editor: Norton S. Rosensweig, M.D.

comments in gastroenterology

Plant fiber. Carbohydrate and lipid

metabolism1 2

James W Anderson and Wen-Ju Lin Chen

ABSTRACT Plant fibers are the portions of plant foods that are not digested in the human
small intestine. During this century, remarkable advances have been made in defining the
characteristics and importance of most nutrients such as carbohydrate, proteins, fats, vitamins, and

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minerals. Plant fibers have largely been neglected because they were considered to have no nutritive
values. In the last decade, however, considerable attention has been focused on the various plant
fibers because of their influence on gastrointestinal physiology. Evidence is emerging that plant
fibers have profound influences on human nutrition because they alter the absorption and
metabolism of many nutrients. We will review the evidence that plant fibers greatly influence the
absorption and subsequent metabolism of carbohydrates and fats. Am. J. Clin. Nut,. 32: 346-.
363, 1979.

Classifications can be divided on the basis ofwater solubility

into 1) insoluble fibers and 2) soluble fibers.
In their natural state plant fibers are amal- The insoluble fibers include cellulose, lignin
gamated into many different anatomical and and many hemicelluloses and are measured
biological functional units. Because of the by the neutral detergent residue method of
hundreds of different polysaccharides com- Goering and Van Soest (3). The soluble fibers
posed of over a hundred different sugars, a include pectins, some hemicelluloses, gums,
definitive chemical or structural classification mucilages, algal polysaccharides, and the
of plant fibers cannot be made at present. At storage polysaccharides (1) that are not in-
our primitive stage of knowledge, three pro- cluded in the neutral detergent residue anal-
visional classifications are useful. From a bi- ysis of plant fibers (2) but are estimated by
ological viewpoint, plant fibers can be di- the technique of Southgate (4).
vided into 1) structural fibers, 2) gums and Ultimately, one classification of plant fi-
mucilages, and 3) storage polysaccharides (1,
bers will relate to their chemical structure.
2). The structural fibers make up the plant The chemical structure of most plant fibers
cell walls and include cellulose, lignin, many has not been determined but the composition
different hemicelluloses and pectins. The
of several dozen has been delineated. The
gums and mucilages have special functions
chemical structure, anatomical relationships
such as the repair of injured areas. Plants and proposed physiological role in humans
store energy as complex carbohydrates which of selected fibers will be reviewed.
are digestible (e.g., starch and dextrins) and
as indigestible fibers termed storage polysac-
charides. The storage polysacchandes from From the Medical Service, Veterans Administration
Hospital and University of Kentucky College of Medi-
certain beans (e.g., guar gum) are of particu-
cine, Lexington, Kentucky.
lar interest because of their profound influ-
2Address reprint requests to: James W. Anderson,
ence on carbohydrate and lipid metabolism. M.D., Veterans Administration Hospital, Lexington,
From an analytical standpoint plant fibers Kentucky 40507.

346 The American Journal ofClinical Nutrition 32: FEBRUARY 1979, pp. 346-363. Printed in U.S.A.
 PLANT FIBER: CARBOHYDRATE AND LIPID METABOLISM 347

Cellulose (1). than 5% of ingested pectin is recovered in the

stool (1). The pectins can form gels and have
This is the best known and most widely
distributed component of the plant cell wall. considerable water holding capacity. They
can also bind cations and organic material
It is an unbranched glucose polymer with
approximately 3000 glucose units. Cellulose such as bile acids.
is not soluble in water but its main physiolog- Gums, mucilages, and storage polysaccharides
ical action in the human gut is to bind water. (1).
Cellulose can take up about 0.4 g of water
These substances are not components of
per gram of fiber.
the cell wall but have biochemical and phys-
Lignin (1). iological properties that resemble the pectins
and certain hemicelluloses. Plant gums are
All of the major plant fibers are polysac- the sticky material that is extruded at the site
charides except the lignins that are phenyl- of injury to plants. These gums are a complex
propane polymers with molecular weights be- group of highly branched polysaccharides
tween 1000 and 4500. Lignin is synthesized containing glucuronic and galacturonic acid
from derivatives of these alcohols: coumaryl, as well as xylose, arabinose, and mannose.

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coniferyl, and sinapyl alcohol. Lignin is vir- Gum arabic or gum acacia, one of the better
tually insoluble in strong acid or alkali and is known gums, is a galacturonic acid polymer
not digested or absorbed in the colon. Lignin with side chains containing arabinose and
can bind bile salts and other organic materials rhamose. It has a molecular weight of ap-
(5) and may delay or impair the small intes- proximately 250,000 and is water soluble. The
tinal absorption of associated nutrients (6). various gums are used in the food industry as
emulsifiers, thickeners, and stabilizers. Some
Hemicelluloses (1). gums are used in pharmaceutical prepara-
tions and as bulk forming laxatives while
This family of fibers containing over 250
others are sold commercially as adhesives.
different polysaccharide polymers is derived
Mucilages and storage polysaccharides are
from pentose and hexose sugars. These struc-
usually dispersed throughout the endosperm
tural components of the cell wall are inti-
and are mixed with the digestible polysaccha-
mately associated with cellulose and thus
have been termed hemicelluloses. In general rides (e.g., starch). Guar gum highlights the
current problems with classification and no-
the hemicelluloses have xylose backbones
menclature. Guar gum is classified as a mu-
with various substitutions and degrees of
cilage (1) but is a major storage polysaccha-
branching. Most molecules have 150 to 200
ride. This galactomannan with a molecular
sugars and contain two to four different
weight of approximately 220,000 is derived
sugars. The most common sugars are xylose,
from the Indian cluster bean Cyamopsis tetra-
arabinose, mannose, galactose, glucose, rha-
gonolobus. After separation from the husk
mose, and uronic acid. Various hemicellu-
and germ, the endosperm is ground to a fine
loses have important physiological action in
particle size and marketed as guar gum. Be-
the gut where they have water-holding capac-
cause of its water absorption and binding
ity and can bind cations.
properties guar gum is widely used in the
Pectins (1). mining and paper industry, in cosmetics and
pharmaceuticals, and in the manufacture of
The basic structure is a polymer of galac- explosives and tobacco products. In the food
turonic acid with variable degrees of esterifi- industry small amounts of guar gum are
cation ofthe uronic acid with methyl or acetyl added to sauces, dressings, ice cream, sher-
groups. In most plants, the basic galacturonic bert, frozen foods, pie fillings, and dog foods
acid polymer has side chains composed of because of its water binding and stabilizing
galactose, arabinose, xylose, rhamose, and properties (8). This neutral polysaccharide
fucose. Their molecular weights usually range has little cation binding property but does
from 60,000 to 90,000. Pectins are virtually form gels in the small intestine and binds bile
completely digested in the colon (7) and less acids and other organic materials.
348 ANDERSON AND CHEN

