Cooperative Extension Service

The University of Georgia College of Agricultural and Environmental Sciences

Cotton Growth
and Development
 Table of Contents

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Inside the Seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Germination and Seedling Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
The Cotyledons and First True Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Soil Effects on Germination and Early Root Growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Root Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
The Meristems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Vegetative Growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Leaf and Canopy Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
The Source to Sink Relationship . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Development of Fruiting and Vegetative Branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Formation of the Cotton Bud from Square to Bloom . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
The Cotton Flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Stages of Flowering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Nodes above White Flower and Cutout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Defoliation and Harvest Timing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Fruit Shedding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Boll Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Yield Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Heat Units or DD60s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
 Cotton Growth and Development
Glen L. Ritchie, Research Coordinator
Craig W. Bednarz, Cotton Physiologist
Philip H. Jost and Steve M. Brown, Extension Agronomists

Introduction seed. Despite its inherent perennial growth habit, how-

ever, cotton is managed as an annual crop plant, and
Domestic cotton has a unique origin and history growers try to produce as much lint and seed as
among cultivated crops. The wild ancestors of modern possible. Continued vegetative growth after flowering
cotton species were perennial vines that inhabited sev- diverts the plant’s energy away from lint and seed
eral distinct geographic areas, including Africa, Arabia, production, so the perennial nature of even modern
Australia and Mesoamerica. During the past several cultivars opposes our current production system.
centuries, people native to these regions developed four The cotton plant also produces fruit on two different
distinct species of cultivated cotton, including upland types of branches, each unique in growth habit, further
cotton (Gossypium hirsutum L.), the primary species complicating crop management. In addition, cotton
grown in the United States. Despite the selective breed- growth is very sensitive to temperature and soil condi-
ing efforts of humans, many of the wild characteristics tions. As in other crops, producers use chemicals in
of cotton have not been removed, making cotton man- cotton to control weeds and insects, but cotton is unique
agement difficult and unique. in that crop growth must also be regulated and eventu-
Wild cotton is a tropical perennial plant with an ally terminated by chemical means. Understanding the
indeterminate fruiting habit, meaning that it continues growth and development of the cotton plant helps pro-
to produce new foliage even after it begins to create ducers grow a high-yielding, high quality crop.
The following discussion is intended to provide
applicable information on the growth and development
of the cotton plant. The Georgia Cotton Production
Guide (updated annually) is an excellent data source for
the agronomic inputs required for producing cotton. The
production guide and other useful cotton links can be
found on the University of Georgia Cotton Web Page at
http://www.griffin.peachnet.edu/caes/cotton.

Inside the Seed

A mature cotton seed contains all of the organs
necessary to produce a small seedling. The seed is
pointed on one end (the micropyle) and rounded on the
other (the chalaza). The tip of the primary root, or
radicle, faces the micropyle, and the precursors of the
stem and cotyledons are plainly visible within the seed
(Figure 1).
The chalaza is the primary site of water and oxygen
absorption during germination. The tip of the primary
root, or radicle, is the first part of the plant to emerge
through the micropyle. The cotyledons that will nourish
the new seedling are folded inside the seed, with the
hypocotyl below them ready to elongate and push the
seedling through the soil. The gossypol glands visible
Figure 1. A small, dormant seedling rests inside a
mature seed. When the seedling emerges, the radicle throughout the inside of the seed are also visible in the
will be the primary root, the hypocotyl will be the stem tissues of the growing plant.
under the cotyledons, and the epicotyl will be the stem
above the cotyledons from which shoot growth occurs.

