Neuroscience & Biobehavioral Reviews, Vol. l I, pp. 131-153. r Pergamon Journals Ltd.. 1987. Printed in the U.S.A.

0149-7634/87 $3.00 + .00

Carbohydrate Taste, Appetite, and

Obesity: An Overview I
ANTHONY SCLAFANI"
Department of Psychology, Brooklyn College and the Graduate School
City University of Nelr York, Brooklyn, N Y 11210
Received 5 September 1986

S C L A F A N I , A. Carbohydrate taste, appetite, and obesity: All overview. N E U R O S C I B I O B E H A V REV 11(23 131-153,
1987.--This paper reviews previous research on sugar appetite and sugar-induced overeating and obesity in the rat, and
previews new findings reported in this issue on the rat's taste and appetite for starch and starch-derived polysaccharides. A
variety of behavioral methods demonstrate that rats are very attracted to the taste of sugars. At low molar concentrations
their order of preference is maltose > sucrose > glucose = fructose, while at high concentrations it is sucrose > maltose >
glucose = fructose. New findings indicate that rats are also very attracted to starch-derived polysaccharides, such as
Polycose. In fact, Polycose is preferred to sucrose and maltose at low concentrations, and is second only to sucrose in
preference at high concentrations. Furthermore, rats taste Polycose as qualitatively different from sucrose. These and
other findings suggest that rats as well as other rodents have different taste receptors for sugar and starch. The role of these
two taste systems in carbohydrate-induced overeating and obesity is discussed. Rats tend to overeat and become obese
when fed sugar or polysaccharide diets, but this response depends critically on the form of the diet. Presenting the
carbohydrate as a solution or gel supplement to the diet is much more effective in promoting hyperphagia and obesity than
is presenting it as a powder supplement or as part of a composite diet. The differential response to hydrated and dehydrated
foods may occur because carbohydrates are absorbed at a faster rate in hydrated forms than in dehydrated forms. The
postingestive actions of carbohydrates also modulate the rat's preference for carbohydrate tastes and flavors. Other factors
that influence carbohydrate appetite and intake include the availability of the carbohydrate source and the macronutrient
composition of the diet. Thus, several factors in addition to taste determine the response of rats to carbohydrate foods.

Sugar Sweet taste Starch Polysaccharide taste Hyperphagia Obesity

Conditioned food preferences Diet form Diet availability Macronutrient balance

CONTENTS F.Qualitative Differences in Polycose and Sugar

I. I N T R O D U C T I O N . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Tastes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
A. Carbohydrate Structure . . . . . . . . . . . . . . . . . . . . . . 132 G. Influence of Saccharide Length on Polysaccharide
Appetite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
11. S W E E T TASTE APPETITE . . . . . . . . . . . . . . . . . . . . . 134
H. Effects of Gustatory Deafferentation on Sac-
A. Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134 charide Appetite . . . . . . . . . . . . . . . . . . . . . . . . . 142
B. Hedonic Response . . . . . . . . . . . . . . . . . . . . . . . . . . 135 I. Effects of Anosmia on Polycose Appetite . . . . . . 142
I11. S U G A R - I N D U C E D H Y P E R P H A G I A AND OBE- J. Sham-Feeding Response to Polycose and Sucrose 142
SITY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135 K. Saccharin-Carbohydrate Taste Interactions . . . . . 142
A. Composite Diet Studies . . . . . . . . . . . . . . . . . . . . . . 135 L. Species Differences in Saccharide Taste Prefer-
B. Diet Option Studies . . . . . . . . . . . . . . . . . . . . . . . . . 138 ences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
M. Sex Differences in Saccharide Taste Preferences 143
IV. P O L Y S A C C H A R I D E - I N D U C E D H Y P E R P H A G I A
AND O B E S I T Y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 N. Starch Preferences in Rats . . . . . . . . . . . . . . . . . . . 143
VI. I N T E G R A T I O N . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
V. P O L Y S A C C H A R I D E T A S T E AND APPETITE . . . 140
A. Carbohydrate Taste . . . . . . . . . . . . . . . . . . . . . . . . . 143
A. Effects of Saccharide Type, Form, and Taste .. 140
B. Carbohydrate Appetite . . . . . . . . . . . . . . . . . . . . . . . 145
B. Effects of Taste and Form on Polycose Preference
C. Importance of Carbohydrate Taste . . . . . . . . . . . . 147
......................................... 140
C. Origin of Polycose Appetite . . . . . . . . . . . . . . . . . . 141 D. Other Nutrient Receptors? . . . . . . . . . . . . . . . . . . . 148
E. Relevance to Human Feeding and Obesity . . . . . 148
D. Hedonic Response to Polycose and Sugar Solu-
tions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 VII. C O N C L U S I O N S . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
E. Polycose and Sugar Taste Preference Thresholds 141 VIII. R E F E R E N C E S . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