Other plant cell wall substances (1). These comparisons lead to the erroneous con-
clusion that adding variable amounts of bran
Many additional compounds are inti-
or other fiber-rich materials to the diet would
mately linked to the plant cell wall and are
decrease transit time and increase fecal
largely indigestible. This list includes phytic
weights. Since different fibers have different
acid, silica, lipids, plant sterols, proteins, sa-
properties this simplistic concept has not been
ponin, tannins, and many additional sub-
substantiated (I 1, 12). Plant fibers may alter
stances. The phytates bind cations and may
gastric emptying time, small intestinal transit
lead to fecal losses of iron and zinc (1). Indi-
time, and colonic emptying times in different
gestible plant lipids such as cutins and waxes
fashions. Since some fibers appear to be com-
may account for a substantial amount of the
pletely digested in the colon, they may not
fecal fat of individuals on high fiber diets (I).
increase fecal bulk. The effects of different
fibers or different fiber-rich foods on these
Physiological effects of fiber in gut
physiological parameters have not been de-
The physiological effects in the gut of fiber- lineated. In general, adequate quantities of
rich foods, semipurified fibers, and highly cellulose-rich foods (e.g., wheat bran) de-
purified fibers have been extensively studied. crease intestinal transit time and increase fe-

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Interpretation of these results are difficult for cal bulk (2). On the other hand, soluble fibers
many reasons. Fiber-rich foods contain a (e.g., pectin or guar gum) may delay gastric
mixture of different fibers and it is difficult emptying, increase small intestinal transit
to assign an effect to a specific fiber compo- time because of their gel formation and have
nent. In the raw plant, different fiber mole- only a small effect on fecal bulk (13-15).
cules are intermeshed to meet structural and
biological requirements of the plant. The Water absorption
anatomy of the plant is destroyed by cutting, Most polysaccharides swell in the presence
cooking, mastication, and digestion and, thus, of water to form gels. Pectins, gums, muci-
the fiber-nutrient relationships are altered. lages, and storage polysaccharides have high
Certain functional properties of these fibers affinities for water and form gels in the small
that are retained are modified by environ- intestine. Some of the structural hemicellu-
mental factors in the gut. The physiologic loses also have the capacity to form gels while
effects of fibers in the gut may be quite other hemicelluloses and cellulose bind water
different from those observed in vitro. Fac- to a limited extent (9). The water holding
tors such as osmolality, pH, the presence of capacity of plant polysaccharides is deter-
other fibers or nutrients, the digestive process, mined by their chemical and structural prop-
water retention and the presence of bacteria erties and also by the pH and osmolality of
will profoundly influence the physiological the surrounding fluids (9). There is almost a
effects of fibers (9). Furthermore, different 15-fold range in the in vitro water-holding
fibers may have synergistic or antagonistic capacity of various plants. When expressed
effects when fed together. Nevertheless, cer- as water retained per gram of fiber, a range
tam general properties of plant fibers can be of 1.5 g water per gram of fiber for maize to
listed. 23.7 g water per gram of fiber for lettuce was
observed (9, 16). Because of the wide varia-
Altered transit time
tion in the fiber content of plants the water
High fiber diets are accompanied by rapid holding capacity per gram of raw vegetables
passage of food residues through the gut and ranged from 0.37 g water per gram of turnip
by increased fecal bulk. These effects are to 4.5 g water per gram of wheat bran. Some
illustrated when African villagers eating high purified plant fibers such as carragenan are
fiber diets are compared with British males so hydrophilic that they can form gels con-
on low fiber diets. Gut transit time in the taming 99% water (17). These water absorp-
Africans averaged 36 hr compared to 77 hr tion properties of most fibers contribute to
for the English. Daily fecal output averaged alterations in intestinal transit time and also
470 g for the Africans while values for the to increases in fecal water associated with
English men were 108 g (see Reference 10). certain foods or fibers.
 PLANT FIBER: CARBOHYDRATE AND LIPID METABOLISM 349

