNA enter and leave the
same side and H1 sealing off the two turns of DNA.
Association of DNA: On prolonged nuclease digestion, beyond the cleavage between

nucleosomes, the DNA was removed from both the free ends yielding particles containing
only 146 b.p. instead of 200 b.p. This reduced form of nucleosome is called core-particle.
Adjacent to core particles, each with core DNA, are joined with each other through DNA
segment called linker DNA which is removed during prolonged nuclease digestion. The
core DNA has 146 b.p. but linker DNA can have 8 b.p. to 114 b.p.
Three dimensional structures as revealed from X-ray crystallography shows that the
nucleosome core particle is neither spherical, nor perfectly flat, but wedge-shaped, about
11 nm in diameter and 6 nm high. The study of crystals suggested that DNA double helix
is coiled into a large helix or super helix by making two turns around the histones in the
middle of the particle. From the structure of DNA and diameter of core particle, it has
been estimated that 2 super helical turns would have 160 b.p. Since the length of DNA in
the core is only 146 b.p. therefore it makes only 1¾ turns. The particle is therefore thin
on one side accounting for the wedge shape.

Solenoid Model: The 10 nm fibre of nucleosome gets coiled upon itself to form 30 nm
wide helix with 5 to 6 nucleosomes per helix. In this helix successive turns come close
together so that their centre to center distance was about 10 nm. This 30 nm structure
was called solenoid (Fig. 3.10). The H1 proteins helped in folding of the 100 A fibre into
300 A solenoid because when H1 was removed this ordered folding was absent. H1
molecules aggregate by cross linking to form polymers and may thus control the
formation of solenoid.
The packaging of DNA in a chain of nucleosomes is about 5 to 7 times more compact than
free DNA and in a 30 nm solenoid it is packed about 40 fold.
Loops, domains and scaffold: The 30 nm solenoid fibres is then organized into looped
domain structure (Fig. 3.11). Each loop has approximately 50 turns of solenoid. Each loop
is with 85 Kb of DNA and a length of 10-30 μm. These loops surrounded a central core of
scaffold or matrix, made up of non-histone fibrous network of chromosomal proteins.
This scaffold is involved in condensing solenoid fibre into tightly packed metaphase
chromosome (Fig. 3.12).
The scaffold proteins include two abundant proteins–the larger DNA topoisomerase II
and the smaller matrix protein. Both initiation and continued replication of DNA occur in
association with matrix proteins and topoisomerase II binding sites are found on matrix
associated DNA. The binding sites for topoisomerase II are called Scaffold Attachment
Regions (SAR) and matrix proteins bind to Matrix Attachment Regions (MAR). It is
believed that the same sequence of DNA work as MAR in inter-phase nuclei o and as SAR
in metaphase chromosomes.
Order of organization: Thus the chromosome fibre is presumed to be formed by three

levels of coiling (Fig. 3.13).
1. The first level of condensation involves packaging DNA as a super coil into
nucleosomes. This produces the 100 A diameter inter-phase fibre.

2. Second level of condensation involves an additional folding and on super
coiling of 100 A nucleosome to produce the 300 A solenoid.
3. Finally solenoid fibre of 300 A is organized into loops around a central

scaffold to make the tightly packed metaphase chromosome.

Special type of chromosomes
Accessorys Chromosome
Accessory chromosome otherwise termed as supernumerary chromosome or B

chromosome, in its simplest form, can be defined as an extra chromosome that is not
necessary for the individual and not homologous to any of the normal chromosomes
(Muntzing, 1967). These chromosomes differ from the normal ones in their variable
number, smaller size and greater degree of heterochromatinization. They may vary in
number within different tissues of the same individual, between different individuals of
the same population and even between populations of the same species from different
regions. They are usually acro-or telocentric in nature. Submetacentrics and metacentrics
have also been recorded. They do not usually pair with normal chromosomes during
meiosis.

