998 CELLULOSE
complex behavior of animals is possible only because about 120 days. Under normal circumstances human
of this cellular specialization. All somatic cells (body
cells, as opposed to the gametes, or sex cells) contain
the full complement of genetic information. With
cellular specialization, however, the capacity to de-
velop in a variety of ways (sometimes called cellular
plasticity) is sacrificed. In some cases, cells become so
constrained and devoted to their special task in the
body that they even lose the ability to divide and
provide more of the same type. Such is the case with
neurons; these cells make up the main controlling and
information-transfer system of the body. Neurons
detect various environmental stimuli and pass this
information on to centers in the nervous system that
interpret this information. Having determined the
appropriate response, the nervous system sends mes-
sages to the muscles and glands of the body to effect
this response. All this is achieved by transmitting
electrical signals around the body. When neurons
are injured and die, they are not replaced. Muscle
cells, too, are incapable of dividing. These cells devote
their energies to building contractile machinery
within them. When these cells are activated, they
contract, bringing about movement of various
organs, limbs, etc., of the body. Adipocytes, or fat
cells, are specialized to store large amounts of fat to
such an extent that the nucleus of the cell becomes
confined to a small part of the cell squashed against
the cell membrane. It is thought that mature adipo-
cytes do not divide. Erythrocytes are packed with the
protein, hemoglobin, which is responsible for trans-
porting the blood gases, particularly oxygen. It
enhances the oxygen-carrying capacity of the blood
about 60-fold. Human erythrocytes are peculiar cells
in that they have no nucleus, losing their nucleus
during maturation. They have a limited lifespan of
CELLULOSE
M T Holtzapple, Texas A & M University, TX, USA
Copyright 2003, Elsevier Science Ltd. All Rights Reserved.
Background
0001 Cellulose is the world’s most abundant biological
material. An estimated 1011 tonnes are produced an-
nually, a daily production of about 45 kg per person.
Because of its rigidity, cellulose provides structure to
plants. It is almost always associated with other com-
ponents as a composite material termed ‘lignocellu-
lose.’ (See Hemicelluloses; Lignin.)
erythrocytes are produced in the body at a rate of
2  106 per second. Hepatocytes are specialized cells
of the liver. These cells contain large amounts of
specific enzymes involved in the metabolism and de-
toxification of many different molecules in the body.
These are just some of the many specialized cells
in the body; there are numerous others, including
osteocytes (bone cells), endocrine cells (hormone-
producing cells), and chondrocytes (cartilage cells).
(See Adipose Tissue: Structure and Function of White
Adipose Tissue; Structure and Function of Brown
Adipose Tissue.)
See also: Adipose Tissue: Structure and Function of
White Adipose Tissue; Structure and Function of Brown
Adipose Tissue; Exercise: Muscle; Fatty Acids:
Metabolism; Glucose: Function and Metabolism; Fats:
Classification; Nucleic Acids: Physiology; Protein:
Synthesis and Turnover
Further Reading
de Duve C (1984) A Guided Tour of the Living Cell, vols.
1 and 2. New York: Scientific American Library.
Fawcett DW (1981) The Cell, 2nd edn. Philadelphia, PA:
WB Saunders.
Ganong WF (1999) Review of Medical Physiology, 19th
edn. Upper Saddle River: Prentice Hall.
Keeton WT and Gould JL (1986) Biological Science, 4th
edn. New York: WW Norton and Co.
Martini FH (1998) Fundamentals of Anatomy and Physi-
ology, 4th edn. Upper Saddle River: Prentice Hall.
Reid RA and Leech RM (1980) Biochemistry and Structure
of Cell Organelles. Glasgow: Blackie.
Tribe MA, Morgan AJ and Whittaker PA (1981) The Evo-
lution of Eukaryotic Cells. London: Edward Arnold.
The most-studied celluloses are from cotton, ramie, 0002
Valonia algae and Acetobacter xylinum bacteria.
Structure
Polymer Structure
Cellulose is a polymer of the monomer glucose (see 0003
Figure 1a). The free-aldehyde glucose conformation is
very unstable, so it cyclizes into a six-membered pyra-
nose ring. (In aqueous solutions at 25  C, only about
0.02% of the glucose is in the free aldehyde form.) The
C1 hydroxyl of the cyclic ring can be in the equatorial b
 CELLULOSE 999

6
H H2COH
O
HO 4 O
1 CH H
H5
2 H C OH 2 OH 1
HO OH b
3 HO C H H3 H
4 H C OH
5 H C OH
1
6 H2COH H a
OH
(a)

1.038 nm

H H H H2COH H H H H COH
HO OH O O HO OH O
2 O
H H H H
H H H H
HO O HO OH O O HO OH OH
H H2COH H H H H2COH H H
n
Nonreducing Anhydrocellobiose Reducing
end end
DP = 2n + 2 (500 −15 000)
(b)