Cation binding (9, 16) Digestion offibers

The cation exchange properties of fibers Plant fibers excepting lignin appear to be
are related to the acidic sugars, especially the partially digested or fermented in the colon
uronic acid groups. The binding of cations by (1 24-30). Williams
, and Olmstead (24) dem-
these polysaccharides may alter the physio- onstrated that only a small percentage of the
logical properties of these fibers for example fiber of plants commonly eaten by man could
by effecting gel formation. The nutritional be recovered in the feces. Recent studies
consequences of cation binding by fibers is (25-27, 30) using modern techniques for mea-
uncertain at present. Short term balance stud- suring plant fiber have confirmed these ob-
ies suggest that the initial weeks of a high- servations. For example, when 16 normal
fiber diet may enhance fecal excretion of subjects were fed 8.5 g cellulose per day an
calcium, iron, magnesium, and zinc (2, 18, average of only 43% was recovered in the
19). Further studies are required to evaluate feces (30). The digestibility of fibers depends
the long term effects of high fiber diets on on their chemical structure, the bacterial flora
mineral balance. of the colon, the length of residence in the
colon, and other factors. Digestibility may be

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A bsorption of organic substances (20) greatly influenced by the physical character-
istics of the bolus that enters the colon; a
Bile salts are absorbed by lignin and other complex gel filtration system may exclude
plant fibers (5, 9, 2 1 ). Conjugated bile salts in bacteria and diminish digestion (9). In gen-
the small intestine are only weakly absorbed eral pectin and structurally similar soluble
but after chemical transformation by bacteria fibers are almost completely digested (1, 7),
in the colon, deconjugated bile salts are hemicelluloses are digested to a variable de-
strongly bound and excreted in the feces (9). gree (56 to 87%, Reference 1) and various
Fecal loss of bile salts may play a major role estimates suggest that about 40% of cellulose
in the hypocholesterolemic effects of certain is digested ( I ). A cellulytic anaerobic bacte-
plant fibers. Other organic material is also rium has been isolated from the human in-
bound by plant fibers (2, 9, 10). The binding testine (29). The metabolic products of bac-
of carcinogenic substances or alterations in terial action on fibers are largely methane,
bacterial metabolism (10) associated with C02, water, and short chain or volatile fatty
high fiber diets may protect against the de- acids (VFA) such as acetate, butyrate, and
velopment of colonic cancer in man (2, 22, proprionate (28). Van Soest and Robertson
23). (28) estimate that half of the polysaccharide
carbons are converted to VFA which can
Gelformation (9) either be absorbed or excreted in the feces.
While most authorities (1, 25, 28) agree
With hydration, a variety of fibers form that the polysaccharide fibers are partially
gels in the small intestine and colon. A gel digested in the colon, there is disagreement
filtration system similar to that used in col- about the extent to which nutrients might be
umn chromatography may be established. absorbed. Some investigators (10, 25) suggest
This system could exclude molecules or bac- that negligible amounts of VFA are absorbed
teria on the basis of size or ionic charge. A from the colon and that the products of fiber
gel filtration system would tend to delay the digestion are insignificant sources of calories.
digestion and absorption of nutrients in the Other workers (24, 28, 3 1) maintain that over
interstices ofthe system. In the small intestine 50% of the VFA produced are absorbed from
this gel system may delay the absorption of the colon and that plant fibers may provide
carbohydrate and other nutrients. In the co- small quantities of energy in the human.
lon, a gel system may greatly alter the avail- From available data on the digestibility of
ability of fiber, bile salts and nutrients for plant fibers in the colon (2, 24, 28, 3 1) and
metabolism by bacteria. The physiological the likelihood that some of these VFA may
and nutritional consequence of these gel fil- be absorbed (24, 28, 32) we feel that plant
tration systems has not been examined. fibers may be a source of energy for man.
350 ANDERSON AND CHEN

The energy available from fibers would be serum insulin changes. All feedings contained
offset by fiber-induced malabsorption of nu- 60 g of carbohydrate. Eating whole apples
trients (2). Thus, further studies are required was accompanied by significantly greater sa-
to delineate the net effects of various fiber- tiety than with either the apple puree or apple
rich foods on energy balance. juice (Fig. 1). Similar effects of plant fiber on
satiety have been reported by others (2 15
Altered digestion and absorption ofnutrients 36). After ingestion ofwhole apples there was
When the intake of plant fiber is increased a smaller rise in plasma glucose at 30 mm but
the digestibility of nutrients is decreased (25). glucose values returned to baseline values at
Fecal excretion of both fat and nitrogen is 60 mm and remained there for the next 2 hr.
increased when the diet is supplemented with In contrast, apple juice induced slightly
fiber (25-27, 33). With only a small increase greater early hyperglycemia but was associ-
in plant fiber intake a significant increase in ated with plasma glucose values that were
energy loss in the stool can be observed (26). distinctly lower than fasting values between
When the plant fiber intake of normal vol- 60 and 180 mm. The serum insulin rise was
unteers was increased from 14 to 27 g/day by 2-fold greater after apple juice than after
feeding bran biscuits (27), fecal values in- whole apples. Undoubtedly the significantly

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creased: for energy from 108 to 166 kcal/day greater insulin rise after apple juice induced
(P < 0.025), for total nitrogen from 1.2 to 1.7 the significant reduction in plasma glucose
g/day (P < 0.005), and for fat from 2.8 to 3.8 values during the 2nd and 3rd hr of the test.
g/day (not significant). The influence of high The effects of apple puree were intermediate
fiber diets (e.g., 65 g/day, Reference 34) con- between that ofwhole apples and apple juice.
taming a variety of plant fibers on nutrient This is of considerable interest because it
and energy excretion has not been evaluated.
This energy loss as a result of nutrient mal-
absorption on high fiber diets may be fairly
well compensated by the energy provided by +6 SATIETY
VFA. Conventional balance studies may not
provide adequate information to resolve these
questions. The critical question relates to the

amount of available nutrients which enters

the colon and may require studies of patients
-6
with ileostomies (7). Ifavailable carbohydrate
is not absorbed in the small intestine but is
metabolized and absorbed from the colon,
this results in less available energy. It is ob-
vious that further definition of the impact of E

fiber-rich foods on nutrient absorption in the

small intestine, metabolism in the colon and 40 N: 10

excretion in the feces will require innovative APPLE.