Such chromosomes were first recorded by Longley and Randolph in maize in 1927-1928,
who referred to them as B chromosomes as against the normal number of 2n=20 in a
normal maize plant. These chromosomes were much smaller than the normal
chromosomes and occurred in varying frequencies in different strains of maize. In Allium
stracheyii too, individuals differ in the number of B chromosomes, whereas absolute,
constancy of such accessory chromosomes in all individuals of a population has been
recorded in Tradescantia virginiana.
In plants the number of B chromosomes present may increase or decrease during the
development of the germ line, particularly the archesporium and/or during gametophyte
development. In Trillium, Lilium, Tradescantia and Plantago, univalent B chromosomes
segregate preferentially to the functional megaspore in female meiosis. In certain cases,
increase in B-chromosomes was observed in plants growing under different
environmental conditions e.g. clay soil and dry climates.
B-chromosomes are usually heterochromatic, showing differential staining behaviour–

Tradescantia virginiana, Allium stracheyii and Secale cereal. The centromere, however,
shows irregular behaviour resulting in irregular an aphasic separation and uneven
segregation.
The aberrant anaphasic separation or non-disjunction, which is responsible for the

numerical variation of B-chromosomes, may lead to variations within different
populations of the same species such as in rye, Urginea indica, Poa alpine and Allium
stracheyii. On the other hand, in the genus Festuca, the number of B chromosomes is
always constant.
Meiotic pairing of B chromosomes shows a wide range of variations in plants. It may be

homologous between two similar B chromosomes, without chiasma formation or non-
homologous in exceptional cases with A chromosomes such as in species of Clarkia. A
typical pairing may involve an end to end association between A and B chromosomes as
in Haplopappus spinulosus.
The degree of heterochromatinization may range from the totally heterochromatic B

chromosomes of Anthoxanthium sp. to the totally euchromatic ones of Tradescantia sp.
The amount of repetitive DNA seems to be involved as well.
The common property of accessory chromosomes in their deleterious action or in certain

cases, their lack of activity. B chromosomes have been shown in certain cases to modify
chromosome pairing in hybrids between closely related species, such as in Lolium, wheat
and Festuca.
The origin of accessory chromosomes in rather obscure. They may have been derived
from the ordinary chromosomes sometime in the remote past, starting possibly as
heterochromatic centric fragments.
The concept of B chromosome as being heterochromatic in nature has led to the

understanding that these chromosomes are highly variable in structure and behaviour.
Their nature and the effects of their presence can be explained more plausibly taking into

account the presence of repetitive DNA in large amounts in most of them, which in some
cases, may be shared with the regular chromosomes.
Lamp Brush Chromosome
The oocytes of some amphibians possess giant sized chromosomes. These chromosomes
can sometimes be seen even without a microscope. Their length is maximum during
diplotene of meiotic prophase and in some cases may even reach 1000 μ. During this
stage each chromosome can be distinguished into a central heterochromatic axis and a
large number of paired loops projecting from it. Because of their brush like appearance at
this stage, these chromosomes are known as “lamp brush chromosomes” Autoradiography
has revealed that the loops are sites of RNA synthesis, i.e., represent active chromatin. In
oocytes, diplotene is of comparatively longer duration and during this period large
quantities of messenger RNA are synthesized and stored for use during rapid protein
synthesis at the time of embryo development. A DNA double helix forms the main axis of
the loop. The DNA of the main axis of chromosomes in inactive and, therefore, very much
condensed, but the DNA of loops is active and stretched. When the activity ceases, loop
DNA also condenses and, therefore, the loops disappear. Loop DNA appears thick because
it is covered with non-histone proteins and nascent RNA molecules.
Polytene Chromosome
These also are examples of giant chromosomes. Most of the studies have been done with
salivary gland chromosomes because these chromosomes are the largest. They arise due
to many longitudinal splitting of chromatids and consequent non separation of split
portions. The process of chromosomes duplication without cell division is called
endomitosis. The polytene chromosomes of Drosophila are of special importance because
they have transverse bands on them. Dark staining bands alternate with lightly staining
inter band regions of chromosomes.
Many polytene chromosomes show various types of swellings or puffs in their inter-band
or band regions, specially during growth and differentiation of larvae. Polythene
chromosomes in plants arising out of endomitosis have also been recorded in endosperm,
haustoria and suspensor cells.
Sex Chromosome
In plant system where the male and female plants are distinct from each other, the
chromosomes associated with sex can be located. However, only a few species
(Melandrium, Coccinia) have been shown to have distinct sex chromosomes. Details of
sex chromosomes and sex determination have been dealt with in another chapter.
However the most common sex determining mechanism is XY where X and Y

