10/6/2021

Principle of Horticulture
Course code: HRT 111, Credit hours: 3 (2+1)

PC: Google

Himal Bhusal, M.Sc. Horticulture

Email: [email protected] 9849708265

Course outline
Unit 1. Introduction
Unit 2. Climate in horticultural crop production
Unit 3. Soil in relation to horticultural crops
Unit 4. Establishment of horticultural enterprises
Unit 5. Training and pruning
Unit 6. Flowering and fruiting
Unit 7. Plant growth regulators
Unit 8. Plant growth and development
Unit 9. High-tech horticulture
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Unit 10. Principles and practices of environmentally controlled production
Course contain:
Plant growth and development
• Seed, bud and tree dormancy:
• Germination, growth, development
• Juvenility, maturity, flowering,
• Fruit set, fruit growth,
• Ripening and senescence.

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 Concept of growth and development
Plant growth process:
• The life cycle of a plant start form seed or vegetative propagule to
its maturity and death is a very complex process.
•Plant growth may be defined as an irreversible increase in the size of
cell, organ or whole organism. This occurs by cell division and cell
enlargement.
  Cont.

•Mature form of plants develop form cell differentiation process

which involves specialization cells, tissues development and
maturation.
•Thus, different organs such as stem, leaves, roots, flowers, fruits and
seeds are formed in which growth and development both occurs.

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 Basic concept on which the growth of higher plants
depends are:
Photosynthesis for food supply
Respiration for energy release
Assimilation for utilization of food
Absorption for water and nutrients
Translocation for movement of manufactured food
Other metabolic process

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  Photosynthesis

The most important process that involves biochemical

reaction for manufacturing the food materials by plants.

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 Cont.
Photosynthesis has two phases:
i. Light phase:
This phase is independent of temperature.
It consist of trapping of light energy by
chlorophyll thereby removing electrons from water.
4H2O + light = O2 + 4H+ + 4e- + 2H2O

Photophosphorylation

Photophosphorylation is the process of utilizing light energy from photosynthesis

to convert ADP to ATP. It is the process of synthesizing energy-rich ATP molecules
by transferring the phosphate group into ADP molecule in the presence of light.
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 Cont.
ii. Dark phase (Blackman reaction):
This phase is independent to light but
dependent to temperature.
Dark phase is not the same for all
horticultural crops.

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 Cont.
ii. Dark phase (Blackman reaction):
•There are three CO2 reduction pathways,
each being favored by different environment
to form carbohydrates:
1.C3 pathway (Calvin cycle)
2.C4 pathway (Hatch and Slack cycle)
3.CAM pathway (Crassulaean acid metabolism).

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 Respiration

• It is a biological oxidation (combustion) process through which

energy required to run the machinery of the cell is released.
• Food mainly carbohydrates manufactured during photosynthesis is
used as source of energy, CO2 and water are the end product

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Respiration cont.

• Biological combustion of glucose (carbohydrate) are divided

in to two major parts:
o Glycolysis
o Krebs cycle (TCA cycle or citric acid cycle)
• Several factors like temperature, water level of oxygen and CO2
affects the rate of respiration,
• Respiration rate is higher in meristematic region and lower in
dormant bud or seed.

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 Assimilation:

• The process of utilization of food material for growth.

• The first step here is the conversion of food into simple compounds
through digestion which involves several enzymes.
Example:
• Starch is converted into simpler compound sugar with amylase
enzymes. Certain minerals may also be needed as component of
enzymes. Deficiency of such minerals may adversely affect the
cellular process. The simpler forms are then used as building blocks
for forming more complex compounds needed in the protoplasm.

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 Absorption

• Absorption is a means whereby water and nutrient enter into the plant.
• Cell is a mass of protoplasm surrounded by a differentially
permeable membrane, which allows passage for water and inorganic
salts but not the complex molecule like sugar.
• Molecule move through such membrane by diffusion.
• The movement of water form a solution of higher water potential to
one of lower water potential across a differentially permeable
membrane is called osmosis.

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 Absorption cont.

• Besides this, the cells also act as metabolic pump, not accountable by
diffusion and such a active uptake requires energy that comes form
respiration.
• The movement of molecules through cell membrane also depend on
the size ionic charge and solubility of the molecules.
• Ionic concentration of the nutrient medium also affects the
permeability. Monovalent like K+ Na+ increase permeability whereas
divalent like Mg++ and Ca ++ decrease.