investigative techniques. PUREE 0

40 . JUICEA

Carbohydrate metabolism

Normal subjects
When carbohydrates and plant fibers are
60 (20 (80
ingested together less hyperglycemia ensues
MINUTES AFTER FEEDING
than when the same amount and type of
carbohydrates are ingested without plant fi- FIG. 1. Effects of different apple preparations on
satiety, plasma glucose and serum insulin. Ten normal
ber. This was elegantly demonstrated by the
subjects ingested 60 g ofcarbohydrate from whole apples,
studies of Haber et al. (35). They compared apple puree, or applejuice (35). The hunger-satiety score
the influence ofwhole apples, apple puree, or ranged from painfully hungry (-10) to full to nausea
apple juice on satiety, plasma glucose, and (+10). Redrawn from the data of Haber et al. (35).
 PLANT FIBER: CARBOHYDRATE AND LIPID METABOLISM 351

suggests that physical disruption of the plant (41). When 24 g of wheat bran was added to
fibers alters their physiological properties. the daily diet of 37 nondiabetic patients with
When large quantities of soluble plant fi- diverticular disease, a significant improve-
bers were incorporated in test meals, even ment in the oral glucose tolerance test was
greater effects on blood glucose and insulin observed after 6 months (38). Improvement
values were observed. Jenkins et al. (37) fed in glucose tolerance (38-40) has been associ-
594-kcal breakfasts containing carbohydrate ated with diets rich in insoluble plant fibers
(106 g), protein, and fat with or without plant such as wheat bran whereas diets rich in
fiber. The fiber-rich meals contained 16 g of soluble plant fibers such as pectins have not
guar flour and 10 g of pectin. The meals had favorable long-term influences on glu-
containing these soluble fibers were followed cose tolerance (41). Differences in the types
by significantly lower blood glucose and se- of plant fiber or in the amount and type of
rum insulin levels than observed with the carbohydrate used (42, 43) may explain these
fiber-free meals (Fig. 2A). After this large divergent findings.
glucose load the blood glucose concentration
Patients with diabetes mellitus
increased only 19 mg/dl when plant fibers
were ingested concomitantly. These two stud- Less postprandial hyperglycemia occurs

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ies (35, 37) clearly demonstrate that when when patients with diabetes ingest fiber-rich
carbohydrate is ingested with plant fibers the meals than observed with fiber-free meals.
resultant rise in blood glucose and insulin Jenkins et al. (44) fed test meals containing
concentrations are significantly lower than pectin and guar gum to patients with chemi-
when the same carbohydrate loads are taken cal diabetes using a similar protocol to that
without fibers. described above. Blood glucose and insulin
The long-term effects of increasing fiber levels were significantly lower after the fiber-
intake on glucose tolerance of normal mdi- rich meals than after the fiber-free breakfasts
viduals are not well delineated. Some studies (Fig. 2B). Likewise when these fiber-rich
(38-40) suggest that high fiber diets may meals were fed to insulin requiring patients
improve glucose tolerance whereas another with diabetes, postprandial hyperglycemia
study failed to note significant alterations was distinctly less than values after fiber-free

A. CONTROLS N:? B. DIABETICS N:8

150 300
GLUCOSE %GU0SE

100 #{149} 200

E FIBER-FREE
#{149} MEAL
50 o FIBER-RICH MEAL 100

* p-O.05

V 60 120 C
0 60 20 80

00 50
NSULIN I NSULIN
E

a.
25

. . . I

0 60 120 0 60 20 80

MINUTES AFTER FEEDING

FIG. 2. Effects of fiber-free and fiber-rich meals on blood glucose and serum insulin reposnses. A, seven normal
. subjects were given test meals without additional plant fiber or meals containing 10 g of pectin and 16 g ofguar flour
(37). B, eight noninsulin-requiring patients with diabetes mellitus were fed similar fiber-free and fiber-rich meals
(44). Redrawn from the data of Jenkins et al. (37, 44).
352 ANDERSON AND CHEN

meals (Fig. 3). To extend these observations, GLYCOSURIA

Jenkins et al. (45) supplemented the diet of
seven overtly diabetic patients with 25 g of
guar gum for 5 to 7 days. This addition of
soluble plant fiber to the diet resulted in a
50% reduction in glycosuria in these patients
(Fig. 4). Miranda and Horwitz (46) compared .

the influence of low (3 g of crude fiber per

day) and high (20 g crude fiber per day) fiber
diets on the plasm glucose values of eight w
C,,
patients with diabetes maintained on constant 0
C-)
insulin doses throughout the study. Postpran-
-J

dial glucose values were significantly lower

w
on the high fiber diets (Fig. 5). During the 10
days on the high fiber diet mean plasma
glucose values (120 10 mg/dl, mean
SEM) were significantly lower than values

