Dietary Fiber: Properties and Sources
R Mongeau and SPJ Brooks, 3W Banting Research Centre, Ottawa, QC, Canada
ã 2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the Encyclopedia of food sciences and nutrition, vol. 3, pp. 1813–1822, ã 2003, Elsevier Science Ltd., with an
updated Bibliography section supplied by the editor.
Background
Although dietary fiber has been known for more than 2000
years under various terms (e.g., bran and roughage), the term
‘dietary fiber’ first appeared in 1953 and referred to
hemicellulose, cellulose, and lignin. The term ‘fiber’ is some-
what misleading since only a fraction (cellulose) of dietary
fiber is fibrillar in nature. To correct this misnomer, other
terms (e.g., plantix) have been proposed, but despite these
efforts, the term ‘dietary fiber’ has survived.
This section deals with several different aspects of dietary
fiber, including its determination and many of its physiologi-
cal effects. Most of this information is relatively new and is the
result of progress in research brought about in the last 30 years.
The first part of this article will focus on the nature and com-
position of dietary fiber, its properties, and examples of sources
of dietary fiber. This is followed by a definition of dietary fiber.
Although this should be a relatively straightforward descrip-
tion, there are many different viewpoints concerning the
nature and physiological effects that dietary fiber should
have. These will be discussed in relationship to the physiolog-
ical effects of dietary fiber.
Chemical Structure
The composition and structure of dietary fiber differ from plant
to plant. It is also a function of the portion of the plant that is
edible and the stage of maturation and is largely composed of the
cell wall (structural) components that give the plant physical
stability. As such, it is made of highly interlinked sugar-based
and phenolic-based polymers (hemicelluloses, pectic substances,
phenolics, glycoproteins, and proteoglycans) in a matrix of amor-
phous structure with some enmeshed cellulose microfibrils. The
cell wall components are intimately linked together through
various linkages including protein–sugar bonds.
Polysaccharide Fiber Components
Dietary fiber comprises carbohydrate and noncarbohydrate
polymers; most of these are structural components. The carbo-
hydrate polymers are often described as nonstarch polysaccha-
rides (NSPs) to differentiate them from the (relatively) easily
digested starch components of food. Figure 1 shows the pro-
portion of the monomeric constituents of fiber polysaccha-
rides from three food categories (11 cereals, 13 fruits, and 11
vegetables) and from canned mushrooms, peanuts, and baked
white beans. As can be seen, arabinose is a major NSP compo-
nent of cereals, peanuts, and beans. Xylose is present in large
amounts only in cereals. Pectic substances (uronic acids) rep-
resent a substantial part of fruit, vegetable, peanut, and bean
NSPs. Mushroom NSP is formed mostly of glucose residues.
404 Encyclopedia of Food and Health
These results were based on the analysis of composites made of
up to 20 individual foods purchased over a 30-month period
using the Englyst gas–liquid chromatography method. A more
recent version of this method would have given a higher pro-
portion of xylose relative to glucose.
The different monomeric residues of Figure 1 are the basic
units that form the major NSP macromolecular constituents
shown in the upper part of Table 1. Cellulose and b-glucans are
generally unbranched polymers containing mostly glucose res-
idues. The b-glucans have mixed b-1,3 and b-1,4 interresidue
linkages and are a constituent of dietary fiber from barley and
oats. The glucose residues of cellulose are joined by b-1,4
linkages. In addition to their resistance to the enzymes of the
human digestive tract, these linkages allow greater inter- and
intrapolymer hydrogen bonding than the a-1,4 linkages of
starch do. This allows the formation of tightly condensed
crystalline regions within the cellulose microfibrils. Cellulose
includes structurally amorphous regions where the presence of
other sugars can be found. The final cellulose structure is a
function of the glucose and nonglucose contents: the degree of
cellulose crystallization is inversely proportional to the non-
glucose residue content. Cellulose is a minor fraction of the
total cereal dietary fiber but is a major fraction of dietary fiber
in other foods. The proportion of cellulose in the dietary fiber
of a food is not reflected by the glucose content of the NSP
fraction, since glucose is also a major component of other
fractions such as b-glucans and hemicelluloses.
Xylose is a part of the primary chain of hemicelluloses and
gums and the secondary chain of gums and pectin. Arabinose
is in the primary chain of gums and the secondary chain of
hemicelluloses and pectin. Hemicelluloses constitute a major
fraction of dietary fiber; this fraction is largely made of arabi-
nose and xylose in cereals and of xylose and glucose in fruits
and vegetables (Table 1). Pectic substances are prevalent in
citrus fruits. They are also present in vegetables, legumes, and
cereals in small amounts (Table 1). Other polysaccharide con-
stituents are mainly represented by galactose or mannose in
bananas. Nonstructural NSPs represent a small portion of
dietary fiber. The monomeric NSP composition is of limited
usefulness in predicting the properties and functions of dietary
fiber. This is because the structural architecture of dietary fiber
plays a large role in many of its physiological properties, and
this structure cannot be predicted by its composition. Indeed,
two dietary fibers with a similar component profile (e.g., wheat
bran and corn bran) can have clearly different structures, even
under optic microscopy, and can show different behaviors in
the gastrointestinal tract.
Nonsaccharide Fiber Components
The minor components of dietary fiber that are not polysac-
charides and, hence, not NSPs also play an important
http://dx.doi.org/10.1016/B978-0-12-384947-2.00784-4
 Dietary Fiber: Properties and Sources 405