chromosomes differ from each other. The female plant is represented by two sex
chromosomes XX, in addition to autosomes. Male is represented by a heteromorphic pair
XY, forming a bivalent. Both the chromosomes show heterochromaticity, Y being more
heterochromatic than X–chromosome. Their segregation during meiosis is normal. In the
female, all the gametes contain X, chromosomes whereas in male half the gametes

contain X–chromosome and half the gametes Y–chromosome. Sex chromosomes in
general are late replicating, compared to autosomes and they can be differentiated from
autosomes by staining behaviour.
Summary
Chromosomes are the bearers of hereditary material and vehicles of transmission from
cell to cell and generation to generation. The number and size of chromosomes are
constant for a species but vary from species to species.
Each chromosome is a thread like structure termed chromonema bearing beads, called
chromomere. Morphologically a metaphase chromosome has two symmetrical
chromatids. The primary constriction or centromere helps in movement of chromosomes
to two poles by spindle attachment through microtubules. On the basis of the position of
centromere, a chromosome may be metacentric, submetacentric, aerocentric or
telocentric. The centromere may be localized or diffused in nature. The chromosome may
bear secondary constriction which is also termed as NOR (nucleolus organizing region) as
it is associated with nucleolus organization. Telomeres represent the enlarged terminal
ends of chromosome bearing simple tandemly repeated repeated sequences.
Chromosomes are composed of euchromatin and heterochromatin. These two

components vary in condensation, staining behaviour, replication and function.
Heterochromatin may be constitutive (centromere, telomere, NOR) or facultative (X
chromosome of human female) in nature. Chemically chromosome consists of nucleic
acid (DNA and RNA) and protein (histone and non-histone).
At the ultra-structural and molecular level each eukaryotic chromosome consists of a

giant molecule of 20 A DNA that extends from one end of the chromosome through the
centromere to the other end of the chromosome. In inter-phase, the chromatin with its
characteristic 100 A diameter fibre has beaded structure, each bead in the string called as
nucleosome. Each nucleosome is made up of histone protein octamer (2H2A + 2H2B + 2H3
+ 2H4) which is surrounded (one and three quarter turns) by a DNA molecule of 146 b.p.
Successive nucleosomes are linked through linker DNA associated with H1 protein.
Folding of nuelcosomes produces solenoid of 300 A with 6 nucleosomes per turn. Finally
solenoids are organized into loops around a central scaffold to make the metaphase
chromosome.
Apart from normal chromosomes there are some special chromosomes in some
organisms; accessory or B chromosome (Trillium, Tradescantia, Urginea), Lamp brush
chromosome (oocytes of amphibians). Polytene chromosome (salivary gland chromosome
of Drosophila) and sex chromosome (Metandrium, Coccinia).

Chapter 8: [From Plant Physiology]:
Plant nutrition, absorption,
translocation and metabolism of
nutrients
Introduction
Of the naturally occurring 92 elements of the periodic table, about 20 are essential to
plants. Water and CO2 provide the plant with the elements C, H and O; the remaining
necessary elements are obtained by flowering plants as inorganic mineral ions, mostly
from the soil solution.
Essential elements
An element is classed as essential to a plant if:
1. The plant cannot complete its life cycle without it.
2. There is no other element can substitute for it.
3. The effect of the element must also be direct, i.e. it should not act by promoting
the uptake of another essential element, or by retarding the absorption of a toxic
one.
To test for the essentiality of an element, the test plants must be placed in an
environment totally free from that element. This means growing the plants in a liquid
culture medium of precisely known composition.
In addition to essential elements there are beneficial elements. They are not absolutely
necessary for survival but promote the growth and vigour of plants.
Non-essential elements are also taken up by plants; any element present in the
environment will be absorbed at least in small amounts. For plants grown in the soil,
large amounts of Al and Na are frequently present as these are common in soils. Though
inessential, such elements are not inert. They influence the ionic balance and osmotic
potential of the cells and may affect the uptake of essential ions.
Many non-essential elements are toxic also and their uptake is detrimental to the plants
and to the animals which feed on them.
Macronutrients and Micronutrients
The essential elements are classified as macronutrients and micronutrients.