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during the 10 days on the low fiber diet (169
12, P < 0.001). All of these observations
(44-46) strongly suggest that the addition of
CONTROL GUAR GUM
plant fibers to the diets of patients with dia-
25G/d
betes may reduce postprandial hypergly-
cemia and glycosuria and may allow insulin FIG. 4. Influence of guar gum supplements on gly-
cosuria of patients with diabetes. Urine glucose excretion
doses to be reduced.
was measured on control diets and on diets containing
The therapeutic utility of high fiber diets 25 g of guar gum per day. Redrawn from the data of
for treating insulin-requiring patients with Jenkins et al. (45).
diabetes is supported by our own observa-
tions (34, 47-49). We have fed high-carbo-
hydrate (70% of kcalories), low-fat (10% of
kcalories) diets to 16 insulin-treated lean men
POSTPRANDIAL GLYCEMIA with diabetes. These weight-maintaining
400
diets provided approximately 60 to 80 g/day
CONTROL of plant fiber from a variety of natural
MEAL sources. The response of one representative
patient is presented in Figure 6. On a control
300
diet containing 43% of calories as carbohy-
drate and 26 g/day of plant fiber these men
E
required an average of 27 3 (mean SEM)
C,)
w
0
units of insulin daily to maintain average
C., 200 MEAL WITH
fasting plasma glucose values of 168 10
-I FIBER
mg/dl and 3 hr postprandial plasma glucose
0
0
0
values of 189 16 mg/dl. After an average
-J
of 18 days on the high carbohydrate, high
00 fiber diets values averaged: insulin dose, 12
4 units (P < 0.001); fasting plasma glucose
N: 3 values 140 7 mg/dl (P < 0.02); and 3 hr
____ postpranclial glucose values, 157 10 mg/dl
0
(P < 0.01). Insulin therapy was discontinued
HOURS
in all eight men who were receiving less than
20 units/day as demonstrated in Figure 7
FIG.
3. Effects of fiber-rich meals on blood glucose
response of three insulin-treated patients with diabetes.
(34). The dramatic reduction in insulin doses
Control meals and fiber-rich meals were fed as described in these men is attributable in part to the
in Figure 2. Redrawn from the data ofJenkins et al. (44). generous carbohydrate intake and to the fat
 PLANT FIBER: CARBOHYDRATE AND LIPID METABOLISM 353

restriction. However, previous investigators 300-

FASTING
(50, 5 1) have noted no changes in insulin or PLASMA 57 WM

sulfonylurea requirements when patients

were fed high carbohydrate (60 or 62% of E
kcalories), low-fat diets. It seems very likely 45
that the abundance of a wide variety of plant ADA HIGH CHO, FIBER DIET
0
fibers in our diets contributed to the improve-
ment in diabetic control. Furthermore, 10
>. 20-
individuals who have had significant reduc- INSULIN IN DOSE
0
tions in insulin requirements on these high-

:
CI, 0
carbohydrate, high-fiber diets in the hospital I-
z
have been maintained on the same doses of 0
D
insulin as outpatients for an average of 15
months. The maintenance diets provide only WEIGHT
U) $65 -

55 to 60% of calories as carbohydrate but 0

z
provide 50 to 70 g of plant fiber per day (49). 0 155 - 60 160
a.
The reports of Douglass (52, 53) also sug-

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gest that high plant fiber, high-carbohydrate
=4 0 5 0 5 20 25
diets are of long term benefit in reducing
insulin requirements of patients with diabe- DAYS 1.77

tes. Four patients who consumed large quan- FIG. 6. Fasting plasma glucose values and insulin
tities of uncooked vegetables, seeds, nuts, and dose of diabetic man fed control (ADA) and high car-
fruits had sizeable reductions in insulin doses bohydrate (CHO), high-fiber diet as described in text.
while eating these raw diets. These diets,
like ours (47-49), are high in carbohydrate
and low in fat and contain plentiful quantities
of plant fibers in their natural or undisrupted
form. Since cutting, mincing, or cooking
Miranda 1978
220 foods alter the physical and physiological
A.
properties of plant fibers (35, 54), these im-
200
pressive reductions in insulin doses may have
0
been related in part to the uncooked plant
fiber content of these diets.
I 80
w
U)
0
Proposed mechanisms
U 160
Plant fibers alter intestinal physiology in
0 many different ways as outlined above. Car-
I40
4 bohydrate absorption and/or metabolism
CI, may be altered by one or more of the follow-
4 120
-J ing mechanisms.
a.
1) A delay in gastric emptying may slow
I00
the absorption of carbohydrate. While bran
_______________________________ appears to accelerate gastric emptying (13),
... I I I 1

0730 1130 $700 2100 soluble fibers such as guar gum may slow the
gastric emptying time ( I 3, 15).
FIG. 5. Plasma glucose values of patients with dia- 2) The gel forming properties of most sol-
betes on low-fiber and high-fiber diets. Eight insulin-
treated patients were fed low-fiber diets for 10 days and uble fibers such as pectin and guar gum may
high-fiber diets for 10 days in random order (46). Fasting result in slower absorption of carbohydrate
plasma glucose values were similar but postprandial from these intraluminal gels. This would ac-
glucose values were significantly lower (P < 0.001) on count for the lower insulin levels and the
the high fiber diet (solid line) than on the low-fiber diet
(broken line). (Reproduced with permission from P. M. blood glucose patterns seen in Figures 1 to 3.
Miranda and D. L. Horwitz, Ann. Internal. Med. 88: 3) Intestinal transit time may influence the
482, 1978.) rate ofabsorption ofcarbohydrate. With bran
354 ANDERSON AND CHEN

0
0
N:8
In

LaJ

0
0

z
-J

Downloaded from www.ajcn.org by guest on June 5, 2011

C,,

-5 0 5 10 5 20
DAYS I

FIG. 7. Changes in insulin doses on high-carbohydrate, high-fiber diets. Eight lean men required an average of
16 units of insulin per day on the control (ADA) diet to maintain fasting plasma glucose values of 153 15 mg/dl
(mean SEM). On the high-carbohydrate, high-fiber diet, insulin therapy was stopped in all men after an average
of 14 days (arrow) and fasting plasma glucose values were 130 6 mg/dl on no insulin therapy.