tissues. In most fruits and vegetables, lignin represents only a

100
small part of dietary fiber, although it can be as high as 4% of
dry matter in mature pears.
Cutin, waxes, or suberin may also be present in some
Distribution (%)

tissues. These compounds are mixtures of lipids, proteins,

and carbohydrates that form the waterproof covering and cuti-
50 cle on the outer cell wall of plants. They are especially resistant
to digestion and fermentation. As such, they impair the digest-
ibility of other cell wall components and are usually found in
the feces.

Mushrooms
Fruits

Peanuts

White Beans
Vegetables
Cereals

Figure 1 Distribution of the main polysaccharide constituents of dietary
fiber from some foods and food categories. The values are means of
n ¼ 11 cereals, n ¼ 13 fruits, and n ¼ 11 vegetables. Other values are
for n ¼ 1. ▪: glucose; : arabinose; : xylose; : uronic acid; and □:
other sugars.
Table 1 Main macromolecular constituents of dietary fiber
Fruits and
Constituent vegetables
Polysaccharides
Hemicelluloses
Xyloglucans X X
Glucuronoxylans X
Arabinoxylans
Glucuronoarabinoxylans
Galactomannans
Cellulose X X
b-D-Glucans
Pectic substances (pectin) X X
Others
Lignin
Phenolic esters X X
Protein
Glycoproteins X X
structural role. Table 1 shows the distribution of the nonsac-
charidic polymers that are intimately associated with dietary
fiber structure. These polymers include glycoproteins (fruits,
vegetables, and legumes), proteins (cereals), phenolic esters
(cereals), and lignins (lignified tissues of fruits, vegetables,
and cereals). Lignin is of special interest because of its role in
slowing down the fermentation of dietary fiber. It is a complex
group of phenyl propane polymers formed by the condensa-
tion of aromatic alcohols and is especially important in con-
ferring structural stability. As such, highly lignified tissues are
found in the stems of plants such as trees and bushes and in the
stalks of cereals. Lignification occurs at the expense of water
and pectin as cells differentiate and mature. This increases cell
wall rigidity in critical areas. Since edible plant tissues are
consumed when relatively immature, their cells are largely
undifferentiated and, largely, unlignified. Wheat bran and the
small seeds covering strawberries are examples of lignified
Structure
Models of primary cell walls of plants were often obtained from
nonfood materials such as wood and tobacco. Because the cell
wall material forms a large part of these plants, the models
provide a wealth of information on the interactions between
cell wall components such as carbohydrates and lignin and
protein and carbohydrates. More recent reports have provided
some structural information on the cell walls of plant foods.
These reports have shown that, in carrots, protein–protein or
protein–polysaccharide linkages may contribute to the forma-
tion of a rigid, inextensible cell wall. Other data have shown that
protein may form up to 10% of cell walls in immature plants
and appears to be important for structural cohesion. The major
Cereals Legumes part of the flesh of fruits and vegetables contains undifferen-
tiated types of cell wall that have significant amounts of highly
branched hemicellulosic types of polysaccharides. The cellulose
content is usually low, and cellulose is laid down in a more
oriented arrangement in the matrix. This may reflect stresses on
the plant. The cell walls are often rich in pectic substances and
X
X
contain
10% protein, as described earlier. These cell walls are
X thin and elastic with the cell form maintained by osmotic
X pressure.
X In contrast, the outer layers of many seeds and nuts contain
X thick lignified cell walls that are difficult to break. These serve
as mechanical protection to the seed within and as a vapor
X barrier to prevent desiccation.
X
Physical Properties
Soluble and Insoluble Dietary Fibers
It is possible to separate the total dietary fiber (TDF) into
soluble and insoluble components based on solubility in
water, but quantifying the amount of soluble fiber remains
problematic. This is because the conditions utilized to measure
the proportion of soluble fiber differ among laboratories (e.g.,
time, temperature, pH, and type of buffer; see the section
‘Definition (Based on Function and Structure)’). It should
be noted that the distinction between soluble and insoluble
fibers is somewhat artificial, since it is difficult to predict the
actual fiber solubility in the gastrointestinal tract.
For many years, it was thought that soluble fiber was not
commonly found in foods. Reports on pectin, guar, legume
fiber, and oat bran appeared in the early 1960s, and currently,
it is estimated that about a third of the daily TDF intake is
soluble fiber. Pectic polyuronides are the major soluble dietary
fiber components of vegetables and fruits, but the exact com-
position varies from plant to plant. When measuring dietary
406 Dietary Fiber: Properties and Sources
fiber, one must also take into account the method of food
preparation because the soluble fiber content of a fiber source
is influenced by the method of food preparation. For example,
some polyuronides are susceptible to depolymerization at the
high temperatures used to cook the foods. Similarly, some
insoluble fiber is susceptible to depolymerization at higher
temperatures and may be found in the soluble fiber fraction.
In addition to the distinction between soluble and insolu-
ble fibers, viscosity may also be an important fiber characteris-
tic. This is because viscous fibers modestly lower blood
cholesterol and triacylglycerol concentrations and attenuate
the postprandial glucose response. It is thought that the
increased viscosity of the intestinal lumen reduces the rate of
diffusion of glucose, cholesterol, and triacylglycerols to their
respective receptors along the intestinal wall, reducing absorp-
tion. The results of a recent meta-analysis using several differ-
ent clinical studies showed a statistically significant lowering of
plasma triacylglycerols and cholesterol in subjects fed for at
least 14 days with a diet supplemented with oats, psyllium,
pectin, or guar gum, but the effect was modest: a decrease of