1. The macronutrients are required in large amounts relative to the micronutrients.
In culture solutions, macronutrients are supplied at 10-3 to 10-2 mol L-1.
2. The micronutrients are required in concentrations as low as 10-7 mol L-1. Most of
the micronutrients become toxic at quite moderate concentrations, that is above
10-4 mol L-1.
Element Symbol Form absorbed
Essential macronutrients
CO2, CO 32- (carbonate), HCO3-

1. Carbon C
(bicarbonate)
2. Hydrogen H H2O
3. Oxygen O O2, H2O, CO2
4. Nitrogen N NO3- (nitrate), NH4+ (ammonium)
5. Sulphur S SO42- (sulphate)
6. Phosphorus P H2PO4-, HPO42-, PO43- (phosphates)
7. Calcium Ca Ca2+
8. Potassium K K+
9. Silicon* Si H4SiO4 (silicic acid)
Essential micronutrients
10. Iron Fe Fe2+ (ferrous)
11. Magnesium Mg Mg2+
12. Manganese Mn Mn2+
13. Copper Cu Cu2+ (cupric)
14. Zinc Zn Zn2+
15. Boron B H3BO3 (boric acid)
16. Nickel Ni Ni2+
17. Cobalt* Co Co2+
18. Molybdenum * Mo MoO42- (molybdate)

Element Symbol Form absorbed
19. Chlorine* Cl Cl- (chloride)
20. Sodium * Na Na+
Beneficial elements
Selenium Se SeO 42- (selenate)
Rubidium Rb Rb+
Strontium Sr Sr2+
Aluminium Al Al3+
Note: An asterisk* marked element has so far been found to be essential only in some
species; of these, silicon, chlorine and sodium are beneficial in numerous other species.
Role of essential elements in plant physiology
Element Role in plant physiology
Carbon
Carbon, hydrogen and oxygen are constituents of all organic
Hydrogen molecules and the majority of organic molecules of living cells.
They are most abundant in the plant biomass.
Oxygen
Nitrogen, too, is a constituent of many cellular molecules, in

particular proteins and nucleic acids, the key macromolecules of
life.
There are many lower molecular weight nitrogenous organic

compounds vital to cell metabolism - vitamins, cofactors,
hormones, the chlorophyll pigments and the phytochrome
photoreceptors.
Flowering plants additionally contain a variety of nitrogenous

Nitrogen
secondary compounds not involved in basic metabolism. These
include alkaloids. Plants also contain numerous non-protein
amino acids, which are not incorporated into normal proteins.
Both the alkaloids and the non-protein amino acids are toxic and
often bitter tasting; and one possible function is protection against
herbivores.
In seeds, non-protein amino acids, with a high proportion of N by

weight, can act as N storage compounds.

