Cottonseed
E Hernandez, Texas A&M University, College Station, TX, USA
ã 2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 1, pp 343–348, ã 2004, Elsevier Ltd.
Topic Highlights
• Traditional varieties of cottonseed contain gossypol
(0.6–2.0% in raw cottonseed kernels).
• Gossypol is a polyphenolic compound that is toxic to man
and monogastric animals, presumably functioning as a
natural insecticide for the plant.
• Gossypol is located in “glands”, appearing as black specks
in the stems, leaves, and seed.
• The breeding of “glandless” cotton varieties (since the
1960s) means that cottonseed is available without gossypol.
• Meal and oil from glandless cottonseed are similar in com-
position to these products from the traditional glanded
seeds.
• Glandless cottonseed offers the advantages for processing
of reduced energy costs, elimination of prepress operations,
and the absence of color in the resulting oil.
• Food products from glandless cottonseed include whole
roasted seeds, flour, protein concentrates, and isolates.
• Glandless cottonseed is suitable to feed monogastric ani-
mals, not only due to the absence of gossypol but also
because of higher available energy.
Learning Objectives
• Understand the important differences between glandless
and the traditional glanded cotton varieties.
• Become familiar with the processing involved in producing
the various products from cottonseed for food and feed
uses.
Introduction
Most varieties of cotton (Gossypium hirsutum L., G. aboreum L.,
G. barbadense L., or G. herbaceum L.) are grown mainly in warm
climates around the world. Over 98.5 million tons (Mt) of
cotton were produced worldwide in 2001–02 and over 33.6
Mt of cottonseed. Cotton is grown for its fiber (over 80% of its
value) and the seed is used mostly for oil recovery and feed.
Whole cottonseed can be fed to dairy cattle, and the meal
resulting from oil extraction is fed primarily to ruminants
and, in limited amounts, to poultry and swine.
Traditional varieties of cottonseed contain gossypol, a
yellow-green polyphenolic compound considered toxic to
man and monogastric animals (Figure 1), reportedly affecting
the heart, liver, and reproductive organs. It has been used in
China as a male contraceptive but the practice was abandoned
because of permanent side effects. Gossypol is dispersed in
the plant as deposited structures or “glands,” which can be
seen as black specks in the stems, leaves, and seed (Figure 2).
Reference Module in Food Sciences http://dx.doi.org/10.1016/B978-0-08-100596-5.00025-1
The glands in the seed are ovoid structures containing 35–50%
gossypol and are 0.025–0.178 mm in diameter. These gossypol
glands are difficult to break by mechanical means, but
heat generated in extraction of oil by pressing, binds gossypol
to protein, turning it nontoxic. Switching to solvent extraction
with hexane, where no appreciable heat is generated, increases
the free gossypol content in the meal over ten times.
Raw cottonseed kernels may contain 0.6–2.0% free gossy-
pol. The Food and Drug Administration in the US (FDA) limits
free gossypol in human food products and ingredients at
450 ppm, and the Protein Advisory Group of the United
Nations Food and Agriculture Organization (FAO) and
World Health Organization (WHO) has set maximum guide-
lines of 600 ppm for free gossypol and 12 000 ppm total gos-
sypol. The feed industry in the US set limits for free gossypol
levels in poultry diets at 100 ppm maximum for broilers and
40 ppm for laying hens.
Removal by mechanical separation of the gossypol-contai-
ning glands has also been reported. Also addition of iron
salts, such as ferrous sulfate, which bind the gossypol in
feeds and render it biologically inactive, is practiced in some
countries.
Solvent extraction with hexane of cottonseed oil, com-
monly used commercially, removes only small amounts of
gossypol with the oil. However, extraction with more polar
solvents is effective in removing gossypol. Examples of polar
solvents used include aqueous acetone, a mixture of acetone
and hexane, and sequential extraction with hexane, aqueous
acetone, and anhydrous acetone. Blends of methylene chloride
and hexane-acetic acid have also been used. These solvents are
not widely used commercially.
Techniques have also been reported that mill cottonseed in
the presence of hexane and then remove the intact, heavier
gossypol glands by the liquid cyclone process. Also, an air
classification process has been developed to separate intact
gossypol glands from solvent-extracted ground flour.