1 mg dl1 g1 of soluble fiber was observed. If we consider
that an individual with moderately higher blood cholesterol
has a total serum cholesterol value of
300 mg dl1, the
change brought about from the consumption of viscous, solu-
ble fibers is on the order of 0.3%. Considering that oat bran
contains about 20% fiber by weight and only half of this is
soluble, one can calculate that 10 g of oat bran per day is
needed for each 0.3% change in total cholesterol.
Insoluble dietary fiber is, by definition, the fraction of the
TDF that is not soluble in hot buffer solution. However, the
measured proportion of insoluble fiber can vary depending on
the dietary fiber methodology. Insoluble dietary fiber consists
primarily of hemicellulose, cellulose, and lignin. It is often
incorrectly believed that insoluble fiber is not fermented to
any great extent by the bacteria present in the large bowel.
However, actual measurements of fiber fermentability have
shown that a significant proportion of insoluble dietary fiber
is fermented. This is true even for wheat bran, which has a
relatively lower insoluble fiber fermentability of between 30%
and 40% (
90% of the wheat bran TDF is insoluble dietary
fiber). The fermentability of oat bran is even greater – up to
80% of the insoluble fiber (approximately half of the TDF) is
fermented in the human large intestine. The degree of ferment-
ability is a property unique to each dietary fiber and depends
largely on the nature and the structural arrangement of the
fiber components and also on other physical characteristics
such as particle size. For example, the structure of coarse
wheat bran favors a high water-holding capacity (see later)
and a slow rate of fermentation. This permits the microtrap-
ping of slowly released fermentation gases, increasing fecal
bulk and stimulating defecation.
Water-Holding Capacity
Dietary fiber holds water by adsorption and absorption. Some
water is also retained outside the fiber matrix (free water). The
particle size, chemical composition, and structure of dietary
fiber influence the water-holding capacity. The in vitro water-
holding capacity cannot be used to predict the impact of highly
fermentable fiber on colonic function.
Table 2 Effect of grinding on the properties of wheat bran insoluble
fibera
Mesh Apertureb MPS Glycocholate
sieve (mm) (mm)c WHCd bindinge
As is 800 9.5  0.1 26.5  1.5
20 840 420 8.1  0.1 24.9  1.2
40 420 280 6.6  0.2 23.8  0.8
60 250 180 5.8  0.1 22.1  1.0
80 175 160 5.6  0.1 21.5  0.3
a
Residue after neutral detergent and porcine pancreatic a-amylase treatments.
b
Aperture size of sieve used with a Wiley mill, intermediate model.
c
MPS, geometric mean particle size.
d
WHC, water-holding capacity, grams of water per gram of insoluble fiber, using the
centrifugation method; mean  SEM of three measurements.
e
Grams of glycocholate bound per 0.2 g of insoluble fiber; mean  SEM of four
measurements.
Although lettuce fiber isolates retain much water, the water-
holding capacity expressed in grams of water held by fiber per
gram of edible portion of food has been found to be higher for
wheat bran fiber (4.5 g) and carrot fiber (2.1 g) than for that of
lettuce and various other foods (0.3–1.3 g). The water-holding
capacity of fiber from the bran of ready-to-eat breakfast cereals
is positively correlated (r  0.85 and P < 0.05) with its mean
particle size (MPS): the water-holding capacity of 160 mm MPS
wheat bran fiber is 59% of that of 800 mm MPS wheat bran
fiber (Table 2). A large portion of the water held by wheat bran
fiber appears to be free water.
The water-holding capacity has a significant effect on fecal
output and stool hardness, as discussed earlier. These factors
are important physiological effects of wheat bran and contrib-
ute significantly to its laxative effects.
Bile Salt Binding and Serum Cholesterol Reduction
There is considerable interest in the cholesterol-reducing prop-
erties of dietary fiber. This is because the influence of serum
cholesterol (low-density lipoprotein (LDL) and high-density
lipoprotein (HDL) cholesterol, in particular) on cardiovascular
disease is considerable. Three different mechanisms have been
proposed to account for the serum cholesterol-reducing prop-
erties of dietary fiber. As discussed earlier, a certain soluble
viscous fiber may modestly lower cholesterol by increasing
luminal viscosity and preventing cholesterol (re)absorption.
Acetic and propionic acids, produced by the fermentation of
dietary fibers in the gut, are thought to inhibit liver cholesterol
synthesis at the metabolic level. Dietary fibers also have the
ability to bind cholesterol directly: deconjugated bile salts are
bound to pectic substances by hydrogen bonding, and lignin
appears to bind bile salts through hydrophobic interactions.
The actual in vivo mechanism is unknown, and it is likely that
different fibers operate through different combinations of all
three mechanisms to influence blood cholesterol levels.
In vitro evidence suggests that the physical form of the fiber
may also play an important role in binding bile salts. In com-
monly ingested ready-to-eat cereals, glycocholate binding
(r ¼ 0.90, P < 0.001) and taurocholate binding (r ¼ 0.86,
P < 0.05) have been positively correlated with the MPS of the