and other sources of insoluble plant fibers or absorbed in the small intestine. This avail-
intestinal transit time is usually reduced al- able carbohydrate may escape into the colon
lowing less time for carbohydrate to be ab- and be digested by bacteria or even be lost in
sorbed in the upper jejunum (2). Thus avail- the feces. As a result of their gelling proper-
able carbohydrate may be absorbed at a later ties, soluble fibers appear to produce malab-
time from the lower jejunum or ileum. Solu- sorption of carbohydrate (15) but the insolu-
ble fibers such as guar gum increase the ble fibrous substances may insulate their
mouth to cecum in transit time and theoreti- starch in such a manner that portions of it
cally would allow more time for carbohydrate cannot be digested or absorbed (57).
absorption in the upper jejunum (13, 15). 6) Other factors such as alterations in gut
This theoretical advantage may be counter- hormones or pancreatic glucagon secretion
balanced by the gelling properties of these (46) may enhance glucose metabolism in the
fibers that retard digestion ofcarbohydrate in liver and account for lower postprandial
the jejunum. plasma glucose levels. The mechanisms out-
4) Available carbohydrate that is packaged lined above (1 through 5) would reduce post-
in plant fiber may be insulated from diges- prandial hyperglycemia and reduce glycos-
tive enzymes by its protective fibrous coat or uria but would not enhance glucose disposal.
plant fibers may partially inactivate digestive If high-fiber diets decrease insulin require-
enzymes (55, 56). ments and enhance glucose metabolism,
5) There is evidence (57) that some of the other mechanisms must be activated to pro-
starch in fiber-rich foods may not be digested mote glucose utilization. Further studies are
 PLANT FIBER: CARBOHYDRATE AND LIPID METABOLISM 355

required to explain the long-term effects of have little influence on serum cholesterol val-
plant fibers on glucose metabolism. ues. Pectin, guar gum, and soluble fibers,
exert distinct hypocholesterolemic effects.
Cholsesterol metabolism Extracts of certain pulses or legumes com-
monly eaten in India have profound effects
Animal studies on the serum cholesterol. When diets were
Extensive studies in experimental animals supplemented with cholesterol, serum choles-
indicate that the addition of certain plant terol values were 5-fold higher in control
fibers to the diet is accompanied by signifi- animals than in those receiving blackgram, a
cant reductions in serum cholesterol concen- soluble storage polysaccharide. Whereas the
trations. Representative studies of rats are soluble fibers reduce the total cholesterol con-
presented in Table 1. Wheat bran or cellulose tent, the influence of these fibers on the cho-
lesterol content of specific lipoprotein frac-
tions has not been well delineated.
TABLE I
These reductions in serum cholesterol val-
Influence of plant fibers and fiber-rich
plants on serum cholesterol ues were accompanied by decreases in the
values of rats cholesterol content of liver, aorta, and other

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tissues. In some animal models, supplement-
Cholo-

No. of
terol ing the diet with soluble fibers retarded the
Ingredient values Reference
studIes
.

I I. of
progression of experimental atherosclerosis
control) (89-93). These observations lend credence to
Bran the exciting hypothesis that a fiber-rich diet
Wheat 6 100 58-63 may delay the development of atherosclerosis
Soy 1 82 64 in humans (94-96).
Rice 2 73 65, 66
Agar 3 108 63, 67, 68 Human studies
Alginate 3 106 67-69
Cellulose 9 96 61, 63, 67, 68, 70-74 Certain plant fibers also exert a distinct
Carrageenan 2 84 63, 75 serum cholesterol-lowering effect in humans
GuarGum 3 81 60,75,76 (Table 2). The insoluble plant fibers, as noted
Pectin 13 80 61, 63, 68, 69, 74-82
in animal studies, are relatively ineffective in
Pulses
lowering the serum cholesterol while the wa-
Bengal gram 2 72 70, 83
Horsegt.sm 2 42 83, 84 ter soluble fibers are moderately effective.
Redgram 2 3 1 83, 84 Our summaries (Tables I and 2) are oversim-
Blackgram 5 20 83, 85-88 plified because different quantities of fibers

TABLE 2
Effects of plant fibers on blood lipids of man

Blood lipids
(I; of control)
Source of fiber No. Plant fiber References

Cholesterol Triglycerides

g/dav

Wheatbran 14 17(7-25) 101 97 97-110

I 25 75 111
Whole oats 1 12 89 1 12
Bagasse 2 10.5 100 99 113, 114
Cellulose 3 16 100 100, 115, 116
I 100 75 117
Psyllium 2 9.6 87 118. 119
Pectin 10 25(12-45) 87 99 104,115,120-127
Guargum 3 24(12-36) 84 104, 121, 128
Bengal gram I Unknown 78 129
Multiple plant fibers I 60 97 86 130
1 Unknown 84 76 131
1 82 75 86 47

I: Numbers are mean va lue with rang es in parentheses.