Glandless Cottonseed
Some of the first studies on the development of glandless
cottonseed in the US were done in the early 1950s. These
studies described that selection of plants from the “Hopi
Moencopi” variety could result in almost complete elimination
of pigment glands from leaves and bolls. When crossing Moen-
copi with upland cotton varieties (G. hirsutum), it was found
that glandless seed appeared in later segregating generations.
Production of glandless cotton was first conducted in Iguala,
Mexico in 1960.
Basically two genes, gl-2 and gl-3, in the plant are found to
control the production of gossypol pigments in the seed. When
present in the homozygous recessive condition (gl2gl2–gl3gl3),
1
2 THE OILSEEDS | Cottonseed
Figure 1 Gossypol.
Figure 2 Cross-section of glanded cottonseed (left) and glandless
seed (right). (Data from Miravalle RJ (1972) J. Am. Oil Chem. Soc. 49:
24–26.)
all parts of the plant aboveground, including the seed, have
no pigment glands. The presence of pigmented glands can
be easily seen as black specks by cross-sectioning a seed
(Figure 2).
Recently, the value of the whole seed as cattle feed has
increased making it less profitable to extract the oil and sell
the meal. In some cases cottonseed is left at the gin in exchange
for ginning and baling services. Sometimes glandless cotton-
seed is considered to produce fiber of lower yield and quality
than traditional “glanded” varieties. Some studies have shown
that the glandless factor in itself does not decrease fiber yield or
quality as long as care is taken to prevent weed and insect
infestation.
Gossypol is regarded also to be a natural insecticide. Reports
show that glandless cotton attracts more insects than glanded
varieties. Entomology studies have shown that cotton bollworm
(Heliothis zea Boddie), tobacco budworm (H. virescens F.), pink
bollworm (Pectinophua gossyprella Saunders), and lygus insects
prefer glandless cotton. It is generally recognized that glandless
varieties require closer supervision to intercept and control
insect infestations.
Processing and utilization of glandless cottonseed was
reported in the early 1960s in the US. Glandless cottonseed
oil and meal were evaluated in poultry broiler and layer
rations. Glandless cottonseed meal was found nearly as effec-
tive as soybean meal in achieving broiler gains and did not
produce the green yolk discoloration in eggs commonly expe-
rienced with feeding glanded cottonseed meal.
The first commercial glandless cotton variety was a storm-
proof, boll-type cotton. It was evaluated in farm-scale trials and
made available for commercial planting in the late 1960s in the
US. Commercial sale of glandless cottonseed began in the early
1970s, with FDA approval. In 1976, under title 21 (Food and
Drugs) of the Code of Federal Regulations, sale of glandless
cottonseed kernels and cottonseed flour as food additives was
allowed. It included restrictions on gossypol (450 ppm). In
1978 the National Cottonseed Products Association (NCPA)
in the US established grades of glandless cottonseed products:
class A, to contain not more than 400 ppm of total gossypol;
class AA, to contain not more than 100 ppm total gossypol;
and class AAA, to contain not more than 10 ppm total gossy-
pol. Glandless cottonseed kernels started to be sold for use as
snack foods, in baked goods and in soft candy.
Research and production of glandless cotton for food and
feed is also reported in Europe, Africa, and Asia. Some US
cotton varieties were adapted to African crops. Development
and study of some Egyptian varieties of glandless cotton have
also been reported. From studies done in the Ivory Coast, it has
been suggested that glandless cottonseed cultivation and oil
mill processing is more economically viable in that part of
Africa. Conditions there may be better for glandless cotton
considering that in larger industrialized countries other com-
petitive oil seed proteins are more readily available. An addi-
tional advantage is that large, isolated land areas can be
dedicated to this crop and less problems of cross-pollination
would be encountered.
Processing and Products
Other than gossypol content very few differences are found in
glandless compared with glanded cottonseeds. Evaluations of
eight varieties each of glanded and glandless cottonseed were
reported (Table 1). Amino acid profiles were similar for
glanded and glandless seed. The storage and processing char-
acteristics of glandless and glanded cottonseed are also
reported to be essentially identical with regard to development
of free fatty acids, moisture levels, and refining losses in the
crude oils.