neutral detergent fiber (NDF). Reducing the particle size of
wheat bran NDF from 800 to 160 mm reduces its glycocholate
binding by 19% (Table 2). The bile salt-binding capacity of
fiber isolates, although method-dependent, may reflect events
in the terminal small intestine. Purified fiber fractions may be
poor (cellulose) or strong (lignin) binders. Some rice bran
appear to have a high capacity to bind bile salts.
Human cholesterol-lowering studies have linked the con-
sumption of dietary fiber with lower total plasma cholesterol
and increased LDL/HDL ratios in addition to reductions in
serum triglycerides. Studies such as these have suggested that
cardiovascular risk may be reduced by consuming particular
types of foods high in dietary fiber, most notably whole cereal
grains. However, it should be noted that coronary heart disease
(CHD) is a complex disease and is affected by many factors. For
example, the reduction in CHD observed in a recent study was
larger than would be expected from the beneficial effects of
soluble dietary fiber on serum cholesterol levels only. Thus,
reductions in CHD are likely to be the result of many factors,
one of which is TDF intake.
Cation Exchange Capacity
Cation exchange is partly dependent on the presence of uronic
acid in the nonesterified form. The preparation of the fiber
material may decrease the number of nonesterified carboxyl
groups and the apparent cation exchange capacity. Wheat bran
contains little uronic acid, and its cation exchange capacity is
mainly due to diffusion within the dietary fiber network. The
cation exchange capacity of fiber from different sources is
difficult to compare, unless it is expressed per edible portion.
According to the affinity of minerals for carboxylic acid groups,
cabbage and coarse wheat bran show a high cation exchange
capacity compared with pectin.
There has been some concern that the ability of dietary fiber
to bind minerals (as measured in vitro) may lead to mineral
deficiencies in individuals consuming high-fiber diets. In
North American diets, calculations show that mineral intakes
far exceed the potential binding capacity of dietary fiber so that
no need for concern arises. In addition, minerals that are
bound to fibers or that are trapped in the (as yet undigested)
cell wall matrix may not be absorbed in the small intestine but
could be partially released and absorbed in the colon when the
fiber is degraded by bacteria. Absorption of minerals in the
colon has been suggested as a mechanism for accounting for
increased mineral absorption in rats fed with fructooligosac-
charides. This may explain why cation adsorption has not been
consistently related to mineral bioavailability. However, wheat
bran fibers can permanently bind heavy metal ions to decrease
their toxicity.
Viscosity and Gelling Properties
As mentioned earlier, certain soluble fibers, such as oat b-D-
glucans, are viscous when dissolved in water, while others,
such as pectins, show gelling properties. These fibers influence
gastric emptying and absorption rates in the small intestine.
Direct measurement of the viscosity produced by concentra-
tions of fiber likely to be used in diets has shown little effect of
low-methoxy pectin, a slight increase with wheat bran, and a
Dietary Fiber: Properties and Sources 407
significant increase with high-methoxy pectin. Some of the
nonstructural dietary fiber components can also increase
viscosity.
Microbial Degradation (Fermentability)
Most dietary fiber remains undegraded until it reaches the large
intestine, where the extent of fermentation depends on the
source and several other factors, including the physical struc-
ture of the fiber, presence of specific components in the fiber
matrix, nitrogen source, bacterial adaptation, and transit time.
It is generally accepted that soluble fiber is almost completely
fermented in the large intestine, but unfortunately, it is also
generally believed that insoluble fiber is not fermented. Actu-
ally, both insoluble and soluble fibers are extensively fermen-
ted. On average, 70–80% of the TDF from mixed diets (e.g.,
fruits, vegetables, legumes, and most cereals) is degraded by
colonic bacteria. Since insoluble dietary fiber is the major
dietary fiber fraction, representing approximately two-thirds
of the TDF in Western diets, this means that insoluble fiber is
highly fermented. Indeed, as indicated in the preceding text,
70–80% of the insoluble fiber from oat bran and up to 40% of
the insoluble dietary fiber from wheat bran are fermented.
There is some indication that fermentability depends on
particle size, with small particle sizes being more readily fer-
mented, but this has not been consistently reported, and the
effect appears modest. In rats fed with purified diets containing
15% hard red wheat bran, NDF fermentability was largely
unchanged (34–36%) when the MPS varied from 1275 to
394 mm. Similar trends were observed when American Associ-
ation of Cereal Chemists soft and hard wheat bran were fed at
various MPSs. In these experiments, 37.3–37.6% and
30.0–32.8% of the NDF were fermented, respectively.
Interaction between Structure and Physiology
Food Processing
The physicochemical properties of dietary fiber per se are
dependent on the chemical composition and the structural
characteristics of the fiber. However, as noted earlier, knowing
the former does not allow one to predict the latter. When
considering the potential in vivo effect of dietary fiber, one
must also take food processing practices into consideration.
Many different types of food processing practices are currently
employed. Foods can be boiled, canned, frozen, blanched,
parboiled, extruded, adiabatically extruded, and milled. All
processes can have an effect on dietary fiber content, as mea-
sured by current methodologies. For example, heat treatment
can break glycosidic linkages in dietary fiber polysaccharides.
This can solubilize some of the insoluble fiber and reduce the
TDF content if the polymers are broken down into small
molecular mass fragments. Heat treatment can also reduce
the overall length of the polysaccharide chains to lower viscos-
ity and the water-holding capacity of the fiber. The pentosans
in dietary fiber can react with amino acids such as lysine
(Maillard reaction) and form new polymers that do not behave
physiologically like dietary fiber. The boiling of vegetables
increases the apparent TDF content, but this seems to be due
to the loss of nonfiber components to the water.
408 Dietary Fiber: Properties and Sources