Cotton Growth and Development 3

Germination and vegetative and reproductive growth of the plant occurs
through the meristems.
Seedling Development A week or so after seedling establishment, the first
Germination begins as the seed absorbs water and true leaf appears above the cotyledons (Figure 4). The
oxygen through its chalaza after planting. The water first leaf shifts the plant’s primary energy source from
swells the dormant tissues, and cell growth and division storage to photosynthesis and signals the move from
begin to take place. The radicle emerges through the emergence to vegetative growth.
micropyle, turns downward, and grows deeper into the
soil, providing a taproot that will supply water and
nutrients throughout the life of the plant (Figure 2a and
b). The hypocotyl elongates from the radicle and forms
an arch or crook that begins to push up through the soil,
a brief period often referred to as the “crook stage”
(Figure 2c).
Seedling emergence normally takes place 4 to 14
days after planting. At the soil surface, the hypocotyl
straightens and pulls the folded cotyledons out of the
soil (Figure 2d), a process known as epigeal germina-
tion. After the cotyledons are pulled through the soil
surface, they unfold and expose the epicotyl and the
apical meristem, or growing point, which will be the
source of subsequent growth (Figure 2e-f). At this
point, germination and seedling emergence are com-
plete and the plant begins its active vegetative growth.
Figure 3. The cotyledons are storage organs
The Cotyledons and that are formed in the seed and emerge from
First True Leaves the soil as leaf-like structures oriented oppo-
site each other on the seedling stem. The
The cotyledons (Figure 3) serve a dual role in germi- cotyledons provide nutrients for the seedling.
nation. Before they unfold, they supply stored food to The apical meristem emerges through the
the germinating seedling. After the cotyledons unfold, cotyledons and will be the source of new
they produce chlorophyll, become green, and produce growth as the plant matures.
energy through photosynthesis. The apical meristem
emerges at the base of the cotyledons, and all further

Figure 4. The first true leaf emerges about 7

days after seedling establishment. From this
point on, the meristems will produce all veg-
Figure 2. Germination and early seedling development. etative and reproductive structures on the
Root growth dominates the early growth of the plant. plant.

4 Cotton Growth and Development

 Cotton emerges the quickest from warm, moist soil.
Low temperatures (below 60 degrees F) or less than
adequate soil moisture may hinder germination by
slowing metabolic processes (see the discussion on heat
units). Physical impedance, such as crusting, does not
slow germination, but it can prevent the hypocotyl from
emerging. This often causes thickening of the hypocotyl
and a condition referred to as “big shank” or “thick-
legged” cotton, resulting in reduced seedling vigor (Fig-
ure 5). Generally, the longer it takes for emergence to
occur, the greater the risk of plant death and yield loss.
A rule of thumb for planting cotton in most regions of
the U.S. Cotton Belt is that the soil temperature at 4
inches deep should be at least 65 degrees F for 3 con-
secutive days, with warm temperatures in the forecast.
Figure 5. Thick shank in a cotton seedling.
Soil crusting, compaction or other mechanical Root Development
factors can cause thick shank.
As the cotton plant grows, the radicle that originally
emerged from the seed becomes a taproot, from which
Soil Effects on Germination and lateral roots begin to form and grow. Lateral roots and
Early Root Growth the taproot collectively make up the basal root system.
Other “higher order” roots then develop from this basal
Root growth dominates the growth of the cotton root system. These higher order roots have a functional
plant during germination and seedling establishment. In life of about 3 weeks. They form when environmental
fact, the taproot may be as deep as 10 inches by the time conditions are good, and then die when nutrients and
the cotyledons emerge. This is a critical time for the water are depleted in the area in which they developed.
development of the root system. Cold soils, seedling As the plant matures, the roots continue to spread
disease, low soil pH, water stress, hard pans and herbi- and probe deeper in the soil profile for water and nutri-
cide injury all inhibit root growth and development, but ents. Therefore, the distribution of roots tends to match
careful crop management can minimize most of these the most fertile soil zones. Figure 6a shows an example
stresses. The roots absorb water and nutrients that are of the root distribution of an unstressed cotton commun-
vital to the development of the plant, and any hindrance ity. Most of the roots in this case can be found between
of root development in these early stages of cotton 1 and 3 feet deep in the soil, but large quantities of roots
growth may cause a disappointing production season. can still be found more than 4 feet deep in the soil. The

Figure 6. (a) Comparison of root quantity with soil depth for a mature cotton plant. New roots are con-
stantly produced in areas of the soil profile that have water and nutrients. (b) Comparison of root quantity
with cotton growth stage. Roots begin to decline after flowering as the cotton plant shifts its energy from
root to boll development.