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  Translocation:

• The process of movement of synthesized or absorbed food

and minerals nutrient from one part of plant to another is
called translocation.
• Sugar synthesized in the leaf is translocated mostly as
sucrose from leaf to growing tissues mainly by phloem.

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 Translocation Cont.

• If it is not sufficient, translocation of sugar occurs from storage

organ after hydrolysis of starch.
• Small amount of sugar my move upward through xylem in certain
season.
• Water and minerals absorbed by the roots are translocated upward
primarily through xylem.
• Some lateral movement of minerals may occur form xylem to
phloem.

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 Transpiration:
• Transpiration - water loss by evaporation from the aerial parts of the
plant followed by diffusion of water vapor into the outside
atmosphere.

• Small percentage of the total amount of water absorbed by the plant

is utilized for its vital biochemical process as well as maintenance of
the turgor in the cells.
• The major portion of the water loss occurs through
numerous stomata present in the leaves followed by
lenticels, cuticle and hydathodes.

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 Cont.
• Transpiration rate is higher in the day time than at night. High
temperature and dry and windy condition increase transpiration rate.

• Higher the vapor pressure of water around the plant the lower would
be the rate of transpiration.

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 Other metabolic process:
• The ultimate sources of numerous organic materials found in the cell
or the plant is the outcome of photosynthesis and absorption.

• Metabolism – refers to the sum total of all the biochemical reactions

that takes place in the plant body

• Metabolism = Anabolism + Catabolism

• When anabolism exceeds the catabolism, growth occurs in the plant.

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 Major constituents in plant:
1. Carbohydrates
- Monosaccharides, disaccharides and polysaccharides
2.Proteins
3.Lipids
- Fats, phospholipids, glycolipids, waxes and sterols
4. Non-protein nitrogenous compounds
- Purine, pyrimidine, nucleic acids, porphyrins, alkaloids,
vitamins, coenzymes and hormones
5. Other organic compound
- Terpenoids and steroids, organic acids and alcohol, Aromatic
compounds
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 Plant development:
A. Development at cellular level
• Cell division - Mitosis

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 Plant development:
A. Development at cellular level
• Cell division - Mitosis

• Cell division is followed by cell

enlargement which can be very
extensive.
Example:
o Meristematic cells of shoot apex may be 20 times shorter in length
and 5 times less wider than the palisade parenchyma cell of leaf and
cortex of stem.
o Loosening of cell wall by the plant hormones like auxin, cells may be
enlarged. 22
 Plant development cont.
B. Development of the plant body

• Development of plant body can be divided into two phases:

i. Vegetative phase
ii. Reproductive phase

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  Plant development cont.
i. Vegetative phase of plant development

• This can be divided into following events

o Seed germination
o Seed dormancy,
o Juvenility,
o Bud dormancy and
o Maturity

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  Seed germination:

A seed is a small embryonic plant enclosed in a covering called the seed coat,
usually with some stored food.

CO2
H2O Activate growth
substances and C6H12O6
release enzymes
O2
Growth of Cell division
…..Cont..... radicle and
plumule Cell elongation

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 Different methods of training:
There are two kinds of germination:
1. Epigeal Germination 2. Hypogeal Germination

Example: Bean, Groundnut Example: Pea, Gram

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 Seed dormancy:
• Seed dormancy is a state of growth inactivity in seeds under the
favorable environmental conditions to germinate.
• The cause of dormancy may be either physical or physiological and
can be eliminated through various techniques.
• In a way seed dormancy is a natural protective mechanism against
premature germination.
• For example, prior to winter or dry season, if a recently harvested
seed were to germinate immediately, it may not be able to withstand
the adverse climatic conditions.

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 Cont.
Physical causes of dormancy may be due to:

• May be impermeability of seed to water and oxygen because of

hard and impervious seed coat. e.g. legume seeds
• May be due to mechanical resistance to embryo enlargement e.g.
cherry, peach and raspberry.
• Some hard seed resists water entrance through hilum because of one
way value mechanism.
• In other cases impermeability may be due to impregnation of seed
coats of underlying membranes with waxy materials etc.

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 Cont.
Physical causes of seed dormancy can be overcome by:

• Weathering (alternate freezing and thawing) and /or soil microbial

action till the seed coat is weakened to permit water imbibitions.