0.42 nm

O
H H2COH
H
H
H
O H OH
O H
HO
H
n H2COH
O Anhydroglucose
H OH

HO H

H
DP = n + 2 (100−3000) O
H H
H2COH H
HO
OH
H
HO
OH H
Nonreducing Reducing
end end
(c)

fig0001 Figure 1 (a) Glucose, (b) cellulose, and (c) amylose starch. Reproduced from Cellulose, Encyclopaedia of Food Science, Food Tech-
nology and Nutrition, Macrae R, Robinson RK and Sadler MJ (eds), 1993, Academic Press.

position or the axial a position. The b position is
favored thermodynamically and accounts for 62% of
the glucose with the remaining 38% in the a position.
0004 Cellulose is a linear, unbranched polymer of anhy-
droglucose joined with ether linkages between C1
and C4 (see Figure 1b). Cellulose is polymerized
polymerized from a-glucose. In the rightward anhy-
droglucose unit, C1 retains its hydroxyl group, so it
may potentially form free-aldehyde glucose, which
has reducing power. Hence, this terminus is called
the ‘reducing end.’
b-Linked cellulose has dramatically different 0005

from b-glucose, whereas starch (see Figure 1c) is properties than a-linked starch. Starch is helical,
 1000 CELLULOSE

water-soluble, and easily hydrolyzed by enzymes, unbranched, whereas amylopectin starch has
whereas cellulose is planar, water-insoluble, and diffi- branches at C6. (See Starch: Structure, Properties,
cult to hydrolyze. Whereas starch is a widely utilized and Determination.)
animal food, cellulose is used only by a select few,
Crystalline Structure
ruminants being the most prominent example. In
starch, the repeating unit is anhydroglucose. For The cellulose hydrogen atoms are all in the axial 0006

cellulose, anhydrocellobiose is the repeating unit
because adjacent anhydroglucoses are rotated 180 
with respect to their neighbors. This rotation causes
cellulose to be highly symmetrical, because each side
of the chain has an equal number of hydroxyl groups,
whereas starch is unsymmetrical. The cellulose degree
of polymerization (DP) ranges from 500 to 15 000.
When fully stretched, a single cellulose molecule
could extend as long as 7 mm. Amylose starch has a
lower DP (100–3000) and could extend as long as
1 mm if fully stretched. Cellulose is completely
position, whereas the hydroxyl groups are all equa-
torial. These equatorial hydroxyl groups can hydro-
gen-bond with their nearest neighbors, allowing
cellulose to crystallize. The monoclinic crystalline
unit cell for cellulose I (native cellulose) is shown in
Figure 2. The hydrogen bonds run in the a direction
and are medium-strength (15 kcal mol1). In the c
direction, the structure is held by weak van der
Waals forces (8 kcal mol1). Covalent bonds run in
the b direction and give cellulose its strength (50 kcal
mol1). A continuous cellulose strand is about four to

OH
H2COH OH

O O O

OH
OH H2COH
b = 838
OH
H2COH OH

O O O
c = 0.786 nm

OH
OH H2COH
OH
OH H2COH

O O

OH
H2COH OH

OH
H2COH OH

O O O

OH
OH H2COH

OH
H2COH OH

a=
0.8 O O O
17
nm
OH
OH H2COH

b = 1.038 nm

fig0002 Figure 2 Parallel cellulose I unit cell. Reproduced from Cellulose, Encyclopaedia of Food Science, Food Technology and Nutrition,
Macrae R, Robinson RK and Sadler MJ (eds), 1993, Academic Press.