356 ANDERSON AND CHEN

from different sources were used. The studies Intestinal handling of cholesterol. Plant fi-
with bran and with cellulose suggest that the ber supplements are accompanied by in-
response of the serum cholesterol may be creased fecal excretion of neutral sterols such
dose related (1 1 1, 1 17). Most investigators as cholesterol and plant sterols as well as bile
have indicated simply that they used wheat acids ( 105, 1 18, 124, 125). Cholesterol is
bran and the plant fiber content was not bound by certain fibers that may retard its
provided. Wheat bran can be characterized absorption. The formation of gels in the in-
as coarse or fine and also as light (with a low testine and colon may partition cholesterol in
flour content) or heavy (with a high flour such a manner that gut absorption is reduced.
content). For example, light bran has a plant Whether plant fibers alter the incorporation
fiber content of approximately 45% whereas of cholesterol into chylomicrons and very low
heavy bran has a fiber content of only 16% density lipoproteins (VLDL) or influence
(132, 133). For these reasons, many of the their secretion has not been examined. The
studies using wheat bran are difficult to in- several mechanisms by which plant fibers
terpret. may alter bile acid absorption or metabolism
The available data clearly indicate that include: binding of bile acids, sequestration
soluble fibers such as pectin, guar gum and of bile acids in fiber gels, selective binding of

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bengal gram have distinct hypocholestero- deconjugated bile acids to alter the composi-
lemic effects in man. Whole oats contain tion of the bile acid pool, influencing the
about 6% insoluble fiber and 3% soluble fiber bacterial metabolism of bile acids and, fi-
( 133). The hypocholesterolemic effect of nally, by altering the bacterial flora of the
whole oats (1 12) may be related in part to colon with resultant changes in bile acid me-
their soluble fiber content. Our preliminary tabolism (61, 81, 124, 125, 134-137).
studies (J. W. Anderson, unpublished data) Hepatic metabolism of cholesterol. Synthe-
utilizing an oat bran preparation containing sis, metabolism, and release of cholesterol
approximately 25% plant fiber further docu- from the liver may be directly or indirectly
ment the hypocholesterolemic effects of oat altered by plant fibers. Some studies (64, 138)
preparations. These observations with soluble suggest that hepatic cholesterol synthesis is
plant fibers suggest that these substances may increased by plant fiber intake whereas others
have a role in the treatment and prevention (139) suggest that synthesis is decreased. The
of hypercholesterolemia in man. Further major hypothesis for the hypocholesterolemic
studies are required to delineate the effects of effect of plant fibers is that the depletion of
these plant fibers on the low density lipopro- bile acids by fecal excretion diverts choles-
tein cholesterol and the high density lipopro- terol into the bile acid pool and less choles-
tein (HDL) cholesterol concentrations (120, terol is available for incorporation into lipo-
140, 141). Our preliminary observations (J. proteins for release in the venous circulation
W. Anderson and D. Rees, unpublished data) (64, 125, 129, 134). However, plant fibers may
suggest that long-term therapy with a diet alter cholesterol synthesis or liproprotein Se-
containing approximately 60 g of plant fiber cretion in a variety of other ways which have
per day is accompanied by a distinct reduc- not yet been assessed. For example, short-
tion in the serum low density lipoprotein chain fatty acid metabolites of certain fibers
cholesterol content and an elevation of the may alter cholesterol synthesis.
HDL cholesterol content to values above Metabolism of lipoproteins. Cholesterol is
those of our control or normal population. contained in all of the major lipoprotein par-
tides; factors that control the synthesis and
Proposed mechanisms
catabolisms of these lipoproteins are still un-
Various plant fibers may influence serum der intensive investigation. In the gut choles-
cholesterol concentrations by three principal terol is incorporated into chylomicrons,
mechanisms: altering intestinal absorption, VLDL and HDL, (142) while the liver se-
metabolism and release of cholesterol; alter- cretes VLDL that contain cholesterol. Two
ing hepatic metabolism and release of choles- studies (64, 78) suggest that plant fiber intake
terol; or altering peripheral metabolism of enhances cholesterol removal from plasma
lipoproteins. but the mechanism is unclear. Conceivably
 PLANT FIBER: CARBOHYDRATE AND LIPID METABOLISM 357

plant fibers could influence the size of chy- (Table 2). The major effects of plant fibers
lomicrons released from the gut and this on fasting triglyceride values appears to be in
would influence their subsequent metabo- patients with fasting hypertriglyceridemia
lism. Plant fibers may alter the proportion of (101, 102).
cholesterol incorporated into chylomicrons, Our preliminary studies strongly suggest
VLDL, and HDL. Also, plant fibers might that when a high plant fiber intake is coupled
influence the size of VLDL particles released with a high-carbohydrate diet, the anticipated
from the gut or liver. Furthermore, plant rise in fasting triglyceride values does not
fibers might influence the composition of lip- occur. When patients with hypertriglyceride-
oproteins by altering the ratios of: cholesterol mia are fed a high-carbohydrate, high-fiber
esters to cholesterol, triglyceride, or phospho- diet, a significant reduction in serum triglyc-
lipids; lipid fractions to apoproteins; or of one eride values is observed. The composition of
apoprotein to another. These alterations in four diets that we have used (47, 48, 145) are
the composition of lipoproteins could influ- summarized in Table 3. When we fed a high-
ence the subsequent metabolism of these par- carbohydrate, low-fiber diet to 12 men with
ticles. diabetes a consistent and significant rise in
fasting serum triglyceride values was ob-

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Triglyceride metabolism served (Fig. 8A). However, when we fed a
high-carbohydrate, high-fiber diet to 12 in-
Human studies dividuals with diabetes there was a reduction
Plant fibers influence the absorption and in average triglyceride values (Fig. 8B). The
metabolism of triglycerides and fatty acids triglyceride-lowering effects of high-carbo-
but these effects may not be discerned by hydrate, high-fiber diets was more evident in
measuring serum triglyceride values after an patients with hypertriglyceridemia (Fig. 8C).
overnight fast. Most investigators have re- Furthermore, we have treated three men with
ported that wheat bran or pectin do not influ- moderately severe hypertriglyceridemia with
ence fasting serum triglyceride values in hu- weight-maintaining high-carbohydrate, high-
mans (Table 2) or experimental animals (143, fiber diets; triglyceride values have dropped
144). However, high-fiber diets are usually to half of those observed on the control diet
accompanied by reductions in the total lipid (Fig. 8D). These observations suggest that
and triglyceride content of liver in experi- plant fibers may have a role in the treatment
mental animals (59, 68, 74-76, 78, 84, 87, 143, of patients with hypertriglyceridemia.
144). When diets containing generous Plant fiber in the diet appears to exert an
amounts of fiber from a variety of plant even greater effect on postprandial triglycer-
sources are fed, a modest reduction in fasting ide values than on fasting values. On these
serum triglyceride values has been observed diets we have made 1 1 measurements of se-

TABLE 3
Composition of representative 2000 kcal diets

Diets
Component
37C HC. LF 43C HC. HF

g kca! g kcal g #{182}kcal g kca!