Cotton Growth and Development 5

amount of roots generally peaks during the cotton flow- ents can develop very tall, heavy vegetative growth
ering phase then declines as the plant partitions more (Figure 8, page 7). This type of rank growth promotes
carbohydrates to the developing bolls (Figure 6b). boll rot and fruit abscission, and makes a cotton crop
difficult to harvest.
The Meristems The first vegetative structures that appear on the
main stem are main stem leaves (Figure 9, page 7).
The cotton plant has meristems, or growing points, Main stem leaves and branches form at points of attach-
at the top of the main stem and on its fruiting branches. ment on the main stem called nodes. As a general rule, a
These meristems allow the plant to simultaneously grow new node is produced from the apical meristem an
upward and outward. Figure 7a is a micrograph of the average of every 3 days, although nodes develop more
apical meristem and first two fruiting branches, which quickly early in the growing season than later in the
are too small to be seen without magnification. season.
Thrips feed on these young meristems, and plant The stem-like structure that connects the leaf with
injury occurs when the thrips insert their mouth parts the stem is called a petiole. Leaves that arise directly
into the cells to feed. The cells near the insertion point from the main stem are referred to as main stem leaves,
die, but the cells around them continue to expand and while leaves that arise from the fruiting branch are
divide, resulting in crinkling and distortion of expand- referred to as subtending leaves. The fruit produced by
ing leaves. Thrip damage slows plant growth, and thrip- a branch will primarily receive carbohydrates produced
damaged leaves have a puckered appearance and may by the leaf subtending that fruit. However, the main
have holes in them because of this damage (Figure 7b). stem leaf also supplies carbohydrate for fruit develop-
ment. Fruit produced closer to the main stem will
Vegetative Growth receive more carbohydrates from the main stem leaf
than fruit produced at more distal positions.
Cotton has an indeterminate growth habit and can A fruiting bud, called a square, begins to form at the
grow very tall under conditions of unrestrained growth. initiation of the fruiting branch. The first square
Growth regulators, such as mepiquat chloride, are gen- produced on a fruiting branch is referred to as a first-
erally applied to cotton to slow internode elongation, position square. As this square develops, the portion of
especially for well-fertilized irrigated cotton. Otherwise, the fruiting branch between the main stem and the
vigorous cotton varieties with plenty of water and nutri- square also elongates. This portion of the fruiting

Figure 7. (a) Light micrograph of a cotton plant apical meristem magnified 40x. The two
fruiting branches in this micrograph are too small to be seen with the naked eye. (b) The
cells where thrips feed on the meristems die, and the resulting leaves appear crinkled and
have holes in them.

6 Cotton Growth and Development

 branch is also called the internode, similar to the por-
tion of the main stem between main-stem nodes. An
axillary meristem also develops adjacent to this square.
The axillary meristem produces a second position
square and subtending leaf. As many as four squares
may be produced in this fashion on a fruiting branch.