Some non-photosynthetic pigments contain N, e.g. betacyanin, the

red pigment of beetroot (Beta vulgaris).
Sulphur performs an important structural role in proteins where

the disulphide bridges -S-S- stabilize tertiary protein structures.
Sulphydryl groups, -SH, are found in the active sites of many
enzymes.
There are also -SH-containing coenzymes, e.g. coenzyme A, whilst

glutathione, again with a -SH group, is an important antioxidant.
Several iron-sulphur proteins, e.g. ferredoxins, occur in the

electron transfer systems of chloroplasts and mitochondria; these
proteins contain clusters of linked S and Fe atoms at their reactive
Sulphur sites.
Membrane sulpholipids are structural molecules which contain a

sulphate group, found in chloroplast thylakoid membranes.
Numerous flowering plants contain pungent secondary S-

containing compounds appreciated as flavours; these are very
common in the Brassicaceae (cabbage family) which includes
mustard (Sinapis alba). Onions (Allium cepa), garlic (Allium sativum)
and related species are also flavoured with S-containing chemicals.
The presence of such compounds may deter some herbivores.
Phosphorus is contained in nucleic acids and also in membrane

phospholipids which make up the bimolecular lipid leaflet of
biological membranes.
Phosphorus
As a component of the adenosine phosphates (ATP, ADP and AMP)
and related nucleotides, the phosphate group is involved in ‘energy
metabolism’.
Calcium contributes to membrane stability in plant cells by its

association with membrane phospholipids, and it is necessary for
the maintenance of the normal permeability of the plasmalemma.
In plants it also contributes to cell wall structure as calcium

Calcium pectate; this is a major component of the middle lamella which
cements adjacent cell walls together.
The Ca2+ ion is extremely important in stimulus perception; one of

the first effects in the chain of reactions set off by a stimulus,
environmental or hormonal.

Ca2+ is also termed a ‘second messenger’.
It is present in cells as the free K+ ion; it does not enter into

organic combination. It is known to be the activator of some
enzymes.
Potassium
K + ions are concerned in turgor control; such cells are stomatal
guard cells and the pulvinar cells (hinge cells) of leaves and
petioles.
Silicon in the form of silica gel, a hydrated oxide of Si, gives the
cell walls of grasses, including cereals, their characteristic rigidity;
Silicon
this is very conspicuous in the dried-out straw. Si is not known to
take part in any biochemical reactions within cells.
Iron and copper are present in the respiratory and photosynthetic

Iron
electron transfer chain cytochromes. They are also needed for
other oxidative enzymes: Fe for catalase and peroxidase, Cu for
ascorbic acid oxidase and polyphenol oxidase; Fe is present in
Copper iron-sulphur proteins, as mentioned in connection with S .
Fe is also necessary for chlorophyll synthesis.
Magnesium Magnesium and manganese activate many dehydrogenases and

phosphate transfer enzymes and are also important in photo-
synthesis, a Mg atom being part of the chlorophyll molecule
Manganese
whereas Mn is present in the O2-evolving complex.
Zinc is an activator for many enzymes. Particularly important in

plants are alcohol dehydrogenase, superoxide dismutase (which
Zinc degrades the highly reactive and dangerous superoxide radicals
formed during certain oxidative and photosynthetic reactions), and
carbonic anhydrase.
Boron is the element for which the physiological role has proved
most difficult to elucidate. Much of the B in the plant is associated
with cell walls where it cross-links cell wall polymers, such as
pectins.
Boron
It also is needed for normal membrane function. In B-deficient
roots, ion uptake capacity deteriorates but when such roots are
supplied with B, recovery is considerable by 20 minutes and
complete within an hour.
Nickel
Nickel is a constituent of the enzyme urease, which hydrolyses

urea; the enzyme is needed for N metabolism in plants.
Cobalt also is needed in minute quantities only, and is known to be

Cobalt needed for symbiotic N2 fixation, which involves the Co-
containing vitamin B12.
Molybdenum is present in the enzyme nitrate reductase, which is

needed to utilize nitrate, the major source of inorganic N for most
plants, and it is needed for symbiotic N2 fixation. It is also part of
Molybdenum
the cofactor (Moco) for aldehyde oxidase, an enzyme involved in
the synthesis of ABA, and in a few other oxidases. The amount
required is extremely small.
Chlorine is required for the O2-evolving system of photosynthesis.