five times stronger than steel with the same cross-
section. Cellulose I is parallel; that is, all the cellulose
molecules run in the same direction from nonredu-
cing to reducing ends (see Figure 3a).
0007 Native cellulose (cellulose I) can be converted to
other crystalline forms. Cellulose II is formed by (1)
treating cellulose with sodium hydroxide (merceriza-
tion), (2) precipitating from solutions of alkali/salt
(e.g., cuprammonium hydroxide), or (3) removing
the added functional groups from cellulose deriva-
tives (i.e., regenerated cellulose). Cellophane and
rayon are both forms of cellulose II. Table 1 shows
that the unit cell dimensions are slightly expanded in
the c direction and compressed in the a direction. Of
course, the b direction is essentially the same, because
it is the covalent bond. Cellulose II is the most
thermodynamically stable form of cellulose because
30−60 nm
Crystalline
Amorphous
(a) (b)
fig0003 Figure 3 (a) Cellulose I parallel and (b) cellulose II antiparallel
structures. Reproduced from Cellulose, Encyclopaedia of Food
Science, Food Technology and Nutrition, Macrae R, Robinson RK
and Sadler MJ (eds), 1993, Academic Press.
tbl0001 Table 1 Unit cell dimensions of cellulose I and II
Cellulose a (nm) b (nm) c (nm)
I 0.817 1.038 0.786
II 0.801 1.036 0.904
From Blackwell J, Kurz D, Su M-Y and Lee DM (1987) X-ray studies of the
structure of cellulose complexes. In: Atalla RH (ed.) The Structures of
Cellulose. ACS Symposium Series, No. 340, pp. 199–213. Washington, DC:
American Chemical Society.
CELLULOSE 1001
it can always be produced from cellulose I, but not
vice versa. The stability may result from hydrogen
bonds extending in the c direction, which normally
has only van der Waals bonds. There is general agree-
ment that cellulose II is antiparallel (see Figure 3b)
with three to four anhydroglucose moieties required
to make the bend. Precipitation of cellulose II from
solution appears to favor the antiparallel conform-
ation, as occurs with many synthetic polymers.
Cellulose III is formed by soaking cellulose in cold 0008
(about  80  C) liquid anhydrous ammonia, which
is subsequently removed by evaporation. Cellulose I
is transformed into cellulose III1 and cellulose II is
transformed into cellulose III2. When rehydrated,
cellulose III reverts back to its original form.
Cellulose IV is formed by soaking cellulose in hot 0009
(about 200  C) glycerol, with subsequent removal by
washing with 2-propanol and water. Cellulose I is
transformed to cellulose IV1, and cellulose II is trans-
formed to cellulose IV2.
Native cellulose (Figure 3a) forms crystalline 0010
regions (40% bacteria, 60% cotton, 70% Valonia)
interspersed with amorphous regions. The amorph-
ous regions are more porous than the crystalline
regions, allowing water or dyes to penetrate and in-
creasing the reactivity to acid or enzymatic hydroly-
sis. When purified cellulose fibers are subjected to
~15 nm
dilute acid hydrolysis, the amorphous regions select-
ively hydrolyze, leaving the more resistant crystalline
regions, which have a ‘levelling-off DP’ of 100–300 in
the case of cotton.
Cellular Structure
Young plant cells have only the primary wall to resist 0011
osmotic pressure. This wall is composed principally
of hemicellulose and pectin with small amounts of
cellulose and protein and is sufficiently flexible to
accommodate cell growth. When cell growth ceases,
the secondary wall is formed in three layers (S1, S2,
and S3) with S2 the thickest (see Figure 4). (See Pro-
tein: Chemistry.)
All cell wall layers are composed of 7–30-nm- 0012
diameter microfibrils, which are visible using an elec-
tron microscope. In the primary cell wall, the fibrils
are randomly oriented. In the S1 layer, the microfibrils
are oriented helically, with each successive layer alter-
nating between right-handed and left-handed helices.
In the S2 layer, which provides much of the plant
b ( ) strength, the microfibrils are arranged in steep right-
handed helices nearly oriented along the cell axis. The
83.0
62.9 S3 layer is another shallow helix. All higher plants
(softwoods, hardwoods, herbaceous) are thought to
have cell structures similar to that shown in Figure 4.
The plant cell diameters range from 15 to 80 mm, 0013
depending on the species and time of year. Spring cell
 1002 CELLULOSE

Microfibril

S3

Lignin
Secondary Hemicellulose
wall
S2

S1
Primary wall

7−30 nm
Middle lamella
Compound
middle
lamella

15 − 8
0 mm

m Elementary
60n
30− fibril
3−5 nm

Unit cell

0.786 nm
0.817 nm

3−10 nm

fig0004 Figure 4 Schematic of plant cell, microfibril, and elementary fibril. Reproduced from Cellulose, Encyclopaedia of Food Science, Food
Technology and Nutrition, Macrae R, Robinson RK and Sadler MJ (eds), 1993, Academic Press.