Leaf and Canopy Development

Plant growth and development are both functions of
sunlight interception and temperature. As a cotton plant
develops, new leaves appear and expand, increasing
sunlight interception. Initially the carbohydrates pro-
duced by the leaves are used to produce roots and more
leaves. This production of new leaves causes the leaf
Figure 8. Cotton can grow very tall if its growth is not area of the cotton plant to increase rapidly. Once repro-
held in check by environmental factors or management ductive structures begin to develop, carbohydrate sup-
practices. plies are slowly shifted to the developing fruit. As the
fruit load on the plant increases and ages, the carbohy-
drate demand increases, and the development of new
leaves steadily declines. Therefore, fruit development
occurs with a leaf population that is steadily aging.
Leaf photosynthesis does not remain constant as the
leaf grows and develops (Figure 10). A cotton leaf
reaches its maximum photosynthetic capacity at about
20 days of age, after which it declines. Collectively, as
the reproductive growth of the cotton plant is increas-
ing, it is doing so with the support of a leaf canopy that
is aging. Premature aging of the cotton leaf canopy due
to water stress, low fertility and other stresses further
reduces the photosynthetic capacity of the crop.
Figure 9. Growth of a fruiting branch from the main
stem. The branch forms in the axil above a main stem
leaf. The leaves and stems on nodes above and below The Source to Sink Relationship
the one illustrated have been removed. Most of the cotton plant’s carbohydrate energy is
directed to root growth prior to the time reproductive
growth begins. This is a function of carbohydrate
source to sink relationships (Figure 11, page 8). Carbo-
hydrates are transported from supply areas, called
sources, to areas of growth or storage, called sinks. The
leaves are the primary source of carbohydrate produc-
tion during the early vegetative growth of cotton. Car-
bohydrates are produced through photosynthesis in the
leaves and channeled through the phloem to the roots,
which act as the main carbohydrate sinks during this
phase. The source-to-sink phenomenon also applies to
the transport of inorganic nutrients and water. The roots
are the source for all inorganic nutrients and water,
which are transported through the xylem to sinks
throughout the plant. Thus, the root and shoot systems
Figure 10. Photosynthetic capacity of a cotton leaf rela- are very interdependent, and damage to either system
tive to leaf age. Leaves reach peak photosynthetic cap- slows growth and decreases yield.
ability about 20 days after they unfold, after which their As bolls begin to develop, they become much
efficiency decreases over time.

Cotton Growth and Development 7

 they grow straight and erect, much like the main stem
(Figure 12). Vegetative branches can also produce
fruiting branches.
The branches from which fruiting buds arise are called
fruiting branches, or sympodia (meaning “multiple
feet”), because each fruiting branch contains multiple
meristems. Fruiting branches have a “zig-zag” growth
habit, as opposed to the straight growth habit of the
vegetative branches (Figure 13). The initial growth of a
fruiting branch is terminated once a fruiting bud forms.
The fruiting branch, however, initiates a new growing
point, called an axillary meristem. The axillary
meristem is located at the base of a leaf that subtends
Figure 11. Source to sink relationships at two stages of the newly formed fruiting bud. The “zig-zag” growth
cotton growth. During early vegetative growth, most of habit is a consequence of the stop-and-go growth of the
the carbohydrates produced by the leaves are sent to fruiting branch.
the root system. Later in the season, however, most of The first fruiting branch will generally arise at main-
the carbohydrates are sent to the developing bolls, and stem node 5 or 6. A cotton plant will mainly produce
the root system and shoot growth rate decline. fruiting branches, but several common environmental
factors such as low population density, insect and dis-
stronger carbohydrate sinks than roots and shoots. At ease pressure and over-fertilization can cause vegetative
this stage, root and shoot growth slow, and boll devel- branches to form. Vegetative branches are produced
opment dominates plant growth, and the widely estab- after fruiting branches, and develop at nodes directly
lished roots continue to supply large quantities of water below the node at which the first fruiting branch was
and nutrients to the shoot. developed. For instance, if the first fruiting branch is
initiated at main-stem node 5, a vegetative branch may
Development of Fruiting and develop at main-stem node 4.
The cotyledons are oriented opposite each other on
Vegetative Branches the stem, but the true leaves and branches of the cotton
The branches on a cotton plant can be classified as plant occur in a 3/8th alternate phyllotaxy, meaning the
either vegetative branches (monopodia) or fruiting distance from one leaf to the next is 3/8th of a complete
branches (sympodia). Vegetative branches, like the turn around the stem (Figure 14a). Branches on the
main stem, are referred to as monopodia (meaning main stem also show this 3/8th alternate arrangement,
“single foot”) since they have only one meristem. since they grow adjacent to the leaves. Nodes are num-
Because vegetative branches have only one meristem, bered in the same order the leaves are numbered

Figure 12. A cotton plant with leaves removed shows Figure 13. A fruiting branch with leaves removed shows
the straight growth habit of the main stem and the its zig-zag growth habit.
vegetative branch.