It is often difficult to predict beforehand the effect of a food
processing method on the TDF and the distribution between
insoluble and soluble fractions. For example, extrusion cook-
ing of some cereal bran apparently increases its water-holding
capacity in vitro, although this does not translate into a change
in fecal water-holding capacity in vivo. Extrusion cooking of
wheat bran not only causes small decreases in the TDF and
insoluble dietary fiber but also gives rise to small increases in
soluble dietary fiber. Processing of rice (abrasion or parboil-
ing) removes the seed coat and the majority of the dietary fiber.
Some collapsing of the fiber structure also occurs during food
preparation and/or mastication due to the reduction of particle
size, and the fiber structure of sources such as spinach may be
modified by cooking.
Milling is known to significantly affect TDF fermentation in
the human colon. Cereal grains have a tough outer seed coat
that would be completely resistant to digestion if the cereals
were not milled prior to consumption. After milling, the thin-
walled dietary fiber structures from the endosperm are partly
disrupted and, thus, easily broken during fermentation. Milling,
therefore, not only makes the endosperm accessible to the
digestive enzymes in the upper digestive tract but also makes
the dietary fiber more accessible to colonic bacteria. However,
milling can also negatively affect the physiological properties of
the dietary fiber in grains. While coarse grinding and adequate
mastication appear to be sufficient for the digestion of cereals,
fine grinding disrupts much of the cell wall architecture, and
further milling can even interfere with its determination by
gravimetric methods. Table 3 shows the effect of milling on
the dry bulk volume of wheat bran, as measured in a volumetric
cylinder. The volume of the coarse bran was reduced from
5 ml g1 to < 2 ml after grinding in an M5 Wiley mill using a
0.5 mm screen. Fiber from different sources ground under the
same conditions may behave differently during milling, so that
their resulting particle sizes may be different.
As discussed earlier, the particle size of unfermented wheat
bran fiber may be related to colonic function and fecal charac-
teristics in humans and laboratory animals. Finely ground
cellulose or even wheat bran, for example, may have a consti-
pating effect, whereas coarsely ground wheat bran prevents
constipation. The influence of particle size on fecal bulking
has been measured directly in rats. When rats were fed with
diets containing 12% hard wheat bran, the fecal wet density
was 0.796  0.010 (mean  SEM, n ¼ 12) in the coarse bran
group (geometric MPS: 850 mm) but increased to
0.888  0.013 in the fine bran group (geometric MPS:
308 mm). This increased density was not due to a change in
Table 3 Effect of grinding on dry bulk of wheat bran (ml g1):
duplicate measurements
Sieve aperture a
Bran A b
Bran B Bran C Bran D
As is 3.1 3.4 5.1 3.5
2.0 mm (10 mesh) 2.7 2.4 2.8 2.1
1.0 mm (18 mesh) 2.2 2.1 2.4 2.0
0.5 mm (35 mesh) 1.8 1.8 1.9 1.7
a
Using an M5 Wiley mill. Mesh sieve equivalence is indicated in parentheses.
b
Bran A was American Association of Cereal Chemists-certified soft white wheat bran;
bran B–D were soft or hard wheat bran from other sources.
fecal weight but was due to a decrease in fecal volume, parallel-
ing the effect demonstrated for bran in Table 3. In ready-to-eat
breakfast cereals, the fiber MPS has been found to vary between
350 and 2000 mm. Increased particle size normalizes gastroin-
testinal transit time, delays gastric emptying, and may also
affect the extent of insoluble fiber fermentation. Although
dietary fiber has many positive effects on the digestive tract,
excessive milling of dietary fiber to produce small particles in
the range of 7–70 mm may have negative effects. Smaller parti-
cles do not exhibit the transit time normalization associated
with larger bran particles, and the persorption of a small
amount of very fine fiber particles may occur in some regions
of the intestinal wall. Contrary to persorbed starch granules,
persorbed fiber particles are not degraded in the blood and
may impose an increased workload on the kidneys. Data on
persorption are scarce, and much work is needed on this
subject.
Studies with apples, apple puree, and apple juice have
highlighted the link between the physical integrity of the cell
wall and the beneficial effects of unprocessed dietary fiber, in
this case, a lowering of the glycemic index of a food (the rise in
blood glucose that accompanies the ingestion and digestion of
a carbohydrate-containing food). Only the unprocessed apple
had a reduced glycemic index; the apple puree, containing the
physically degraded dietary fiber, was similar to apple juice in
response. This demonstrates the importance of the physical
encapsulation of available macronutrients within the fiber
matrix. This effect has also been observed with rolled oats
where higher glycemic indexes were observed with more finely
cut oats. Studies such as these show that the integrity of the cell
wall is important in bringing about beneficial physiological
effects, even in the presence of viscous, soluble fiber types
(b-glucans in the cell wall; see the section ‘Bile Salt Binding
and Serum Cholesterol Reduction’).
In Vitro Measurements
In vitro properties of fiber isolates have been used to estimate
their potential in vivo effect. However, in vitro data should be
used with caution. The measurements are influenced by the
methods used to follow digestion in vitro and by the methods
used to prepare the fiber. Other confounding influences are
also present: the fiber residue obtained in vitro may have been
altered by heat or chemicals or may still contain residual
digestible materials. For these reasons, in vivo conditions gen-
erally result in a more extensive digestion than that measured
in vitro.
Digestive Events
An important consideration when examining the physiological
effect of dietary fiber is the fact that dietary fiber is concentrated
and denuded only in the terminal small intestine. This means
that the dietary fiber that is partially associated with digestible
materials in the jejunum and duodenum may have different
physical properties from those of an entirely denuded fiber
matrix at the terminal ileum. In the large intestine, most of
the soluble fiber and a large but variable portion of insoluble
dietary fiber are degraded by the colonic microflora. As already
 Dietary Fiber: Properties and Sources 409