diameters are larger than summer diameters, giving
rise to the familiar growth rings in trees. The wall
thickness is typically about 2–5 mm, with thicker
walls formed in the summer. Softwood cells are
about 3–4 mm long, which makes them particularly
useful for paper pulp, whereas hardwood cells tend to
be shorter (0.7–2 mm long). Cotton cells suitable for
textiles are about 25 mm long.
0014 Microfibrils are composed of elementary fibrils of
pure cellulose embedded in a matrix of hemicellulose.
The fringes of the elementary fibrils are paracrystal-
line and intermingle with tightly adsorbed hemicellu-
lose. (An exception to this is cotton in which the
fringes contain only cellulose.) In Fengel’s model of
the microfibril, the elementary fibrils are clustered
into four 4  4 arrays. The lignification process
occurs late in cell life, so lignin is located primarily
on the microfibril exterior, where it covalently bonds
to the hemicellulose. Lignification is initiated in the
middle lamella where lignin constitutes about 70% of
 CELLULOSE 1003

the wall. The wall interior is less lignified, with only cellobiose to glucose). Up to about 10% of cellulose
about 15% lignin. The hemicellulose content is fairly is enzymatically digested by microbes in the human
constant (20–30%), with cellulose the remaining 10– large intestine, so it is not completely noncaloric.
50% of the wall. The cells in fruit pulp are composed Cellulose is fairly stable to base, provided that 0018

mainly of primary walls containing approximately oxygen is excluded. The reaction terminates after
34% pectin, 24% hemicellulose, 23% cellulose, and approximately 50 anhydroglucose units are ‘peeled
19% protein. off’ from the cellulose reducing end; d-glucoisosac-
charinate is the soluble product. Under severe condi-
Properties tions (e.g., 1 M sodium hydroxide, 170  C), alkaline
hydrolysis readily occurs.
Physical Cellulose is more stable to oxidants than lignin, a 0019

property exploited in pulp bleaching and analytical

0015 Table 2 shows the physical properties of cellulose and
cellulosic materials. The listed heat of combustion is
the ‘higher heat’ (i.e., the combustion water is 25  C
liquid). Cellulose is stable up to 300  C.
0016 The DP for various types of cellulose is shown in
Table 3. For cotton, the DP may be as high as 15 000.
Cellulose is insoluble in water because of the strong
hydrogen bonding in its crystal lattice. However, it is
soluble in a number of solvents, including concen-
trated acids (e.g., 85% phosphoric, 72% sulfuric,
and 40% hydrochloric acids) and inorganic salt solu-
methods, which selectively oxidize lignin. However,
oxidants (e.g., chromic acid, permanganate, and
hypochlorite) can damage cellulose by cleaving the
chain or by inserting carbonyl functional groups.
The three cellulose hydroxyl groups are very 0020
reactive, allowing ready formation of the cellulose
derivatives described later.
Specialty Celluloses
Microcrystalline cellulose (Avicel, FMC Corporation) 0021

tions (e.g., cuprammonium hydroxide, cadoxen (i.e.,
cadmium ions and ethylenediamine)). The viscosity of
cellulose dissolved in salts is often used for molecular
weight determinations.
Chemical
0017 Cellulose may be hydrolyzed to glucose by acids or
is prepared by acid hydrolysis of cellulose using 2 M
hydrochloric acid at 105  C for 15 min. The highly
reactive amorphous regions selectively hydrolyze,
releasing the crystallites, which are subsequently
Table 3 Cellulose polymer length tbl0003

enzymes. Acid hydrolysis produces a number of deg- Material Degree of polymerization Molecular weight
radation products, such as 5-hydroxymethylfurfural,
Native cellulose 3500–10 000 600 000–1 500 000
formic acid, levulinic acid, formaldehyde, furfural, Chemical cottons 500–3000 80 000–500 000
and resins. Enzymatic glucose production requires a Wood pulps 500–2100 80 000–340 000
cellulase system containing endo-cellulase (to pro- Rayon filaments 350–450 57 000–73 000
duce nonreducing ends from the chain interior),
From Hamilton JK and Mitchell RL (1964) Cellulose. In: Standen A (ed.)
exo-cellulase (to produce cellobiose generally from Kirk^Othmer Encyclopedia of Chemical Technology, 2nd edn., vol. 4, pp.
the nonreducing end), and cellobiase (to hydrolyze 593–616. New York: Wiley.