For this it is needed in only micronutrient amounts.
However, the element is taken up by cells in large quantities and

Chlorine
the chloride ion Cl- is the chief inorganic anion in cells, often
accompanying K+, e.g. during K+ fluxes in stomatal guard cells, so
that it is beneficial in much larger amounts than required to fulfil
its essential biochemical role.
Sodium is required for C4 photosynthesis in some C4 species where

it seems to be involved with the conversion of pyruvate to PEP. It is
present in cells as the free Na+ ion and like Cl- is tolerated in
relatively high concentrations.
Sodium
Chemically it is very similar to K and to some extent it can
interchange with that element; e.g. in Commelina benghalensis Na
can replace K in the control of turgor of the stomatal guard cells.
In succulent halophytes, plants which live in saline habitats, Na+
acts as an osmoregulatory ion, with Cl-.
Important deficiency symptoms
Symptoms caused by nutrient deficiencies are generally grouped into five categories: 1)
stunted growth, 2) chlorosis, 3) interveinal chlorosis, 4) purplish-red coloring and 5)
necrosis.
Nitrogen (N)
Function:-Nitrogen is the most critical nutrient for optimum farm production. Since it is
readily leached from the soil, its level in soil is typically low, however the levels required
for optimum crop growth are quite high and thus generous application is typically
required. Nitrogen reduces all aspects of crop and pasture production e.g. growth, yield
and quality.

Symptoms of Deficiency :-Plants which have Nitrogen for only limited growth, may
exhibit chlorosis especially in the older leaves. In severe cases, the leaves first yellow and
then tan as they die. Some plants (tomatoes, maize) may exhibit a purplish colouration of
the stems, petioles and on the underside of their leaves.
Phosphorous (P)
Function:-Phosphorous is an essential element for cell division and growth. It is required

for photosynthesis, sugar and starch production, energy transfer and the movement of
carbohydrates within the plant and reproduction.
Symptoms of Deficiency:-Plants exhibit stunted growth and leaves are often dark green
in colour.Oldest leaves become dark brown as they die. Maturity may be delayed.
Potassium (K)
Function:-Potassium is an essential element for protein, carbohydrate and fat synthesis

and is required for the proper functioning of chlorophyll and other enzymes involved in
photosynthesis,respiration and protein formation.It is essential for cell division, cell
electrolyte balance and for the functioning of plant stomates.
Symptoms of Deficiency:-Crops suffering this deficiency appear scorched around the

edges and surfaces are irregularly chlorotic. In legumes, the chlorotic spots form patterns
around the leaf edges. Cereal grains develop weak stalks and their roots may become
more prone to infection by root rotting organisms.These two factors may result in
collapse of the crop by wind or rain.
Iron (Fe)
Function:-Iron is an essential element required for the synthesis of chlorophyll. It is

involved in the activation of many enzymes used in photosynthesis and respiration.
In animals, iron is a key constituent of haemoglobin, the species that carries oxygen in
the blood.
Iron is relatively immobile and is generally in short supply in alkaline soils.
Symptoms of Deficiency:-Young leaves develop chlorosis in the interveinal areas which

may develop into white leaves with necrotic spots.Stunted growth.
Manganese (Mn)
Function:-Manganese is an essential nutrient for the growth of both plants and animals.
In plants it enhances root growth, disease resistance and the development of fruit. It is
required for the synthesis of chlorophyll and assimilation of nitrate. It is involved in the
activation of many enzymes involved in photosynthesis and respiration.
In animals, manganese is essential for growth, reproduction, skeletal growth and

carbohydrate metabolism.

Symptoms of Deficiency:-Symptoms vary with species; in cereals – grey – white spots,
flecks and stripes may appear in the interveinal areas. In legumes, interveinal chlorosis of
young and middle aged leaves and tissue may rapidly become necrotic. Seed disorders e.g.
“split seed” or “marsh spot” may develop.
Boron (B)
Function:-Boron is an essential element in plant nutrition. It is essential for root tip,

pollen tube and shoot growth and the synthesis of DNA and RNA.
Symptoms of Deficiency:-Leaf blades may be distorted and stems may become brittle and
crack e.g. “stem crack” in celery. Shorter intermodal length, retarded growth or necrosis
of the terminal buds and youngest leaves. Reduction or failure to seed and fruit.
Malformation of fruit.
Zinc (Zn)
Function:-Zinc is an essential nutrient required for the functioning of a large number of

enzymes involved in the growth and reproduction of both plants and animals. It is
required for the synthesis and functioning of chlorophyll, is involved in the plant
hormone system and as a catalyst for the plant growth regulator, auxin.
Symptoms of Deficiency:-In plants, shortened internodes with excessive branching