discussed, fermentation plays an important role in mediating
many of the physiological effects associated with dietary fiber.
Definition (Based on Function and Structure)
In the early 1970s, it became evident that cellulose represents
only a small part of the TDF content of foods. Prior to this,
there was a tendency to equate dietary fiber with cellulose
because that is what was measured using the old crude fiber
method. While this helped reinforce the use of the term ‘dietary
fiber’ (cellulose has a fibrillar structure), cellulose represents
only a small part of the TDF content of foods. Thus, at that
time, new methodologies were emerging, and it was clear that
there was a need for a dietary fiber definition that was linked to
the actual physicochemical composition of dietary fiber.
It is in this context that the dietary fiber hypothesis appeared,
a few years after Cleave’s hypothesis in 1956, which related
refined foods and Western diseases. The dietary fiber hypothesis
thus appeared as a refocusing of Cleave’s original ideas, revers-
ing the original emphasis to become a diet high in unrefined
edible plant material that is protective against Western diseases
of the bowel. The dietary fiber hypothesis is important in under-
standing the ‘dietary fiber concept,’ since it links botanical struc-
ture to digestive processes and health outcomes – mainly
noninfective colonic diseases such as constipation, diverticular
disease, irritable colon, ulcerative colitis, appendicitis, hemor-
rhoids, polyps, and cancer of the large bowel. It is this link to
these and other health outcomes that has characterized much of
the work in the dietary fiber field over the last 30 years. The first
dietary fiber definition in 1972 sought to describe the plant
structures and components that were responsible for mediating
these health benefits: “that portion of food which is derived
from cellular walls of plants which is digested very poorly by
human beings. . . Fibre is composed largely of cellulose,. . .
hemicelluloses, pentosans, pectin and lignin.”
There are many different definitions of dietary fiber. How-
ever, they all include certain specific and critical aspects. First
and foremost, a definition must include the fact that dietary
fiber corresponds mostly to the structural material in the edible
part of plant foods in the usual human diet. This has been
consistently reflected throughout the decades by the inclusion
of the term ‘plant cell wall’ when referring to dietary fiber
(Table 4). The term ‘dietary fiber’ itself is something of a
misnomer, since dietary fiber is not fibrillar (except for cellu-
lose). To reflect this fact, many other terms have been proposed
over the years, including unavailable carbohydrate and plantix.
None of these terms have succeeded in replacing ‘dietary fiber.’
Another important aspect of any dietary fiber definition is
that it is resistant to human digestive secretions (including
enzymes) and not absorbed in the small intestine. The obser-
vation that dietary fiber increased fecal weight initially gave the
impression that dietary fiber was resistant to both human
digestive secretions and bacterial digestion in the lower gut
(fermentation). While it is true that dietary fiber is not digested
and absorbed in the small intestine, a large proportion of
dietary fiber is fermented in the colon. The extent of this
fermentation is a function of several different factors (see the
section ‘Microbial Degradation (Fermentability)’), meaning
that the extent of fermentation cannot be used as a criterion to
define dietary fiber.
Using ‘resistance to endogenous digestive secretions’ alone
as a criterion may be confusing. For example, some dietary
fiber may be partly degraded by acids in the stomach, while