Protein 102 20 49 10 98 20 95 19
Carbohydrate
Total available 183 37 370 74 215 43 360 72
Simple 89 30 100 90
Complex 94 340 115 270
Fat 95 43 36 16 83 37 20 9
Plantfiber 20 24 31 74

(I Values were derived from Tables in Reference 133. Thirty-seven percent carbohydrate control diet. High-
carbohydrate, low-fiber diet. Forty-three percent carbohydrate control diet. High carbohydrate, high fiber
diet.
358 ANDERSON AND CHEN

A. B. C.

300

C HC

0
HC
LF HF
C-)
200
-I

1.
4 3C

37C

I-:i
w
(I) 00

I
z
:1)

12 7
I.
P <0.01 N.S. <0.001 N S.

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FIG. 8. Fasting serum triglyceride (FTG) values on control (C) and high-carbohydrate diets. The compositions
of these diets are given in Table 3. A, twelve diabetic men on 37 C diets had FTG ranging from 39 to 209 mg/dl; on
high-carbohydrate (HC), low-fiber (LF) diets, FTG increased significantly. B, twelve diabetic men on 43 C diets had
FTG raninging from 74 to 234 mg/dl: on HC, high fiber (HF) diets, FTG were slightly lower. C, seven diabetic men
on 43 C diets had FTG ranging from 241 to 394 mg/dl; on HC, HF diets, FTG were significantly lower. D, three
nondiabetic men with type IV hypertriglyceridemia on 43 C diets had FTG ranging from 925 to 1 1 15 mg/dl; on HC,
HF diets, FTG were lowered to one-half of control values.

rum triglyceride values over a 24 hr period of further studies are required to delineate the
time. We have noted the highest value (peak effects of high fiber diets on serum triglycer-
value) and calculated the total area (TGA) ide values.
for the 24 hr period. To correct for differences
Proposed mechanisms
in fasting triglyceride values we have deter-
mined the incremental change in serum tri- The three principal mechanisms outlined
glycerides. The incremental change area is for cholesterol metabolism could also explain
the total area - fasting triglyceride x 24. As a triglyceride-lowering effect of plant fibers.
reported earlier (146), the total area, incre- These fibers may alter intestinal handling of
mental changes and peak triglyceride values triglycerides and fatty acids, alter hepatic me-
were significantly lower on high carbohydrate tabolism and release or alter peripheral me-
diets than on control diets (Fig. 9). When the tabolism of lipoproteins. Because most ob-
control diets contained moderate amounts of servers have not been impressed with the
plant fibers, the incremental area and post- influence of plant fibers on plasma triglycer-
prandial peak values were significantly lower ide values, these proposed mechanisms have
than values on the low-fiber control diet. not been examined in detail.
When high-carbohydrate, high-fiber diets Intestinalhandling offats. When plant fiber
were fed, postprandial values were consist- intake is increased, fecal fat excretion is often
ently lower than fasting values and the incre- increased slightly (25-27, 33). It seems un-
mental change was negative. Furthermore the likely that this small increase in fecal fat,
postprandial peak values were lower than the including some waxes and fiber-related lipids,
fasting values (not shown). Thus, on a high- has an important impact on triglyceride me-
carbohydrate, high-fiber diet, postprandial tabolism. However, it does seem likely that
triglyceride values fell below the fasting val- the absorption of fat is slower and may occur
ues and remained below this value through- lower in intestinal tract when fiber is present.
out the Jay. These observations strongly sug- Our studies (Fig. 9) that demonstrate lower
gest that high fiber diets are associated with postprandial peak values for serum triglyc-
lower postprandial triglyceride values than erides suggest that plant fibers may attenuate
observed on low-fiber diets containing similar the postprandial rise in chylomicrons.
amounts of carbohydrate and fat. Obviously, Hepatic metabolism of fatty acids and tn-
 PLANT FIBER: CARBOHYDRATE AND LIPID METABOLISM 359

37C INCREMENTAL CHANGES patic ketogenesis by providing excessive

. 3000 amounts of ketone precursors (see Reference
C 150). All of these interacting influences may
2000 tend to decrease triglyceride synthesis or the
HC,HF
.<
release of VLDL resulting in lower fasting
43C and postprandial triglyceride values.
000
Metabolism of lipoproteins. As outlined in
I
the discussion of cholesterol metabolism, al-
-J
C.,
terations in the size or composition of chy-
t_5O0
A B B C lomicrons or VLDL might influence their
subsequent metabolism. In addition, if tri-
POSTPRANDIAL PEAK
glyceride-rich particles were presented to pe-
C VALUES
C,,
ripheral tissues at a slower and steadier rate,
0 hydrolysis by the lipoprotein lipases might
C.)
proceed in a more optimal fashion. Further
C., studies are required to examine the influence
of plant fibers on the production and removal
(

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rates of triglyceride-rich lipoproteins.
C. A B B C
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