8 Cotton Growth and Development

 Figure 14. (a) A defoliated cotton plant shows the d alternate phyllotaxy of branches. Each branch is d of
a turn around the stem from the branch below it. The branches form from the axils of main stem leaves.
(b) A diagram of the general timing of flower emergence from buds on the fruiting branches by fruiting
position.

where the cotyledonary node is considered node 0 The Cotton Flower

(Figure 14b).
New fruiting branches are generally believed to As discussed previously, the cotton square is actu-
develop approximately every 3 days, although recent ally a flower bud. The first visible structures of the
studies show that this developmental rate varies. square are the leaf-like bracts, or epicalyx. Three bracts
Squares are produced at new positions on a fruiting surround the flower bud in a pyramid-like shape. The
branch approximately every 6 days. The age of fruiting cotton plant produces perfect flowers, meaning the
structures on a cotton plant can be mapped according to flower contains both male and female organs (Figure
this time sequence (Figure 14b). 16, page 10). The first square is typically visible on
node 5 to 7 about 35 days after planting. Anthesis, or a
Formation of the Cotton Bud flower bloom, occurs approximately 21 days after the
first square appears.
From Square to Bloom When a pollen grain reaches the stigma, it germi-
During the 21-day period from square to bloom, nates into a pollen tube. The pollen tube grows through
there are several recognized developmental stages of the the style, the micropyle, and into the ovule chamber,
cotton flower bud. A “pinhead” square is the first stage where fertilization takes place. Anything that reduces
at which the square can be identified. The next stage of egg or pollen viability or tube growth in a flower
square growth is “match-head” or “one-third grown” adversely affects the final yield for that boll.
square. Just prior to the time the flower opens, a candle
shape can be seen (Figure 15d, page 10). This period of Stages of Flowering
square development prior to bloom is called “squaring.”
Flowering is important to cotton production because
Once the cotton begins to bloom, it is said to be
pollinated flowers form cotton bolls. The bloom process
“flowering.” A cotton plant typically blooms or flowers
takes several days, and bloom age can be estimated by
for about 6 weeks. Thus, until the cotton begins to pro-
the bloom characteristics (Figure 17, page 11). On the
duce fruit, the stage of development is discussed in
day a flower opens it is white in color. Pollination of
terms of leaves or nodes. Once fruiting begins, the stage
that flower usually occurs within a few hours after the
of cotton development is discussed in terms of square
white flower opens.
development and the number of nodes. Once blooms are
On the second day the flower will have a pink-like
present, the stage of cotton development is discussed in
color, and a red color on the third day. Approximately 5
terms of weeks of bloom.

Cotton Growth and Development 9

 Figure 15. Development of the bud from a match head square (a) to a flower (e) involves
both a size increase and petal development. Two bracts have been removed from each
square, candle and bloom to show this development.
to 7 days after a flower appears it usually dries and falls ment and ceases flower development. This event is
from the plant exposing the developing boll. Occasion- termed cutout. Cutout generally occurs at 4 or 5
ally a flower will stay attached to the developing boll NAWF. Cutout occurs when carbohydrate supply
for a longer period of time. This is referred to as a equals demand and vegetative growth ceases. At cutout,
bloom tag (Figure 17d, page 11). no more harvestable fruit is set.