• Artificially, the dormancy can be broken through moist storage at

high temperature, mechanical scarification or treatment with acids.

The treatment must not affect the embryo, anyway.

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  Physiological causes of seed dormancy
• Physiological cause of dormancy is presence of germination
inhibitor present in the seed, besides absence of gibberellin(s) or
cytokinin or both.
• A number of chemicals that inhibits germination have been identified
such as coumarin, abscisic acid, phenolics, alkaloids and several
other compounds.

• Germination inhibitors may be present in the flesh of fruit, the seed

coat, the embryo or the endosperm.

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 Physiological causes of seed dormancy :

• Example: dormancy in apple seed is found to be due to the presence of

abscisic acid which inhibits the growth promoting action of gibberellic
acid.

• Germination inhibitors are removed naturally by leaching action of

rain or microbial degradation of artificially through soaking and
washing.

• Some seeds like cherry, peach and apple can be induced to germination if
they are stored under well aerated and low temperature and moist
condition, a treatment referred to as stratification.
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 Double dormancy:

• Seed dormancy caused both by physical and physiological causes is

referred as double dormancy and is the characteristics of several
legumes.

• In such cases, one year is required for weathering or microbial

attack seed coat and the second year is to leach out the inhibitor.

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 Juvenility:
• Juvenility describes the phase of growth following germination from
seed during which flowering does not occur and the bud meristem is
not “competent” to respond to seasonal environmental inductive cues,
and hence remains vegetative
•Physiologically, the juvenile state is a period when the plant is capable
of exponential increase in size, when flowering cannot be readily
induced, and when plant develops characteristics morphological forms
of leaves, stems, thorns etc.

•For example: Juvenile leaves may simple and mature leaves are
compound in bean and citrus.
• https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/juvenility

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 Juvenility :

Seed Embryonic stage Juvenility stage Mature (adult) stage

Plant Juvenile period
Rose 20-30days
Grapes 1 year
Mango 5 year
Tomato, Brinjal 60 days Changing phase
• It is important as it affects when a plant can become reproductive
• Understanding this phenomenon and getting different ideas of accelerating
growth to get early flowering will have great practical significance.
• It affects the plant breeding efficiency, plant propagation, and to the selection of
new cultivars.
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 Juvenility:
• The length of the juvenile phase can vary from a few days to years
depending on the species.
•For example, some herbaceous species remain juvenile few days
(Cauliflower: 2-8 weeks, broccoli: 5 weeks etc) whereas apple
seedlings usually remain vegetative for 7–9 years after seed
germination (Flachowsky et al., 2009).
• Plant hormones – GA and auxin are more likely control the juvenility

Juvenile phase Maturity

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 • Plant development
i. Vegetative phage of plant development

• Seed germination, Seed dormancy, Juvenility, Bud dormancy and

Maturity

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 Bud dormancy:
• Bud dormancy is a suspension of most physiological activity and
growth that can be reactivated.
•It may be a response to environmental conditions such as seasonality
or extreme heat, drought, or cold. The exit from bud dormancy is
marked by the resumed growth of the bud.

•The presence of an inhibitor (mostly abscisic acid) as

well as lock of some promoter(s) probably causes
dormancy.

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 Bud dormancy:
•Bud dormancy is commonly broken through chilling treatment for
a specific period. Example – Peach requires 350-1200 hrs of
below 7ºC temperature depending upon the cultivars to break
dormancy

• Chemicals like ether vapor or a solution of ethylene chlorohydrin can

break dormancy.

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 Maturity

Maturity – a state of plant development, which is capable of

flowering.
• For the expression of flowering potential, environmental condition
should be favorable.
• In mature stage, food, minerals and metabolites are
translocated to reproductive parts.

• This leads to the partial reduction of vegetative

growth, although removal of flowers enhances
vegetative growth.
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 Maturity

• The is evidences that during transition from juvenility to maturity,

the level of auxin and gibberellins are reduced.

• The period of maturity varies depending upon the species and

growth habit of the plants within the species.
Annual, biennial and perennial plant
•Growth habit – determinate and indeterminate

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 Plant development

ii. Reproductive phage of plant development

Flowering, Fruit development, Unfruitfulness and fruit drop,
Fruit maturity and ripening, Tuber and bulb development, and
Senescence

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 Flowering:

Formation of flower primordial is a major event in the

start of reproductive phase of plant development.