No major differences were encountered either in solvent
extraction of glanded versus glandless cottonseed. Extraction
rates of oil from glandless cottonseed flakes are about the same
as from glanded flakes, using commercial hexane. It has also
been suggested that glandless cottonseed can also be a source
of a new type of lecithin. Cottonseed has the highest content of
phospholipids after soybeans, present at about 2.2% in the oil,
but due to gossypol presence it was not commercially viable.
Currently, the major commercial source of lecithin is soybean
oil. Composition of cottonseed phospholipids consists of
phosphatidylcholine, phosphatidylethanolamine, and phos-
phatidylinositiol (33%, 22%, and 37%, respectively) and is
considered more oxidatively stable than soybean phospho-
lipids due to lower content of unsaturated fatty acids.
One of the main advantages of milling glandless cottonseed
is that it simplifies direct solvent extraction. Current commer-
cial practice of refining glanded cottonseed oil requires that the
 THE OILSEEDS | Cottonseed 3

Table 1 Comparative analysis means of eight varieties each

of glanded and glandless cottonseed and their productsa Advantages of processing glandless cottonseed over glanded
seed include:
Glanded Glandless
Product and assay cottonseed cottonseed 1. reduction in electrical energy used for flaking;
2. elimination of prepress operations with larger solvent
Whole cottonseed
extractors with lower maintenance costs, and reduction in
Oil (%) 21.0 21.1
processing energy;
Iodine no. 108.9 109.9
Protein (N  6.25; %) 23.1 22.5 3. since there is no color setting problem, crude cottonseed oil
Wt. 100 fumed kernels (g) 10.0 10.6 can be held longer in storage;
No. fumed seed/100 ml 542 510 4. elimination of more dangerous miscella (oil-solvent solu-
% Kernels in lint-free seed 61.7 59.6 tion) processing or on-site conventional refineries;
Cottonseed kernels 5. lower refining loss of glandless seed oil due to reduced use
Oil (%) 37.8 39.7 of alkali to eliminate color;
Protein (N  6.25; %) 39.3 38.9 6. reduction of bleaching earth needs; and
Crude fiber (%) 1.6 1.7 7. marketing of a light-colored, gossypol-free oil.
Total phosphorus (%) 0.8 0.9
Total sugars (%) 7.4 6.8 Some emulsification problems have been reported in refining
Total gossypol (%) 1.2 0.02 desolventized crude due to higher concentration of lecithin.
Wt. 100 kernels (g) 6.5 7.0 There are some drawbacks when switching to glandless
No. kernels/100 ml 912 844 crops; varying levels of gossypol can exist among individual
Hexane-extracted flour (meal)
seeds from the same plant. It is also difficult growing large
Oil (%) 0.8 0.8
acreages of glandless cottonseed, without cross-fertilization
Protein (N  6.25; %) 63.2 62.6
Crude fiber (%) 2.7 2.8 by windblown or insect-carried pollen of glanded varieties. In
Ash (%) 8.0 7.8 order to preserve the variety and seed lot purity in the produc-
Total phosphorus (%) 1.3 1.4 tion of glandless cottonseed products, it is important to mon-
Total sugars (%) 13.4 13.7 itor both, cross-fertilization from cotton plants in nearby fields
Total gossypol (%) 1.6 0.02 and reversion to the glanded condition during successive
Color, Hunter “L” values plantings due to fertilization between heterozygous plants.
Dry 84.3 89.8 Furthermore, segregated handling of glandless cottonseed in
Wet (5 water:1 flour) 48.1 71.3 gins and oil mills can also be costlier. As a result large cotton-
Crude oil
seed processors may be hesitant to handle glandless cottonseed
Cyclopropenoid fatty acids (%) 0.23 0.23
and dedicate separate processing facilities.