Nodes Above White Flower Defoliation and Harvest Timing

And Cutout Defoliants, or harvest aids, are used to defoliate
The development of the cotton plant in terms of leaf cotton, enhance boll opening, and control regrowth
number, node number and fruiting stage is discussed in prior to harvest. Defoliants effectively terminate the
previous sections. During the flowering period, the cotton crop and prepare it for machine harvest at the
stage of cotton development can also be discussed in end of the growing season. These chemicals also give
terms of Nodes Above White Flower (NAWF). This is a the producer some control over harvest timing and
measurement documenting the number of nodes separa- increase harvest efficiency.
ting the uppermost first position bloom and the terminal
of the plant.
When the cotton plant first begins to bloom there
will be approximately 9 to 10 NAWF (Figure 18, page
11). As the season progresses, the number of NAWF
decreases. This reduction in NAWF can be related to
the source to sink relationship of carbohydrate supply.
NAWF generally decreases more quickly after bloom in
early-maturing varieties than in mid or full season vari-
eties. As the flowers develop into bolls, they become
stronger sinks for carbohydrates and their combined
demand for carbohydrates increases. Eventually the
carbohydrate supply produced by the leaves will be
used primarily by developing bolls, leaving less and
less available for the production of new vegetative
growth. As flowering progresses up the plant, less top
growth is produced, allowing the NAWF to decrease.
As the flowering approaches the top of the plant, the Figure 16. Parts of a mature cotton flower. The cotton
plant eventually puts all of its energy into boll develop- flower contains both male and female parts.

10 Cotton Growth and Development

 Figure 17. Development of a cotton bloom. A white flower emerges on day 1 (a),
then gradually darkens and takes on a red color during days 2, 3 and 4 after
emergence (b and c). The bloom eventually dries up and either falls off or be-
comes a bloom tag (d).

Table 1. Active ingredients of chemicals used for defoliation

and harvest preparation of cotton.
Class of Active
Defoliant Ingredient Comments

Hormonal Thidiazuron Enhances ethylene production and

inhibits auxin transport.
Dimethipin Causes rapid water loss through the
stomata of the leaves, which leads to
ethylene production as leaves become
water-stressed.
Ethephon Increases ethylene production; used
primarily for boll opening.
Figure 18. Comparison of canopy photosynthesis with
boll development. Cutout occurs as photosynthesis Herbicidal Tribufos Injures leaf cells to trigger ethylene
declines and more energy is partitioned to forming bolls. production.
Carfentrazone Inhibits a step in chlorophyll synthesis,
Defoliant performance is affected by temperature, causing destruction of cellular mem-
plant condition, spray coverage and product rate. Tem- branes and ethylene production.
perature is the primary force in determining harvest-aid Pyraflufen Ethyl Inhibits a step in chlorophyll synthesis,
rate. Under optimal conditions, a cotton crop might be causing destruction of cellular mem-
harvestable in as little as 7 days after defoliation, but branes and ethylene production.
cool temperatures will prolong the defoliation process. Paraquat Non-selective desiccant.
Cotton harvest aids can be classified into two modes Chlorates Non-selective desiccant.
of action, herbicidal and hormonal. Herbicidal harvest Glyphosate Used for regrowth control and weed
aids injure the leaf, stimulating the production of ethy- management.
lene. Hormonal harvest-aids increase the ethylene
concentration in the leaves without causing any injury. A detailed discussion of defoliation and harvest
A list of the active ingredients in most defoliants is timing can be found in the University of Georgia exten-
found in Table 1.

Cotton Growth and Development 11

 plant due to the same factors that lead to square shed-
ding (Figure 20). Generally, though, the sensitivity of
squares, flowers and bolls to shedding can be related to
their age. Young fruiting forms are more likely to be
shed than are more developed squares and bolls.