tbl0002 Table 2 Physical properties of cellulose and cellulosic materials

Property Compressed cellulose Paper Pine Oak

Specific gravitya 1.47 0.70–1.15 0.43–0.67 0.64–0.87

Heat capacity (kJ kg1 K1)b 1.3 1.3 2.8 2.4
Heat of combustion (MJ kg1)b 17.6 17.6 20.4 19.2
Thermal conductivity (W m1 K1)b
With grain 0.13 0.35
Across grain 0.13 0.15 0.21
Tensile strength (MPa)a
With grain 62 99
Across grain 2.8 5.5
Water diffusion constant (cm2 s1)c 5.3  107 1.9  107
a
From Baumeister T, Avallone EA and Baumeister T, III (1978) Mark’s Standard Handbook for Mechanical Engineers, 8th edn. New York: McGraw-Hill.
b
From Graboski M and Bain R (1981) Properties of biomass relevant to gasification. In: Reed TB (ed.) Biomass Gasification: Principles and Technology,
pp. 41–71. Park Ridge: Noyes Data.
c
From Summitt R and Sliker A (1980) CRC Handbook of Materials Science: Wood, vol. IV, pp. 26–27. Boca Raton, FL: CRC Press.
 1004 CELLULOSE
mechanically dispersed. Aqueous suspensions of
microcrystalline cellulose have constant viscosities
over a wide temperature range, are heat-stable, and
have good mouth-feel properties. Avicel is used to
extend starches, stabilize foams, and control ice crys-
tal formation. Avicel has found widespread accept-
ance in the food industry for meringue, whipped
toppings, confections, and ice cream, and is also used
as a binder in pharmaceutical tablets and in cosmetics.
0022 Bacterial cellulose (Cellulon, Weyerhaeuser Co.) is
produced by selected strains of Acetobacter xylinum,
which maintain their ability to produce cellulose in
agitated submerged fermentors. The cellulose fibers
are about 0.1 mm in diameter, which is substantially
smaller than softwood pulp fibres (about 30 mm
diameter). Cellulon is a potential noncaloric food
thickener or texturizer.
Modified Celluloses
0023 Alkali cellulose is prepared by soaking cellulose
in concentrated sodium hydroxide (> 14%) in the ‘mer-
cerization’ process invented by John Mercer in 1844.
The sodium ions are incorporated into the cellulose
structure according to the following reaction (eqn (1)):
Rcell OH þ NaOH ! Rcell ONa þ H2 O: ð1Þ
Alkali cellulose
The cellulose structure swells, allowing easy penetra-
tion by dyes or reagents for the manufacture of cellu-
lose derivatives.
0024 Cellulose xanthate is formed by reacting alkali
cellulose with carbon disulfide (eqn (2)):
S Ethyl cellulose is produced by reacting alkali cellu-
k lose with ethyl chloride. In commercial products, the
Rcell ONa þ CS2 ! Rcell O CSNa: ð2Þ
Cellulose xanthate
0025 Regenerated cellulose is made by dissolving cellulose
xanthate in 4–7% sodium hydroxide and contacting
with aqueous sulfuric acid. These steps convert the
cellulose xanthate back into cellulose, which may be
spun into viscose rayon or cast into films. The fibers
are used in textiles (artificial silk), tyre cords, and V
belts. The films are used in packaging (Cellophane) or
sausage casings. Weiner casings (70% regenerated
cellulose, 12% glycerol, and 18% water) are peeled
away after the meat emulsion is cooked. Hemp paper
casings (23% paper, 46% regenerated cellulose, 21%
glycerol, and 10% water) are used in bologna, salami,
pepperoni, summer sausage, and liverwurst.
0026 Cellulose hydroxyl moieties are highly reactive,
allowing a variety of esters and ethers to be manufac-
tured. Because each anhydroglucose has three hydroxyl
groups, the maximum degree of substitution (DS) is
three. Purified wood pulp or cotton linters (short fibers)
are the industrial sources of ‘chemical cellulose.’
Cellulose Ethers
Sodium carboxymethyl cellulose (CMC) is formed by 0027
reacting sodium chloroacetate with alkali cellulose
(eqn (3)):
O
k
Rcell ONa þ CICH2 CONa !
ð3Þ
O
k
Rcell OCH2 CONa þ NaCl:
CMC
Commercially available CMC has a DS range of
0.38–1.4, with 0.65–0.85 more common. The nega-
tively charged carboxyl group makes CMC soluble in
both hot and cold water. The solution viscosity de-
creases as the temperature increases. CMC is ‘gener-
ally recognized as safe’ and is used as a thickener in
many foods such as cheese, frozen desserts, and salad
dressings. It is not metabolized, so it is used in low-
calorie foods.
Methyl cellulose is formed by reacting alkali cellu- 0028
lose with methyl chloride (eqn (4)):
Rcell ONa þ ClCH3 ! Rcell OCH3 þ NaCl: ð4Þ
Methyl cellulose
Methyl cellulose (DS 1.8) solutions form a firm gel
when heated to 50–55  C and return to solution when
cooled. Methyl cellulose is added to salad dressings,
jams and preserves, soda water, and meat patties as a
binder.
0029
DS ranges from 2.0 to 2.6. It is water-insoluble and
may be incorporated into inks used for marking foods
and in vitamin tablet binders.
2-Hydroxypropyl methyl cellulose is formed by 0030
reacting alkali cellulose with mixtures of methyl
chloride and 2-hydroxypropyl chloride. It forms gels
like methyl cellulose, but has a higher gelation tem-
perature. It may be used as an emulsifier, film former,
stabilizer, or thickener in foods such as salad dress-
ings, sherbet, pie fillings, fried foods, whipped top-
pings, breading batters, and baked goods.
2-Hydroxyethyl cellulose (HEC) is produced by 0031
reacting cellulose with ethylene oxide using a sodium
hydroxide catalyst at 30–35  C for about 4 h (eqn (5)):
O
RcellOH ⫹ CH2 CH2 RcellOCH2 CH2OH.
HEC (5)
Because the side-chain also has a hydroxyl group,
ethylene oxide can continue to react and form a