Fatty acids (%)
Myristic 0.9 0.7
Palmitic 23.0 22.6
Stearic 2.2 2.1
Oleic 17.7 17.7 Foods Uses
Linoleic 55.8 56.5
Unknown 0.4 0.4 Several food uses of glandless cottonseed kernels have been
Refined oil reported as well as applications on the use of glandless cotton-
Refined oil color, red 6.9 3.7 seed blended with other foods. Even though glanded cotton-
Bleached oil color, red 2.9 2.2 seed is not considered suitable for human consumption, there
a
Dry weight basis.
have been cases where cottonseed flour with bound gossypol
(Source: Lawhon JT, Cater CM, and Mattil KF (1977) Journal of the American Oil has been used in nutrition intervention feeding products, such
Chemists’ Society 54: 75.) as Incaparina in South America.
Food products from glandless cottonseed include whole
roasted seeds, flour, protein concentrates, and isolates. From
oil be refined in its miscella state (oil–hexane solution) right these products many foods were reported to be prepared. The
after extraction. This is necessary because if the solvent is technologies for the production of high-protein products have
removed from the oil prior to refining, the color fixes into the been well established and apply similarly to all vegetable pro-
oil and is very difficult to remove in subsequent bleaching teins. Specific technologies were developed for the production
operations. This means the equipment in refining of miscella of cottonseed protein products that were later applied for
has to be explosion-proof, which is more expensive and more processing of other vegetable proteins like soybean.
difficult to maintain. This is no longer the case with glandless The preliminary steps in preparing glandless cottonseed
seed processing, where binding of gossypol (by cooking and products are similar to the process used with glanded cotton-
hard press or prepress) or miscella refining become unneces- seed, i.e., ginned cottonseed is first cleaned to remove dirt and
sary. The oil from glandless seed is in lighter colored oil and other impurities, conditioned, de-hulled, and separated to pro-
the meal has greater amounts of soluble protein. Protein duce kernels. The kernels may be size-sorted and the larger
solubility of processed glandless cottonseed flakes or press particles roasted and color-sorted. The whole kernels are sepa-
cake has been reported to be higher than that of glanded seed rated for roasting or conditioning. The smaller or broken
(89.6%) compared with 84.8% for extracted flakes, and 59.3% kernels may then be further conditioned, flaked, and solvent-
for press cake. extracted for production of flours, concentrates, and isolates.
4 THE OILSEEDS | Cottonseed
Whole glandless cottonseed kernels are prepared either for
direct use as whole nut substitutes or for food ingredients.
These are also used in the production of high-quality flours
and food protein concentrates or isolates. The use of glandless
cottonseed kernels for the production of nutlike products by
means of various roasting methods has been reported, includ-
ing dry roasting at several temperatures, roasting under vac-
uum followed by steam injection, pressure steaming followed
by oven roasting, and deep fat frying in various oils (including
corn, cottonseed, peanut, safflower, soybean, and sunflower
oils). When the main product is to produce high-protein
flours, less care needs to be taken to minimize breakage during
de-hulling.
Glandless cottonseed flour can be prepared by extracting de-
hulled glandless seed flakes with hexane. The extracted flakes are
then desolventized and ground to a desired mesh size. In the
production of cottonseed concentrates, the defatted flakes are re-
extracted with acidified water or ethanol to remove the soluble
sugars and flavor compounds, and dried to produce protein
concentrate. A spray dryer can also be used in preparing con-
centrates where the defatted flakes are ground prior to extrac-
tion. Concentrates have been prepared from flour by a dry air
classification method and by acidic water extraction. Figure 3
shows a simplified diagram of the process to be used to generate
protein concentrate from glandless cottonseed.
Three major techniques have been reported for making
glandless cottonseed isolates. These isolates can be conven-
tional protein isolates, storage proteins (SP) (soluble at high
and low pH), and nonstorage proteins (NSP) (soluble at near
neutrality):
1. Ground flour is first extracted with dilute alkali (at pH 10)
and the insoluble residue removed by continuous centrifug-
ing or decanting. The clarified liquor is then precipitated at
one pH (5.0). The resulting solids are concentrated by
centrifugation and then dried to produce a mixture of stor-
age protein and nonstorage protein.