Boll Development
After pollination occurs the boll begins to develop.
Under optimum conditions it requires approximately 50
days for a boll to “open” after pollination occurs. Boll
development can be characterized by three phases:
enlargement, filling, and maturation.
The enlargement phase of boll development lasts
approximately 3 weeks. During this time the fibers
produced on the seed are elongating and the maximum
Figure 19. Square shedding is a common occurrence in volume of the boll and seeds contained therein are
cotton. attained. Also during this time, the fiber is basically a
sion publication Cotton Defoliation, Harvest-Aids, and thin walled tubular structure, similar to a straw. Each
Crop Maturity, by Philip Jost and Steve M. Brown, and fiber develops from a single epidermal cell on the seed
yearly updated specific harvest-aid suggestions can be coat. During the boll enlargement and fiber elongation
found on the University of Georgia Cotton Web Page at phase, the development of the fiber is very sensitive to
http://www.griffin.peachnet.edu/caes/cotton adverse environmental conditions. Low water availabil-
ity, extremes in temperature and nutrient deficiencies
Fruit Shedding (especially potassium) can reduce the final fiber length.
The filling phase of boll development begins during
A phenomenon often seen in a cotton field is square the fourth week after flowering (Figure 21, page 13). At
shedding (Figure 19). The shedding of squares may be this time, fiber elongation ceases and secondary wall
the result of several factors, including water stress, formation of the fiber begins. This process is also
shading (from prolonged cloudy weather), nutrient defi- known as fiber filling, or deposition. Cellulose is
ciencies (especially N), high temperatures, high plant deposited inside the elongated fiber every 24 hours,
populations, high percent fruit set and insect damage. In filling the void space of the elongated fiber. The deposi-
addition, the reproductive cells formed during square tion of cellulose into the fiber cell is also sensitive to
development are very sensitive to environmental condi- environmental conditions. Water, temperature and
tions. High temperatures and humidity, and nutrient nutrients (especially potassium) are the primary envi-
deficiencies (especially boron) can inhibit gamete pro- ronmental factors that influence this stage of boll
duction and result in flower sterility and ultimately development. The filling phase of boll development
square loss. Sterility may also decrease seeds per boll continues into the sixth week after pollination.
and locks per boll. One cause of pollen sterilization and The boll maturation phase begins as the boll reaches
subsequent yield loss is misapplication of glyphosate in its full size and maximum weight. During this phase,
Roundup Ready® cotton. fiber and seed maturation take place and boll dehis-
Flowers and young bolls may also be shed from the cence occurs. The capsule walls of the boll dry, causing

Figure 20. Examples of square and boll shedding. Shedding can be related to several environmental factors such
as population density, water, nitrogen, insect pressure and disease. Newly formed bolls and young squares tend to
be the most susceptible to shedding.

12 Cotton Growth and Development

 Figure 21. Development of the cotton boll. Undeveloped cotyledons have a jelly-like consistency. As the cotyle-
dons develop, they become firm and leaf-like, and a black layer that will become the seed coat begins to form
around the seed. As the black layer matures and the outside of the boll dries, the dorsal suture splits, allowing the
cotton boll to open.

the cells adjacent to the dorsal suture to shrink uneven- of row, first position bolls contribute from 66 to 75 per-
ly. This shrinking causes the suture between the carpel cent of the total yield of the plant, and second position
walls to split, and the boll opens. bolls contribute 18 to 21 percent.
Yield distribution research is an intensive, detailed
Yield Distribution process that involves counting and weighing bolls from
each fruiting position on many plants. First position
The contribution of a single fruiting structure to the bolls tend to fill out more and be heavier than bolls
overall yield of the cotton plant depends largely upon from other positions, so the majority of boll weight on
its position on the plant. First position bolls are heavier plants generally comes from the first position fruit
and produced in higher quantities than bolls at any other between nodes 7 and 20 (Figure 22).
position. In cotton populations of three plants per foot
Heat Units or DD60s
Cotton growth milestones are often given in terms of
days after planting or between growth stages, but the
development rate of cotton is strongly influenced by
temperature. A cotton crop grows more slowly on cool
days than on warm days, so temperature measurements
during the cropping season help estimate when a crop
reaches a specific developmental stage. Heat units, or
DD60s, are an estimation of this accumulated tempera-
ture effect during a day, based on the average of the
maximum and minimum daily temperatures in degrees
Fahrenheit (oFmax and oFmin, respectively). The number
60 is subtracted from this average, because 60 degrees F
is generally accepted as the lowest temperature at which
cotton growth occurs. The formula for calculating heat
units per day is as follows:
DD60 = (oFmax –oFmin) – 60
2
Calculating the accumulated heat units of a crop
over time can then be used to estimate the growth of the
cotton during the season. Table 2 demonstrates how to
calculate accumulated heat units over a 5-day period.
Figure 22. Comparison of cotton yield by fruiting
Scientists at the University of Georgia Tifton
position and main-stem node. First position bolls Campus have measured daily temperature data since
are heavier and more abundant than bolls at the 1928, and the average heat unit accumulation pattern
other positions, making them the primary source of for a cotton crop planted on May 1 at this location is
yield. illustrated in Figure 23.

