CHAPTER 3

GROWTH, PHENOLOGY AND DEVELOPMENT

What will you learn:
• Growth, development and morphogenesis
• Phenological development and internal and environmental
controls of phenology
• Quantitative Crop Growth Analysis (Individual &
communities) and Growth analysis indices; Relative growth
rate, Net assimilation rate, Leaf Area ratio, Crop growth
rate, Leaf area index,
• Light interception and canopy architecture, leaf angle and
orientation
MORPHOGENESIS

• The origin and

development of plant
form and structure.
• Two primary functions:
• Growth &
• Differentiation (or
development)
PHENOLOGY

• Phenology is the study of the timing of recurring biological

events in the animal and plant world, the causes of their
timing with regards to biotic and abiotic forces, and the
interrelation among phases of the same or different
species.
Phenology - Components

• Phenology is the qualitative and quantitative description

of a plant's life cycle from seed to seed.
• Qualitative aspects: include morphological development and
the partitioning of the life cycle into distinct stages of
development, such as seedling emergence, flowering, and
physiological maturity.
• Quantitative aspects: include rate of development and the
duration of the life cycle.
1. Germination : Defined as a plant with 4 or
fewer leaves (i.e., seedlings)

2. Vegetative: Defined as a plant with 4 or more

leaves that has not flowered or fruited.

3. Flowering : Defined as a plant that has

begun to flower, but does have fully developed
fruits yet.

4. Fruiting: Defined as a plant that has

fully developed fruits.

5. Senescence : Defined as a plant that is completely

dead indicated by a lack of green leaves, fruits, or stems
(i.e., a lack of chlorophyll)

Phenophases
Plant and its Environment
ATMOSPHERE
Weather (current & last
season) and climate – micro-
mesomacro scale local
and, gases, light,

BIOSPHERE INDIVIDUAL
diseases, CHARACTERISTICS
pests, genes, age
competition

SOIL
temperature, nutrients, water,
flora & fauna
Phenology - Importance
1. Seasonal dry matter accumulation is a function of the
duration of the life cycle of annual crops.
2. Rates of physiological processes can vary among phases
of the life cycle. For instances:
▪ Dry matter partitioning to the seeds or fruits occurs during the
final phase of the life cycle of annual crops.
▪ Leaf photosynthesis tends to peak – during fully expanded
stage, declines gradually after full expansion.
3. Most crops are more susceptible to adverse
environmental conditions during one or more phases or
stages of phenological development.
Phenology – Duration

• Duration of the life cycle is a genetic attribute, but the

duration is influenced by environmental conditions - the
life cycle of a crop cultivar/hybrid varies with the location
and year in which it is grown.
BBCH Monograph

• The BBCH-scale is a system for a uniform coding of

phenologically similar growth stages of all mono- and
dicotyledonous plant species.
• The BBCH-scale uses a two-digit code system, which is divided
into principal and secondary growth stages
• Phenological development stages of plants are used in
scientific disciplines (crop physiology, entomology and plant
breeding) and in the agriculture industry (risk assessment of
pesticides, timing of pesticide application, fertilization).
The basic principles of the scale:
• The BBCH-scale provides a framework to develop scales for
individual crops.
• Similar growth stages of each plant species are given the
same BBCH code.
• Each code has a description and important growth stages
have additional drawings included.
• The first digit of the scale refers to the principal growth stage.
• The second digit refers to the secondary growth stage which
corresponds to an ordinal number or percentage value.
• Post harvest or storage treatment is coded as 99.
• Seed treatment before planting is coded as 00.
Organization of the scale

• The entire developmental cycle of the plants is subdivided

into ten clearly recognizable and distinguishable longer-
lasting developmental phases.
• These principal growth stages are described using
numbers from 0 to 9 in ascending order.
Principal growth stages
Stage Description
0 Germination / sprouting / bud development
1 Leaf development (main shoot)
2 Formation of side shoots / tillering
Stem elongation or rosette growth / shoot development (main
3 shoot)
Development of harvestable vegetative plant parts or
4 vegetatively propagated organs / booting (main shoot)
5 Inflorescence emergence (main shoot) / Heading
6 Flowering (main shoot)
7 Development of fruit
8 Ripening or maturity of fruit and seed
9 Senescence beginning of dormancy
Retrieved from: https://www.politicheagricole.it/flex/AppData/WebLive/Agrometeo/MIEPFY800/BBCHengl2001.pdf
 Phenological growth stages and BBCH
Identification keys of rice (Oryza sativa L.) - Lancashire et al., 1991
 Stages Management Considerations:

0 Germination to emergence
Water (food translocation), temperature (germination), air (respiration)
1 Seedling stage Care of seedlings care, (sufficient water, fertilizer ), no stress
2 Tillering Water, nutrient, weed management
3 Stem elongation Variety, planting time, short / long maturation

4 PI to booting Sufficient k-fertilizer, water management ( 5-7cm water depth)

5 Heading Manage drought, it will cause significant yield. Protection from diseases
(blast)
6 Flowering Manage drought , temperature, strong wind and rain
7 Milk grain Manage insects (destructive at this stage)
8 Dough grain Drain water