Table 4 Use of the term ‘plant cell wall’ in defining dietary fiber over time

Year Text Reference

1885 The ‘cell membranes’ of Rubner
1929 The ‘skeletal framework of the plant’
1981 The ‘remnants of plant cells’
1985 ‘Derived from plant cell walls and not
digested by human alimentary
enzymes’
1990 The ‘structural plant cell wall
composed of polysaccharides and
lignin’
1995 ‘The “cell wall material of plant foods”
responsible for the physical form
and texture of unprocessed plant
foods’
1995 The ‘plant cell walls’ are the common
characteristic of plant foods that
constitute a high-fiber diet
associated with beneficial effects
1999 ‘The cell walls of edible plant tissues in
the traditional human diet’
Trowell, H. C. (1975). In: Burkitt, D. P. and Trowell, H. C. (eds.) Refined carbohydrate foods and
disease, p. 43. London: Academic Press
McCance, R. A. and Lawrence, R. D. (1929). The carbohydrate content of foods. Medical Research
Council special report series No. 135, pp. 1–173. London: HM Stationary Office
AOAC Fiber Consensus (1981). Association of Official Analytical Chemists 95th Annual Meeting,
October 1981, Washington, DC
Trowell, H., Burkitt, D. P. and Heaton, K. (eds.) (1985). Dietary fibre, fibre-depleted foods and
disease. London: Academic Press
Eastwood, M. (1990). Function of dietary fibre in the large intestine. In: Southgate, D. A. T., Waldron,
K., Johnson, I. T. and Fenwick, G. R. (eds.) Dietary fibre: chemical and biological aspects, pp.
211–219. Cambridge: Royal Society of Chemistry
Heaton, K. W. (1995). The dietary fibre concept – time for a re-evaluation? In: Sørensen, A., Bach
Knudsen, K. E., Englyst, H. N., Gudmand-Høyer, E. and Nyman, M. (eds.) Metabolic and
physiological aspects of dietary fibre in foods: recent progress in the analysis of dietary fibre, COST
92, p. 15. Luxembourg: Commission of the European Commission
Cummings, J. H., Hudson, G. J., Quigley, M. E. and Englyst, H. N. (1995). The classification and
measurement of dietary carbohydrates. In: Sørensen, A., Bach Knudsen, K. E., Englyst, H. N.,
Gudmand-Høyer, E. and Nyman, M. (eds.) Metabolic and physiological aspects of dietary fibre in
food. Recent progress in the analysis of dietary fibre in food, COST 92, pp. 17–36. Luxembourg:
European Commission
Mongeau, R., Scott, F. W. and Brassard, R. (1999). Definition and analysis of dietary fiber. In: Cho, S.
S., Prosky, L. and Dreher, M. (eds.) Complex carbohydrates in foods, pp. 305–316. New York &
Basel: Marcel Dekker
410 Dietary Fiber: Properties and Sources
some small intestine-digestible material can reach the colon
where bacterial fermentation occurs. In addition, a definition
based on the resistance to digestibility by endogenous digestive
secretions could be problematic if the material is not specified
as coming from the edible part of plant foods. Indeed, such a
definition could be erroneously interpreted as meaning that
dietary fiber is any material resistant to in vivo digestion or even
as any material measured as dietary fiber by a specific dietary
fiber method. Among the materials to be excluded from the
dietary fiber definition are ‘ruminant fiber’ and the products
formed during processing and/or cooking. There have been
requests to include some other plant components and manu-
factured polymers, and this is a subject of much debate. Among
the criteria that could be used to accept or reject a material as
dietary fiber, only ‘edible plant cell wall’ is specific enough to
allow an absolute definition. In addition, it links current die-
tary fiber definitions to the original ‘dietary fiber concept,’
which recognized that a diet high in minimally processed
edible plant foods provided many health benefits. Confusion
often arises because of the overlapping characteristics and
physiological effects of the different ‘candidate’ materials with
those of a dietary fiber ‘reference.’ Examples of ‘candidates’ are
polydextrose (a randomly bonded synthetic glucose polymer),
inulin and fructooligosaccharides (naturally occurring
glucose–fructose oligomers and polymers), and resistant starch
(formed by food processing methods and largely undigested in
the small intestine). This overlapping means that it becomes
virtually impossible to distinguish between fiber and nonfiber
material based on resistance to digestion alone. Debates such
as this started shortly after dietary fiber became popular in the
1970s. They are still ongoing, as demonstrated by the publica-
tion of a recent international survey of 147 professionals.
While a complete discussion of the merits of either side is
beyond the scope of this article, it is important to remember
the origins of the dietary fiber definition and the dietary fiber
hypothesis that firmly linked fiber to physiological outcome.
The definition of dietary fiber was first centered on the