A wide spectrum of physiological and morphological

changes occurs in the transformation of a vegetative stem
primordium (apex) to a floral primordium.

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 Flowering:

This take place only after the plant has attained a certain minimum vegetative
growth provided the environmental condition for flowering are within
satisfactory range.

The conical shape in vegetative condition turns to more flattened shape with
small protuberances in a whorl arrangement around the meristem known as
flower bud differentiation.

This is accompanied by a rise in synthesis of DNA and RNA and increased

mitotic activity.

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 Factors determining the onset of flowering:

a. Photoperiodism

The growth response of plant to the length of day i.e. length

of light and dark period is called photoperiodism.

It is evident that the flowering behavior in response to

photoperiod does not depend on the day length alone, but
upon a factor called the critical night length (CNL).

CNL is the night duration of the photoperiod required to

induce flowering.
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 Cont.

• Based on the response to light, plant can be divided in to

three groups:
i. Short day plant:
• They flower with night longer than the critical night length or
initiates flower when the day length is below 12 hours.
Example: Chrysanthemum, coffee, poinsettia etc.
ii. Long day plant:
• They flower with nights shorter than CNL or initiates flower
when day length is exceeds 12 hours.
Example: Beetroot, Radish, Spanish etc.

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 Cont.
Based on the response to light, plant can be divided in to
three groups:

iii. Day neutral or intermediate plant:

• They flower irrespective of the photoperiod.
Example: tomato, broad bean etc.

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 Cont.
Florigen concept:
•Florigen is a systemic signal that initiates flowering in
plants. It is synthesized in leaves and transported to the
shoot apical meristem (SAM) where it promotes floral
transition

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Phytochrome and flowering.

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  Cont.
Vernalization:
There are plants for which flowering is either quantitatively or
qualitatively dependent on exposure to low temperature, this
phenomenon is termed vernalization.
In short, vernalization refers specially to the promotion of
flowering by a period of low temperature.
In general, exposure of plant at temperature -2 to 10ºC (optimum:
2-5ºC) for several weeks (2-8 weeks) depending upon the species is
required.
Example: sugar beet, cauliflower, radish etc
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  Cont.

Devernalization:
The vernalized plants can be reverted back to their original
non-flowering condition by exposure to high temperature a
process called dervernalization.

Example: Onion sets stored in cold storage are exposed to

above 27 degree C for 2-3 weeks before planting they will
form bulb instead of flowers.

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  Cont.

Carbohydrate: Nitrogen ration (C:N ratio):

• It has been established in many plant that if C:N ratio is
optimum, it favors the flowering induction.
• on the other hand, if nitrogen level is more, it will favor
only vegetative growth and reduce or delay flowering.

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  Regulation of flowering and sex expression:

• Several known plant hormones seem to play role in flowering.

Example:
• Gibberellic acid can promote flowering in some long-day
plant or some plants require vernalization (eg. Cabbage, beet
and carrot).
• In contrary, some growth retardant regarded as anti-gibberellin
can promote flowering in some species (eg. Rhododendron).
• Auxin induces flowering in pineapple (at low
concentration). Ethylene- generating compound called
‘ethephon’ replaced it for commercial use.
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  Regulation of flowering and sex expression:

Sex expression:
• GA3 (10-25 ppm), IAA (100 ppm) and NAA (100 ppm)
when sprayed at 2- 4 leaf stage in cucurbits, they have
been found to increase the number of female flowers

• Whereas, GA3 (1500-200 ppm), Silver nittrate (300-400

ppm) and Silver thiosulphate (300-400 ppm) sprayed at 2-4
leaf stage induces male flower production in cucurbits.

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  Fruit development:

A fruit is simply a structure arisen by development of tissue

supporting the ovules.

Such a view is valid even for seedless fruits. Because, ovules are
also initially present in seedless fruits.

Usually, fruit development is considered to start after anthesis. But,

in the real sense, the events occur right after flower induction and set
the pattern of development.

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 Stages in fruit growth and development:

Sigmoid growth curve Double sigmoid growth curve

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  Cont.
Pollination:
- Pollination is the act of transferring pollen grains from the
male anther of a flower to the female stigma.
- Pollination does not allow formation of flower or fruit
abscission layer and provide male gametes for fertilization.
- It has been demonstrate that simply artificial application of
auxin or water extracts of pollen grains result into fruit set.

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  Cont.