side-chain with several units. HEC is soluble in both
hot and cold water. The solution viscosity decreases
as the temperature increases. HEC is not permitted as
a direct food additive, but it may be used in food-
packaging adhesives and coatings.
0032 2-Hydroxypropyl cellulose is produced using pro-
pylene oxide, rather than the ethylene oxide used for
HEC. It has a thermal gel point, like methyl cellulose,
and is used in food coatings and glazings.
Cellulose Esters
0033 Cellulose acetate is the most important cellulose ester.
The cellulose is first ‘activated’ in aqueous acetic acid
to ensure uniform acetylation. It is then dehydrated
and reacted with acetic anhydride using a catalyst
(e.g., sulfuric acid) in a solvent (e.g., anhydrous acetic
acid) (eqn (6)).
O O
k k
Rcell OH þ CH3 CO CCH3 !
ð6Þ
O O
k k
Rcell O CH3 þCH3 COH:
Cellulose acetate
The resulting cellulose triacetate (DS  3) product
may be subsequently acid-hydrolyzed to lower
the DS. Cellulose triacetate is water-insoluble and
hydrophobic, whereas cellulose monoacetate is water-
soluble. Cellulose acetate is used in fibers, plastics,
photographic films, lacquers and reverse osmosis or
dialysis membranes.
0034 Other esters (e.g., cellulose formate, cellulose pro-
pionate, cellulose butyrate) may be formed, but they
do not have the widespread commercial applications
of cellulose acetate. Also, mixed esters (e.g., cellulose
acetate propionate, cellulose acetate butyrate, cellu-
lose acetate phthalate) may be produced.
0035 Cellulose nitrate is made by reacting cellulose with
nitric acid/sulfuric acid for 20–30 min. The acids are
then removed by washing with water. The water must
be removed with great care because dry cellulose
nitrate is extremely explosive. It is often shipped
wetted with water or alcohol. Highly nitrated (DS
2.4–2.6) forms are used as explosives (gun cotton).
Less nitrated forms (DS 2.1–2.3) find applications in
plastics, films, and inks.
Composition of Cellulosic Materials
0036 Table 4 shows the composition of many cellulosic
materials. The cellulose content in vegetables ranges
from 1 to 21%; fruits range from 0.6 to 4.2%; seeds
range from about 2 to 12%; agricultural residues
range from 31 to 59%; wood ranges from 41 to
53%; flax and hemp have about 70% cellulose; and
CELLULOSE 1005
cotton (the purest natural cellulose) has about 95%
cellulose. The free sugars shown in Table 4 are the
sum of glucose, fructose, and sucrose. The ‘crude
fiber’ is reported as the sum of cellulose and lignin
in the case of the seeds. The indigestible portion of
some fruits, called ‘dietary fiber,’ is reported as
the sum of cellulose, lignin and hemicellulose. (See
Dietary Fiber: Properties and Sources; Fructose;
Sucrose: Properties and Determination.)
Cellulose Isolation and Analysis
Gravimetric Methods
Van Soest procedure The raw plant material is con- 0037
tacted with dilute acid and washed with solvents to
remove starch, pectin, hemicellulose, fats, oils,
protein, free sugars, and soluble minerals. The resi-
due, called ‘acid detergent fiber,’ contains cellulose,
lignin, and insoluble minerals (mainly silica).
Air-dried plant material (1 g), ground to particles of 0038
less than 1 mm, is placed in a beaker with 100 ml of
acid-detergent solution (49 g l1 H2SO4, 20 g l1 cetyl
trimethylammonium bromide). Decahydronaphtha-
lene (2 ml) is added, and the contents are slowly
boiled for 1 h while the reflux condenser maintains a
constant liquid volume in the beaker. Then, the
beaker contents are filtered through a tared Gooch
crucible, washed with boiling water, acetone, and
hexane (optional). The residual acid detergent fiber
(ADF) is dried at 100  C and weighed.
The cellulose content in ADF can be measured by 0039
two methods: (1) cellulose removal with acid or (2)
lignin removal by oxidation.
Method 1 The ADF prepared above is placed in a 0040
tared Gooch crucible containing an equal volume of
asbestos as a filter aid. The crucible contents are
contacted with room-temperature 72% sulfuric acid
for 3 h and then thoroughly washed with hot water.
The sample is dried at 100  C for 8 h and weighed.
The weight loss corresponds to the cellulose content.
Method 2 The ADF (0.5–1.0 g) is placed in a tared 0041
Gooch crucible. A saturated potassium permanganate
solution (50 g l1 KMnO4, 0.05 g l1 AgSO4) and
buffer solution (6 g l1 Fe(NO3)39H2O, 0.15 g l1
AgNO3, 500 ml l1 glacial acetic acid, 5 g l1 potas-
sium acetate, 400 ml l1 t-butyl alcohol) are mixed in
a 2:1 ratio (by volume). This mixture (25 ml) is added
to the crucible for 90 min at room temperature to
oxidize the lignin in the ADF. The spent reagent is
then removed by vacuum filtration. The crucible con-
tents are soaked for 5 min in demineralizing solution
 1006 CELLULOSE