(resetting) of small, dark green deformed leaves. In cereals and grasses – chlorotic bands
(yellow, red) may appear either across or within the veins. Stunted growth and necrosis
of older leaves.
Copper (Cu)
Function:-Copper is an essential nutrient required for the functioning of a large number

of enzymes involved in the growth and reproduction. It is required for the synthesis and
functioning of chlorophyll, is involved in the plant hormone system and acts as a catalyst
for the plant growth regulator, auxin.
Symptoms of Deficiency:-Young leaves become dark green,twisted and deformed.

Necrotic spots may appear. In grains and grasses, seed production is reduced and seed
heads may be white and empty.
Molybdenum (Mo)
Function:-Molybdenum is an essential element for both plants and animals. In plants,

Molybdenum is required for protein synthesis.It enhances both photosynthesis in plants
and nitrogen fixation in legumes. In animals, Molybdenum is a constituent of several
important enzymes, and plays a role in animal fertility, the estrus cycle, and mammary
anticarcinogenesis.
Symptoms of Deficiency:-In plants, reduced and irregular leaf blade formation,

interveinal mottling and chlorosis around the edges of older leaves. Necrotic spots at leaf

tips and edges, smaller root nodules coloured white or green (not pink), growth
inhibition in legumes.
Cobalt (Co)
Function:-Cobalt is a key constituent of Vitamin B12 and Propionate (themajor source of

energy in ruminants). Cobalt enhances thenitrogen fixing ability of legumes and
improves the efficiency of ruminal digestion.
Symptoms of Deficiency:-Small root nodules on legume species.
Uniformly pale green – yellow leaves,most severe on old leaves. Some crops may develop
red leaves, stems or petioles. Stunted growth – tops may be less leafy. Grain or seed
production may be retarded.
Magnesium (Mg)
Function:-Magnesium is an essential nutrient for the synthesis of chlorophyll. It is

involved with the functioning of several enzymes associated with
photosynthesis,respiration and reproduction.
Symptoms of Deficiency:-Interveinal chlorosis of older leaves.
Mineral absorption from the soil by plant roots
With the exception of C, H and O, which are derived from water and CO2 and are
incorporated by photosynthesis, plants acquire all other elements as inorganic ions.
The ions which serve as sources of the essential elements for flowering plants are listed
in a table earlier. For terrestrial flowering plants the chief source of mineral ions is the
soil. The mineral rock particles of the soil yield ions by weathering which gradually
brings them into solution; ions are also released by the action of microorganisms on dead
organic material. The ion concentration of the soil solution rises as the water content of
the soil falls, but except under very dry conditions the solution is very dilute.
Not all the ions in the soil are totally free in the soil solution. The colloidal matter in the
soil, both inorganic clay particles and organic particles ‘humus’ also serve to retain ions
by adsorption.
The colloidal constituents of the soil usually carry a net negative charge; cations, being
positively charged, are adsorbed to the negatively charged groups on the clay and on the
organic particles. These ions are held at the surface of the soil particles by electrostatic
attraction only loosely and can be exchanged for other cations.
Plants release H+ ions in the soil which can displace many cations from the surface of soil
particles and make them available in the soil solution freely.
There are two mechanisms of increasing H+ concentration is the soil.
1. Active pumping of protons by the root cells