Cotton Growth and Development 13

 Table 2. Calculation of daily and accumulated heat units based on daily high and low temperatures.
Daily High Daily Low Average Daily
Temperature Temperature Temperature Daily Heat Units Accumulated
Day (oFmax) (oFmin) (oFmax+oFmin)/2 (oFmax+oFmin)/2 – 60 Heat Units

1 81 61 71 11 11
2 83 63 73 13 24
3 82 62 72 12 36
4 85 66 75.5 15.5 51.5
5 80 62 71 11 62.5

Table 3 shows typical heat unit accumulations at of the development of the cotton plant is crucial for
which a cotton crop reaches various growth milestones, making management decisions and maintaining profit-
as well as the average number of days after planting that able production.
these heat units are accumulated in South Georgia.
These numbers will vary according to location, year and References
cotton variety.
Constable, G.A., and H.M. Rawson. 1980. Effect of leaf
Table 3. Accumulated heat units required for a normal position, expansion and age on photosynthesis,
cotton crop to reach a specific growth stage. These transpiration and water use efficiency of cotton.
values will differ by variety. The third column Aust. J. Plant Physiol. 7:89-100.
shows the average number of days after planting at Dumka, D., C.W. Bednarz, and B.W. Maw. 2004.
which these heat units are accumulated, based on Delayed initiation of fruiting as a mechanism of
historical data from Tifton, Ga. improved drought avoidance in cotton. Crop
Growth Stage Heat Units Days Sci.:44:528-534.
Stewart, J.M. 1986. Integrated events in the flower and
Emergence 50 5
fruit, In J. R. Mauney and J. M. Stewart, eds.
First Square 550 38 Cotton Physiology, Vol. 1. The Cotton Foundation,
First Flower 950 59 Memphis, Tenn.
Open Boll 2150 116 Wullschleger, S.D., and D.M. Oosterhuis. 1989. Water
Harvest 2600 140 use efficiency as a function of leaf age and position
within the cotton canopy. Plant and Soil 120:79-85.
Summary
Cotton is a unique crop plant, and its innate growth
pattern makes it challenging to grow. However, the
plant develops in a somewhat predictable pattern. Initi-
ally, leaf area and vegetative structures are developed
that will then support future reproductive growth. If this
initial vegetative growth is compromised, subsequent
reproductive growth also suffers. Unlike many other
crops, the cotton plant continues vegetative growth after
flowering begins. The development of fruiting struc-
tures ultimately reduces vegetative growth as the plant
matures. The environment regulates every developmen-
tal process of the cotton plant, both vegetative and
reproductive. Heat unit accumulation dictates develop-
ment as much as time. Figure 23. Accumulated heat units during the growing
Due to increasing production costs and decreasing or season based on historical data at Tifton, Ga., from
stagnate commodity prices, cotton producers must be 1928 to 2003, assuming a May 1 planting date. The
able to critically evaluate every input. An understanding heavy bar represents the average accumulated heat
units, and the light bars are ± 1 standard deviation.

14 Cotton Growth and Development

Cotton Growth and Development
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