‘original dietary fiber concept’ that linked the consumption of
‘plant cell walls’ in unrefined, traditional foods to positive
physiological outcomes. Table 5 shows two definitions that
were proposed after the turn of the millennium. These two
definitions have taken the original ideas of the dietary fiber
hypothesis into account and included a reference to some
specific physiological effects in humans.
While the newer definitions of dietary fiber specifically
mention the fact that dietary fiber has physiological benefits,
Table 5 Recent proposed definitions of dietary fiber
Year Text
2001 ‘. . .the edible parts of plants or analogous carbohydrates that are
resistant to digestion and absorption in the human small
intestine. . .promotes beneficial physiological effects including
laxation, and/or blood cholesterol attenuation, and/or glucose
attenuation’
2002 ‘Dietary Fiber consists of nondigestible carbohydrates and lignin that
are intrinsic and intact in plants. Functional Fiber consists of
isolated, nondigestible carbohydrates that have beneficial
physiological effects in humans’
the list is rather specific and deals primarily with effects
observed with viscous fibers (lowering postprandial serum
glucose and lowering serum cholesterol) or effects brought
about from partial fermentation and water-holding capacity
(laxation). These effects are well supported by clinical experi-
ments with laboratory animals and with humans. However,
the original dietary fiber concept related dietary fiber intake to
the diseases of Western man, including heart disease, cancer,
and obesity. As mentioned earlier, the link between viscous
dietary fiber and heart disease is supported in the literature
through its effect on serum cholesterol and triglycerides, but
the link with other Western diseases has been much harder to
demonstrate. It has been well established that eating patterns
that favor foods high in dietary fiber and low in red meat
significantly reduce the risk of cancer, heart disease, and obe-
sity. However, it has been relatively difficult to isolate this
effect to dietary fiber consumption per se. Part of the problem
comes from the almost exclusive reliance on epidemiological
studies to demonstrate any association. There are several rea-
sons why epidemiological studies are generally favored. First,
these studies provide data on free-living humans, rather than
inbred laboratory animals. Second, these studies follow the
long-term effect of dietary fiber consumption on health out-
comes. Third, the sample size can vary from a few hundred
(case control studies) to many thousands of individuals (pro-
spective cohort studies). This gives the studies good statistical
power and lends weight to the conclusions. However, epide-
miological studies also suffer from a number of shortcomings
including problems with assessing food intake for the past 2–3
years (case control studies) or with monitoring food intake 2–3
times over a period of 5 years and using this to predict disease
outcome 15 years later (prospective cohort studies), problems
that arise from the use of incomplete dietary tables to calculate
dietary fiber intake, and problems arising from the use of
inappropriate methods for correcting energy intake.
The most striking conclusion from epidemiological studies
is a failure to provide support for an inverse correlation
between dietary fiber intake and colon cancer. This concept
has long been entrenched in the literature largely because of
studies with laboratory animals that have provided much data
in favor of a protective effect of dietary fiber. The animal
studies are supported by plausible physiological and biochem-
ical mechanisms that can account for the observed results.
However, the epidemiological data are inconsistent in its sup-
port for an inverse relationship between colon cancer and
dietary fiber intake, and clinical trials that examined the effect
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 Dietary Fiber: Properties and Sources 411

of diet on the reappearance of colon polyps (colon cell growths content expressed on an ‘as is’ basis. Since some methodolog-
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Corn kernel (canned), potatoes (boiled), EMP serving of bran flakes will provide
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(raw), onion (cooked), green peppers (raw) will provide about 1.8 g of TDF, and 1 serving of corn will
Cauliflower (cooked), tomatoes M provide about 2 g of TDF.
Apple, bananas, orange, pears, strawberries, EMP
cherries, grapefruit, plums
Apple pie, blueberry pie, pasta (cooked) M
2.3–3.0 Rye bread M See also: Cellulose; Dietary Fiber: Bran; Dietary Fiber: Determination.
Mushroom (E ¼ 2.0) MP
Beets (canned), rutabaga (raw), carrots EMP
(raw), broccoli (raw)
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