tbl0004 Table 4 Composition of cellulosic materials (g per 100 g of dry matter)

Cellulose Lignin Hemicellulose Pectin Starch Free sugars

Vegetablesa
Leafy
Broccoli 7.2 0.26 24 1.7 15
Brussels sprouts 9.04 2.1 26 8.7 18
Cabbage 8.9 4.3 26 4.3 47.3
Cauliflower 13.4 Tr 13 5.0 29
Lettuce 20.6 Tr 9.2 0 17
Legumes
Beans, haricot 5.3 0.9 22.0 51.3
Beans, runner 17 3 21 2.9 39.6
Peas 14 2 36 66 6.9
Root
Carrot 12.9 Tr 19 0.9 24.8
Turnip 11 Tr 23 Tr 47.7
Fruiting vegetables
Pepper 3.5 Tr 10 Tr 34.2
Tomato 9.1 5.3 11 Tr 46.4
Tuber
Potato 1.2 Tr 9.2 84.6 1.5
Fruitsa
Apples 2.9 Tr 5.8 2.3 1.8 63.9
Apricots 15 3.3 0 56.6
Banana 1.3 0.93 3.83 10.4 55.8
Blackberries 44 1.9 0 34.1
Cherries, sweet 1.2 0.3 4.5 0.4 0 65.4
Grapefruit 0.6 0.9 4.9 0 67.7
Lemons 35 3.4 0 21.1
Oranges 14 3.3 0 59.6
Peaches 1.8 5.1 12.2 3.3 0 74
Pears 4.2 2.7 8.2 Tr 63.5
Pineapples 7.64 0.25 0 74.1
Strawberries 3.6 8.4 10 3.5 Tr 61.8
Seedsb
Barley 5.3 64.1
Corn 2.4 71.8 1.9
Grain sorghum 2.7 2.5 70.2 1.4
Oats 11.9 60.6
Peanut 2.8 2.5 4.0 4.7
Wheat 2.1 74
Agricultural residuesc
Barley straw 44 7 27
Cottonseed hulls 59 13 15
Oat straw 41 11 16
Rice straw 33 7 26
Sorghum straw 31 11 30
Sugarcane bagasse 40 13 29
Wheat straw 39 10 36
Treesd
Hardwood
Aspen 53.3 16.3 26.2
White birch 41.0 18.9 36.2
Red maple 44.1 24.0 29.2
Softwood
Balsam fir 44.8 29.4 23.6
Jack pine 41.6 28.6 25.6
White spruce 44.8 27.1 26.1
Bast fiberse
Flax 71.2 2.2 18.6 2.0
Leaf fiberse
Manila hemp 70.2 5.7 21.8 0.6
Seed fiberse
Crude cotton 95.3 0 0 1.0