9 Mature grain Harvest at 80-100% grain maturity or when most of the grains are golden
yellow
Growth Patterns

1. sigmoid growth curve ( organism and population )

2. limited growth curve ( annual plants ) and unlimited
growth curve ( perennial plants )
3. isometric growth ( fish ) and allometric growth (human
organs )
4. intermittent growth curve ( arthropods )
 LIMITED GROWTH CURVE UNLIMITED GROWTH CURVE

• In annual plants, the sigmoid growth curve • In perennial plants, the growth curve for
shows a small reduction in dry mass in the unlimited growth comprises a series of small
initial part of growth, due to seed sigmoid curves.
germination. • This type of curve shows that there is an
• This loss of mass is later replaced when the increase in the mass and size every season.
first leaves develop and start photosynthesis. • In plants that demonstrate unlimited growth,
• Later, growth rate is very rapid, until it there is no fixed rate and the growth
becomes constant at a later stage. Then the continues until enemies, natural catastrophe
rate gradually reduces until it becomes or diseases cause destructive effects on
zero. them.
• e.g.: pea plant • E.g.: woody plants
Plant Growth Analysis

❑Growth - Growth is defined as an irreversible increase in

size of a plant or a plant part. Therefore growth describes
a quantitative change in a plant or a plant part.
❑Development - Development is defined as a qualitative
change occurring in a plant. e.g:-Initiation of leaves, roots
and flowers.
Growth Analysis Method: It describes a
series of techniques to quantify plant growth.

Dry weight (drymass):

Weight of a plant or plant
part after it is oven dried at Leaf area. Total assimilating
800C at least for 48 hours. It area (green, non-green,
is a measurement of the LAI)
actual dry matter present in
a given sample (unit g, kg).
Growth Analysis - Methodology
• Serial harvest: all treatments
have an initial starting point
(T0) with initial weight (Wo)
◼ can be a harvest
◼ can be initial growth
◼ Individual Plant measured

• Plants are harvested at

various times.
◼ weights at different times
are used in subsequent
calculations
Objectives of growth analysis
▪ To compare the growth of different species (Soybean & rice),
varieties (Genotype) of the same species, compare growth of
different plant parts
▪ To estimate effect of different environmental factors on crop
growth. E.g.:- Effect of temperature on rice
▪ To evaluate effect of different levels of a given management
practices. E.g:- effects of fertilizer treatments, irrigation
frequencies, growth hormones on the growth of specific crop.
▪ To determine the net result of different physiological process
within a plant. Net result of photosynthesis, respiration
Growth Curve
 The graphical representation of variation in total dry
weight with the time is called growth curve.

▪Sigmoid shape, therefore

a.k.a. sigmoid curve.
▪Slope of the growth curve
describes the dry weight
increases per unit time.
This is the growth rate.

Any plant parts – leaf, stem, shoot, root,

reproductive etc
Phase Description Physiological basis

1.Lag • During this phase total DW ▪ Occurs just after germination.

increases at a very slow rate. ▪ The plant uses the nutrients
•  the slope of the curve is stored in the seed and begins to
very low. develop above ground shoot
(Stem, leaves ) & root systems.
▪ The foliage canopy is very small
and very little light interception
occurs.  photosynthetic rate is
low.
▪  the dry weight increases very
slowly during lag phase.
Phase Description Physiological basis

2. • During this phase total DW ▪ the foliage canopy grows

Expone increases at an increasing rapidly with increased leaf
ntial rate. initiation and expansion.
• Therefore the slope of the ▪ Consequently light
curve increases rapidly. interception increases,
causing a significant increase
in the phot. rate.
▪ Causes the total DW to
increase at an increasing rate.
▪ This phase produces a high
proportion photosynthetic
tissues (Leaves, green parts)
& non photosynthetic tissue -
Roots, Stems
Phase Description Physiological basis
3. • During this phase total ▪ the foliage canopy continues
Linear DW increases at a to grow and light
constant rate. interception continuous to
• Therefore the slope of increase.
the curve is constant. ▪  a continuous increase
occurs in the crop DW due
to increase photosynthesis.
▪ However in addition to
photosynthetic tissue a
significant proportion of non
photosynthetic tissues also
increases.
▪ Here another important
physiological process
determines the variation of
growth rate of plants i.e.
Respiration.
 Phase Description Physiological basis

4. • total DW increases at a This phase occurs due to two reasons.