Fertilization and post-fertilization developments

• The process of union of male two nuclei of male and
female gametes is called fertilization.
• Double fertilization: In this process, two male gametes
fuse with one female gamete wherein one male gamete
fertilizes the egg to form a zygote, whereas the other fuses
with two polar nuclei to form an endosperm.
Parhtenocarpic fruit – fruits obtained without
fertilization becomes seedless
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  Cont.

Fertilization and post-fertilization developments

• After fertilization, the early stage of fruit growth is

characterized by formation of new cell, but quickly change
to cell enlargement.
• In general, Cytokinins are involved in cell division while
IAA and GA function during cell enlargement phase of
fruit growth and development.

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  Unfruitfulness and fruit drop:

• Unfruitfulness refers to the state where a plant is not capable of flowering and
bearing fruit

‘Fruitfulness’ refers to the state where a plant is not only capable of

flowering and bearing fruit, but also takes these fruits to maturity. The
inability to do so is known as ‘unfruitfulness’ or ‘barrenness’.

Causes of unfruitfulness:
lack of balance between vegetative growth and fruiting
Unfavorable environment for/during flowering and fruit set
Heavy cropping - leading to inhibition of fruit bud production
Sterility- due to impotence, incompatibility or the abortion of
embryo. 59
  Unfruitfulness and fruit drop:

• Fruit drop is the detachment or separation of a fruit from a branch of a tree or

a plant, caused by the formation of a separation of layer of cells (abscission
layer) on the fruit stalk due to a series of physiological and biochemical
events

• It may occur shortly after pollination and fruit set or even at the time
of normal fruit ripening.

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  Unfruitfulness and fruit drop:

Example:
In apple fruit drop occurs twice:
i. Early drop – occurs initial swelling of ovary and start of endosperm
development
ii. June drop – occurs during the period of rapid embryo development
accompanied by heavy fruiting in the tree.

- During the fruit drop auxin level in low

- ABA and ethylene have also been implemented in fruit drop.

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  Fruit maturity and ripening:

Fruit Maturity refers to the processes associated with a fruit reaching

maximum size (and it the time for harvesting).
Ripening refers to the processes which transform the mature fruit
qualitatively so that the fruit may have specific flavor, aroma, color and
consistency with better edible qualities. In fleshy fruit, ripening is accompanied
by softening of flesh.
Fruit vary in ripening behavior. For example: avocado ripening proceeds
only after picking but many fruits ripen while the fruit is still attach to the tree or
may proceed if the fruit picked at maturity.
•Ripening is degenerative process.

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• Paul et al., 2011

 Climacteric and non-climacteric ripening:

Climacteric fruits

Non-climacteric fruits

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• Paul et al., 2011

 Chemical changes during ripening:

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  Tuber and bulb development:
•The development of underground structures like tuber and bulb has several
similarities with reproductive development.

•Morphological differentiation begins to form tuber and bulb like fruits which
will pass on to the ripening process.

•The formation of tuber, corm and bulb is a dramatic expression of the

capacity of higher plants for mobilization of solutes.

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  Tuber and bulb development cont.
•This involves the translocation of nutrients and growth stimulates at the cost of
other plant parts since these underground structure are sinks..

• Tuber formation is very much influenced by day length.

•Example: It is favored by short days as in Jerusalem artichoke and in may wild

species of potato. However, the cultivated European varieties of potato are not
very sensitive to day length.

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  Senescence.

• Senescence refers to such a stage of development in the plant life

that is characterized by deteriorative process leading to the
termination of the functional life of an organism or organ.
• It is an irreversible degenerative phenomenon once the process
sets in and ultimately leads to the death of the plant or plant parts.
Different pattern of Senescence

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  Senescence.

1.Overall Senescence:
This type of senescence occurs in annuals where whole of the plant
is affected and dies.
2.Top Senescence:
This is represented by perennial herbs where senescence occurs
only in the above ground parts, the root system and underground
system remaining viable.
3.Deciduous Senescence:
This type of senescence is less drastic and takes place in woody
deciduous plants. Here senescence occurs in all the leaves
simultaneously but the bulk of the stem and root system remains
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alive.
  Senescence.

4. Progressive Senescence:
This is characterized by gradual progression of senescence and death
of
leaves from the base upwards as the plant grows.

The senescence of the entire plant after a single reproductive cycle

is also known as monocarpic senescence

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Thank You

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