(50 g l1 oxalic acid dihydrate, 700 ml l1 95% etha-
nol, 50 ml l1 12 M HCl), which is then removed by
filtration. The filter contents are washed with ethanol
and acetone. The crucible is dried at 100  C for 8 h.
The crucible contains cellulose and insoluble ash. The
crucible may be placed in a 500  C oven for 3 h; the
weight loss corresponds to the cellulose content.
0042 A simpler approach to removing the lignin from the
ADF has been described by C.S. Edwards. The ADF is
soaked in activated trigol (i.e. 6.3 ml of 32% hydro-
chloric acid dissolved in 1 l of pure triethylene glycol)
in a 121  C autoclave for 60 min. The sample is then
washed with 95% ethanol and acetone. The residue
contains cellulose and insoluble ash.
0043 Other gravimetric methods Cross and Bevan cellu-
lose is the lignocellulose portion remaining after
removing hemicellulose with two boilings in NaOH
and removing lignin with chlorine and bleach.
0044 Monoethanolamine cellulose is the lignocellulose
portion remaining after monoethanolamine treat-
ment, chlorination, and bleaching.
0045 Norman–Jenkins cellulose is the lignocellulose por-
tion remaining after sodium sulfite boiling, bleaching,
and acid treatment.
0046 a-Cellulose is the cellulose fraction insoluble in
room temperature 17.5% sodium hydroxide wash
water (DP > 90).
0047 b-Cellulose is the cellulose fraction in the alkaline
wash water that precipitates upon neutralization
(15 < DP < 90).
0048 g-Cellulose is the cellulose fraction soluble in the
neutralized alkaline wash water (DP < 15).
0049 Holocellulose is the residue that remains after lig-
nocellulose is defatted with hot, azeotropic benzene/
ethanol and delignified with hot chlorous acid
(HClO2). Holocellulose contains the cellulose and
hemicellulose of the original plant material.
Colorimetric Method
0050 Hexosan assays may be used to measure cellulose, the
dominant hexose polymer in starch-free plant mater-
ials (other than softwoods). Holtzapple describes an
assay in which the hexosan sample is placed in a
sealed test tube with 1530 g l1 sulfuric acid and
20 g l1 chromotropic acid, and boiled for 1 h. Hexose
C6 degrades to form formaldehyde, which reacts with
the chromotropic acid to form a purple compound
that is measured spectrophotometrically. There is
Tr, trace.
a
From Southgate DAT (1976) Determination of Food Carbohydrates, pp. 91–93. London: Applied Science.
b
From Zaborsky OR (1981) CRC Handbook of Biosolar Resources, vol. II. Boca Raton, FL: CRC Press.
c
From Marsden WL (1986) CRC Critical Reviews in Biotechnology, vol. 3, issue 3, p. 235. Boca Raton, FL: CRC Press.
d
From Timell TE (1957) Carbohydrate composition of ten North American species of wood. TAPPI 40: 568.
e
From Turbank AF, Durso DF, Battista OA et al. (1979) Cellulose In: Kirk^Othmer Encyclopedia of Chemical Technology, 3rd edn., vol. 5. New York: Wiley.
CELLULOSE 1007
minor interference by pentosans that may be reduced
by lowering the chromotropic acid concentration to
1 g l1 and shortening the reaction time to 20 min.
There is also some interference with lignin, so the
most accurate results are obtained with holocellulose,
rather than the original lignocellulose. (See Spectros-
copy: Visible Spectroscopy and Colorimetry.)
Chromatographic Method
The cellulose content can be estimated from the 0051
glucose composition, because cellulose is the main
source of glucose in plant materials (assuming free
sugars and starch are not present). (See Chromatog-
raphy: Principles.)
See also: Chromatography: Principles; Dietary Fiber:
Properties and Sources; Fructose; Hemicelluloses;
Lignin; Protein: Chemistry; Spectroscopy: Visible
Spectroscopy and Colorimetry; Sucrose: Properties and
Determination
Further Reading
Atalla RH (1987) The Structures of Cellulose. ACS Sympo-
sium Series, No. 340. Washington, DC: American
Chemical Society.
Bogan RT, Kuo CM and Brewer RJ (1979) Cellulose deriva-
tives, esters. In: Kirk–Othmer Encyclopedia of Chemical
Technology, 3rd edn., vol. 5, New York: Wiley.
Edwards CS (1973) Determination of lignin and cellulose in
forages by extraction with triethylene glycol. Journal of
the Science of Food and Agriculture 24: 381–388.
Fengel D (1970) Ultrastructural behavior of cell wall poly-
saccharides. Tappi 53(3): 497–503.
Goering HK and Van Soest PJ (1970) Forage Fiber Analysis.
Agricultural Handbook, No. 379, Jacket No. 387–598.
Washington, DC: US Department of Agriculture.
Greminger GK (1979) Kirk–Othmer Encyclopedia of Chem-
ical Technology, 3rd edn., vol. 5. New York: Wiley.
Holtzapple MT and Humphrey AE (1983) Determination
of soluble and insoluble glucose oligomers with chromo-
tropic acid. Analytical Chemistry 55: 584–585.
Jayne G (1971) Cellulose and cellulose derivatives. In:
Bikales NM and Segal L (eds) High Polymers, 2nd
edn., vol. V, part IV. New York: Wiley Interscience.
Plunguian M (1943) Cellulose Chemistry. New York:
Chemical Publishing.
Southgate DAT (1976) Determination of Food Carbohy-
drates. London: Applied Science.
Turbank AF, Durso DF, Battista OA et al. (1979) Cellulose.
In: Kirk–Othmer Encyclopedia of Chemical Technology,
3rd edn., vol. 5. New York: Wiley.