Declinin decreasing rate. ▪ Further increase in non photosynthetic
g • Therefore the slope of tissues.
the curve decrease ▪ Mutual shading -
gradually until it o When leaves are produced continuously
becomes zero at the the canopy develops several leaf layers.
end of declining phase. o In intercepting the incident solar
radiation leaves in the top leaf layers
would intercept most of the light.
o Therefore leaves in the middle and
bottom layers become shaded and
receive very little light energy.
Phase Description Physiological basis
5. • During this Reasons.
Negativ phase total dry ▪ Stopping of leaf growth.- Production of
e weight new leaves stop after crop passed a certain
stage. Therefore light interception and
decreases with
phot.do not increase further.
time. ▪ Leaf ageing (Senescence) -With
• Therefore the increasing age of leaves their
slope of the photosynthetic rate decreases while the
curve is respiration rates increases. Consequently
negative. after certain stage dry matter production
through photosynthesis becomes lower
than dry matter loss through respiration
▪ Leaf fall (Leaf shedding) -During the
latter part of a crops life cycle senesced
(aged) leaves are shed. This also
contributes to the decrease of total dry
weight during negative phase.
Growth Absolute Growth Rate System Production
Indices (AGR)

Relative Growth Rate (RGR) System Productivity

• The growth curve gives a
general description of the Crop Growth Rate (CGR)
plant/ crop growth. However
when we want to make Leaf Area Ratio (LAR) System size
specific comparisons
between crops, varieties, Net Assimilation Rate (NAR System Efficiency
environment conditions or -ULR)
management practices
different growth indices are Leaf Area Index (LAI)
used.
Leaf Area Duration (LAD)
Absolute Growth Rate (AGR) & Crop Growth Rate (CGR)
AGR CGR
Definition – Increase in total dry
Definition – Increase in total dry
weight per unit land area of a crop
weight per plant per unit time
per unit time.
(W 2 − W 1) W 2 − W1 1
AGR= CGR= t 2 − t1
×
A
(t 2 − t1) Weaknesses
Used to quantify growth of single plants (eg: ▪ Other than giving quantitative estimate of
in pot experiments) where the components of growth of a crop CGR does not provide
a crop (i.e. a plant population) spread over a any insights into the underlying
given land area is absent.] physiological mechanisms.
Weaknesses ▪ CGR is a poor indicator of the biomass
▪ Weakness same as CGR production efficiency of a crop.
▪ The most common units are g m-2 d-1, g m-
▪ Units: - g d-1 or g wk-1 2 wk-1 or kg ha-1 wk-1

Where, W 1 & W2 are total dry weights of crops over a sampling

area of A at times t1 and t2 days after planting.
Solar radiation and irradiance
Why radiation is important to plants

1) The absorbed radiant energy will affect tissue temperature and,

consequently, rates of metabolic processes, and energy
exchanges (i.e.transpiration)
2) The visible fraction of the incident solar radiation (PAR-
photosynthetically active radiance) is needed for
photosynthesis
3) The energy of specific wavelengths will influence
morphogenesis and rate of development (i.e; red:far red ratio,
photoperiod).
Classification of Earth Solar radiation
Solar radiation on the earth can be classified as:

▪ Direct radiation: This radiation comes directly from the sun without
any change in its direction. This type of radiation is characterized
by projecting defined shadow onto the objects that intersect.
▪ Diffuse radiation: This radiation comes from all over the
atmosphere as a result of reflection and scattering by clouds,
particles in the atmosphere, dust, mountains, trees, buildings, the
ground itself, and so on.
Global radiation: Is the total radiation. It is the sum of Direct & Diffuse
radiations. On a clear day with a clear sky, the direct radiation is
predominant above the diffuse radiation.
Light Interception by the canopies

▪ Definition: Sunlight intercepted by canopy relative to total

incoming sunlight
▪ Maximum effective interception in high density orchards
(fruit trees) is 65-75%
▪ Function of:
❖ Tree size (height & width) & shape
❖ Planting distances (within & between row)
❖ Canopy architecture & leaf density
Radiation and Canopy Interception
Plants intercept direct and diffuse sunlight.

▪ The upper leaves receive

both types of radiation, while
the lower leaves intercept a
small portion of direct
radiation.
▪ Diffuse radiation therefore,
becomes more significant in
the lower leaves due to
radiation transmitted and
reflected from the leaves and
the soil surface. Solar
radiation transmitted by the
Crop cover leaves is predominantly Forest cover
infrared.
Radiation distribution within a tree canopy
Radiation/light received as a percentage to full sun

Low Photosynthesis, weak leaf

development & poor fruit color
 ▪ The incident light at the top of the
canopy is absorbed by
successive layers of leaves in the
canopy. This light enters the
Radiation bottom of the canopy by:

distribution ▪ Direct radiation as sun fluxes

through the gaps in the
within canopy

canopies ▪ Scattered light from the

leaves and soil.
▪ Transmittance through the
leaves.
 LAI & Interception
• For an efficient use of solar radiation by crop, must be absorbed by
photosynthetic tissues. Leaf is the principal photosynthetic functional
unit, therefore its efficiency on the capture and use of solar energy
determines the crop productivity.
• The area and arrangement of foliage (the canopy architecture),
determine the interception of solar radiation by a crop and the
distribution of irradiance among individual leaves.
• Leaf area and arrangement change during the life of a crop and,
by leaf movement, even during the course of a single day.
• Deficiencies in water and nutrient inputs may reduce the rate of leaf
growth, reducing yield below optimum levels.
LAI & Interception during cropping period
Effects of nutrient & water status