2. In the “selective precipitation” procedure the protein is
precipitated with alkali at a pH 10, the solids are then
Figure 3 Protein concentrate production from glandless cottonseed. (Data from Lusas EW and Jividen GM (1987) J. Am. Oil Chem. Soc. 64(6):
839–854.)
separated by centrifugation. The solids in the liquor are
also precipitated at pH 7, and the storage protein curd
removed by centrifugation and then dried. The liquor is
further acidified to pH 4 to precipitate the NSP, also
removed by centrifugation and then dried. The soluble
matter remaining in the whey is then removed by either
the selective extraction or selective precipitation method.
A relatively pure storage protein fraction, containing over
90% protein (dry weight basis), can be prepared.
3. The “selective extraction” procedure consists basically of
leaching the proteins soluble with water at neutral pH,
where the solids are then centrifuged out. The liquor is
then acidified to pH 4; this results in a precipitated protein
curd. The curd is dried to produce an NSP isolate. The solids
from the original water leaching are then solubilized in
alkali at pH 10 and centrifuged to remove insolubles. The
liquor is then precipitated at pH 7 and the resulting curd is
concentrated and dried to produce an SP isolate.
A patented process for the preparation of glandless cottonseed
protein concentrate from solvent-extracted flour using industrial
membranes has also been used. Defatted flour is sieved through
an 80-mesh screen to break up any agglomerates and remove
hull particles, and suspended in acidified water at pH 4.0–4.5.
This solution is passed through an ultrafiltration membrane
(100 000 MW cutoff) and the solids retained are either dried
directly or neutralized before drying. The liquid fraction perme-
ate may then be passed through a reverse osmosis membrane to
concentrate the soluble solids and to recover water which may
be reused in the process.
Feed Uses
Glandless cottonseed has been reported to be suitable for
monogastric animal feed not just for its lack of gossypol but
also for higher available energy. The lower heat treatment
required for glandless cottonseed during milling is reported
to degrade less protein than the prepress method normally

used with glanded cottonseed. Commercial glanded cotton-
seed meal intended for poultry feeding usually receives consid-
erable heat during processing to intentionally bind the
gossypol. Also in order to increase the amount of glanded
meal gossypol, addition of iron to the meal is used to inactivate
gossypol. In the cases of glandless cotton meal, it would be
expected that lower heat treatment needed for glandless cotton
would have beneficial effects in the protein availability and
quality of the meal. Some heat pretreatment of the meal is still
recommended, however, before hexane extraction, to improve
protein efficiency. Heating glandless cottonseed with 12%
added water for 10 min at 82  C, followed by 105  C for
20 min before hexane extraction, was reported to appreciably
increase broiler weight gains when compared with glanded
cottonseed feed. It has also been shown that meal from gland-
less cottonseed had an equal protein availability to soybean
meal in supporting chick growth. Feed trials with glanded
cottonseed meal with swine, catfish, fish, and shrimp have
also been described.
Exercises for Revision
• What is the meaning of “Glandless” in relation to
cottonseed?
• What is gossypol?
○ What natural advantage does gossypol offer to the
cotton plant?
○ Where is gossypol located in the cotton plant?
○ What is the toxicity of gossypol to humans, to animals
and to insects?
• Describe the ramifications of breeding glandless cotton for
the cotton-production industry, for the cotton-oil industry
and for the food and feed industries.
• Explain the relative advantages (and disadvantages) of
glandless cotton (versus glanded cotton) to all parts of the
cotton industry.
• List the food uses of cottonseed and compare these with the
food uses of the soybean.
Exercises for Readers to Explore the Topic Further
• Find out about the differences between the three major
species of cotton (Gossypium hirsutum L., G. aboreum L.,
G. barbadense L., G. herbaceum L.). What potential advan-
tages might accrue from the transfer of genes from one of
these to the more common cotton species?
• Consider the possibility of extracting gossypol from glanded
cottonseed for use as an insecticide for growing glandless
cotton.
• Research the uses and value of lecithin, and examine the
possibility of producing lecithin from glandless cottonseed.