2. Release of CO2 as a result of respiration increases the levels of H2CO3 in the soil
(CO2 + H2O  H2CO3). H2CO3 then dissociates into HCO3— and H+ ions.
When in the soil solution, the levels of H+ ions increase, the essential cations become
available freely which can then be taken by the root cells mostly by active transport.
Essential anions are usually available freely in the soil sap, therefore they can be taken up
directly using a suitable membrane transport method.
The long-distance transport of ions takes place in the xylem concurrently with water
transport.
Just as for water, the ions move by an apoplastic, symplastic or transcellular route
Apoplastic ion movement is partly by diffusion, along with the flow of water moving into
the transpiration stream.
The endodermal Casparian strips are believed to form a barrier to ion movement, as for
water. Here, the ionic movement becomes symplastic.
Once in the root xylem, the mineral ions move up into the aerial parts with the xylary
stream of sap. (Please refer to the earlier coverage on Ascent of Sap).
Role of Mycorrhiza
The word mycorrhiza means ‘fungus-root’. It is the name given to a symbiotic

association between a plant root and a fungus, which in most cases enhances the mineral
nutrient supply of the plant, whilst the fungus benefits from a supply of organic C from
the plant.
Mycorrhizal associations occur in most species of flowering plants in the field and it is
highly beneficial.
In a mycorrhizal association, part of the fungal mycelium is free in the soil, part is closely
associated with roots.
Exchange of nutrients takes place over the large area of contact between the fungus and
the root cells.
The surface area of fungus exposed to the soil is also very large, enabling efficient
mineral absorption.

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Our full course will offer you the following.

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Payment option by online transfer into Evolution’s Account

A/c Name: Evolution Educare Pvt.Ltd.

A/C No: 4988002100000930

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Branch: Dr. Mukherjee Nagar, Delhi – 110009

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Payment option by Demand Draft

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The details of fee and dispatch

Distance Learning Course Number of Course Fee Including

GS – Prelims (IAS & IFoS) 3 Rs. 8,500

GS – For IAS Main – All 4 papers 4 Rs. 14,500

GK – For IFoS Main 3 Rs. 8,500

ENGLISH – For IFoS Main 1 Rs. 4,500

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A) Specialized Farming: Specialization means that the farmers specialize in one

iv. Less equipment and labour are needed.

v. Costly and efficient machinery can be used.

Sample Chapters: AGRICULTURE DISTANCE LEARNING COURSE |Page 8

ii. Business risk is reduced due to a crop failure or unfavourable market

i. Because of varied jobs in diversified farming, a farmer can efficiently

i. Mixed farming helps in the maintenance of soil fertility. Byproducts of crop

ii. Draught animals are used in crop production and transport.

iii. It provides regular income and employment.

Sample Chapters: AGRICULTURE DISTANCE LEARNING COURSE |Page 9

Determinants of Type of Farming

2) Economic Factors: Marketing cost, changes in input and output prices,

1. Capitalist or Estate farming: In what is known as capitalistic or estate or

2. State farming: State farming as the name indicates is managed by the

Sample Chapters: AGRICULTURE DISTANCE LEARNING COURSE |Page 10

3. Collective farming: The name, collective farming implies the collective

4. Peasant farming: This system of farming refers to the type of organization in

5. Co-operative farming: Co-operative farming is a voluntary organization in

The working group on co-operative farming defines a co-operative farming

A Co-operative farming society makes one of the following four forms:

i. Co-operative better farming: These societies are based on individual

Sample Chapters: AGRICULTURE DISTANCE LEARNING COURSE |Page 11

Sample Chapters: AGRICULTURE DISTANCE LEARNING COURSE |Page 12

Weed Control Vs Weeds Management

Principles of Weed Management

Important weed prevention practices are:

1. Use clean crop seeds/ weed free crop seed

Sample Chapters: AGRICULTURE DISTANCE LEARNING COURSE |Page 13

Cultural Methods of Weed Control

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Sample Chapters: AGRICULTURE DISTANCE LEARNING COURSE |Page 15

2. Oldest, effective and economical method

3. No special skill is required in adopting physical methods.

1. More labour is required, and tire some.

3. Usually operations limited by too wet or too dry conditions

3. Spudding: Hand weeding, hand hoeing added by a sharp edged sickle.