• Find out about genetically modified (GM) cotton,
the nature of its consequent tolerance to insect attack, its
suitability for producing cotton, oil and meal, and the
possible public resistance to cotton as a genetically modi-
fied organism (“GMO”).
THE OILSEEDS | Cottonseed 5
See also: The Basics: Grain Crops, Overview (00001); Taxonomic
Classification of Grain Species (00002); Grain and Plants, Morphology
(00009); The Oilseeds: Oilseeds, Overview (00026); Soy, Overview
(00028); Grains around the World: Grain Production and
Consumption, Oilseeds in North America (00052); Food Grains and
the Consumer: Genetically Modified Grains and the Consumer
(00075); Fats: Lipid Chemistry (00099); Healthy Fats and Oils
(00100); Non-food Products from Grains: Biodiesel (00138);
Barley, Rice and Maize Processing: Oil from Rice and Maize
(00168); Soybean, Soy Concentrates and Isolates (00170); Soybean,
Processing (00172); Agronomy of Grain Growing: Soybean,
Agronomy (00199); Non-food Products from Grains: Animal Feed
(00231).
Further Reading
Berot S, Chenu M, Coutret J, and Gueguen J (1995) Cottonseed protein-rich products
from glandless African varieties: 1. Pilot-plant scale production of protein
concentrates, Sciences-des-Aliments 15(3): 203–215.
Camire ME, King CC, and Bittner DR (1991) Characteristics of extruded mixtures of
cornmeal and glandless cottonseed flour. Cereal Chemistry 68(4): 419–424.
Cater CM (1968) Processing glandless cottonseed. Oil Mill Gazeteer 72(7): 8–15.
Cherry JP, Gray MS, and Jones LA (1981) A review of lecithin chemistry and glandless
cottonseed as a potential commercial source. Seed-oil content of glanded and
glandless, cottons. Journal of the American Oil Chemists’ Society 58(10): 903–913.
Glandless cotton: Its significance, status, and prospects: Proceedings of a conference,
December 13–14, 1977, Dallas, Texas/sponsored by Agricultural Research Service,
United States Department of Agriculture and National Cottonseed Products
Association, Inc., Agricultural Research Service – Southern Region, US Dept. of
Agriculture, 1978.
Kohel RJ and Lee JA (1984) Genetic analysis of Egyptian glandless cotton. Crop
Science 24(6): 1119–1121.
Lawhon JT, Cater CM, and Mattil KF (1977) Journal of the American Oil Chemists’
Society 54: 75.
La-Rue DC, Knabe DA, Izard RS, and Tanksley TD (1986) Glandless cottonseed kernels
for starter, grower and finisher swine. Journal of Animal Science 64(4): 1051–1059.
Lusas EW and Jividen GM (1987) Glandless cottonseed: A review of the first 25 years of
processing and utilization research. Journal of the American Oil Chemists’ Society
64(6): 839–854.
Miravalle RJ (1972) The plant geneticist’s contribution toward changing the lipid and
amino acid composition of cottonseed. Journal of the American Oil Chemists’
Society 49: 24–26.
Percival AE, Wendel JF, and Stewart JM (1999) Taxonomy and Germ plasm resources.
In: Smith CW and Cothren JT (eds.) Cotton. New York: Wiley.
Robinson EH, Rawles SD, Oldenburg PW, and Stickney RR (1984) Effects of feeding
glandless or glanded cottonseed products and gossypol to Tilapia aurea.
Aquaculture 38(2): 145–154.
Ryan JR, Kratzer FH, Grau CR, and Vohra P (1986) Glandless cottonseed meal for laying
and breeding hens and broiler chicks. Poultary Science 65(5): 949–955.
Simmons RG and Golightly NH (1981) Cottonseed Cookery. College Station, TX: Food
Protein R&D Center, Texas A&M University.
Silva-Ramos LC and Kohel RJ (1987) Seed-oil content of glanded and glandless
cottons. Journal of the American Oil Chemists’ Society 64(9): 1337–1340.
Relevant Website
cottonseed.com http://www.cottonseed.com – National Cottonseed Products
Association (NCPA) is a trade association for the cottonseed processing industry.
Besides commercial information, it has further links to technical information on
cottonseed products.