~The preparation of this paper was supported by grants from the National Institutes of Health (DK-31135) and from the Faculty Research
Award Program of the City University of New York. The author thanks Dr. Israel Ramirez for his helpful comments on an earlier version of
this paper. The author is also most grateful to Dr. George Collier who served as a special reviewer for the present and following papers in this
issue.
-"Requests for reprints should be addressed to Dr. Anthony Sclafani, Department of Psychology, Brooklyn College, Brooklyn, NY 11210

131
 132
Carbohydrate Structure
(Simplified)
Glucose Q
Fructose 0
Sucrose
Maltose 0 - 0
Malt ooligosaccharides ~ E . I ~ J - - - - " U ~
Maltopolysaccharides ~ - - - evoke taste sensations in humans, but the "taste worlds" of
Amylose
Amylopectln
- _
FIG. I. Simplified representation of carbohydrate structure. Note
that maltooligosaccharides contain two to 10 glucose units and
therefore include maltose; maltopolysaccharides contain more than
10 glucose units. Only portions of the amylose and amylopectin
starch molecules are illustrated (each circle represents a glucose
unit).
"Among the three fundamental classes of foodstuffs: carbohydrates,
lipids and proteins, that alone of carbohydrates with their sugary
flavour has given rise during evolution to the differentiation of an
autonomic and homogenous receptor system."
Le Magnen, cited in [186].
INTRODUCTION
CARBOHYDRATES, in the form of sugar and starch, rep-
resent a major source of food energy for a great number of
animal species. An even more abundant carbohydrate is cel-
lulose but only certain species (e.g., ruminants) can utilize
cellulose. Except for some fruits, plants contain much more
starch than sugar, but it is sugar with its sweet taste, that is
the most sought after carbohydrate. The sweetness of sugar
in fact has been described as a supernormal reinforcer that
humans find difficult to resist [25]. Motivated by their
"'sweet tooth," humans have over the centuries perfected
the refinement of pure sugar (sucrose, glucose, fructose)
from sugar cane, sugar beets and, more recently, from corn.
Sugar, once a delicacy, is now a staple. (Man's preoccupation
with sugar has a long and fascinating history, see [123].)
The preference for sweet taste is not unique to humans
but is widespread in the animal kingdom, although there are
SCLAFANI
notable exceptions (e.g., cats, some species of birds)
[100,139]. Compared to sugar, starch is rather bland to the
human palate and has been assumed to be tasteless to other
animals as well [17, 52, 81, 90, 121, 172]. However, recent
studies in my laboratory indicate that rats and other rodents
taste starch and starch-derived polysaccharides and find the
taste quite attractive. Our studies further suggest that rats
taste polysaccharides as qualitatively different from sucrose.
Thus it appears that, contrary to the statement by LeMagnen
quoted above, some animal species have two carbohydrate
receptor systems, one for sugar and one for starch. Other
investigators have speculated on the existence of a starch
taste sensitivity in rats [16] but this possibility has not been
previously explored. As noted by Dethier [51], animal taste
studies have generally been limited to those substances that
animals and humans are not the same.
The discovery of a polysaccharide taste sensitivity in rats
emerged from studies designed to clarify the role of sweet
taste in the hyperphagia and obesity promoting effects of
high-sugar diets. In the course of this research it became
evident not only that rats taste polysaccharides as well as
sugars, but that their appetite for these carbohydrates is
modified by postingestive factors. The following reports in
this issue provide a detailed account of our studies on
polysaccharide and sugar appetite. The present paper sum-
marizes these studies and presents an overview of previous
research on sugar appetite and sugar-induced overeating and
obesity in the rat.
Carbohydrate Structure
To facilitate discussion, carbohydrate structure and ter-
minology will be briefly reviewed (see Fig. 1). The carbo-
hydrates (or saccharides) of interest here include the
monosaccharide sugars: glucose and fructose; the disac-
charide sugars: sucrose and maltose; and starch and its de-
rivatives. Starch exists in two forms: an amylose form which
consists of long linear (unbranched) chains of glucose (glu-
cose polymers) united by c~-l,4-1inkages; and an amylopectin
form, which consists of branched molecules containing many
sl~ort linear chains (20 to 30 glucose residues) connected to-
gether at branch points by a-l,6-1inkages (Fig. 1) [86]. When
st~trch is hydrolyzed, shorter glucose polymers are
prbduced, i.e., maltopolysaccharides and maitooligosac-
ch'arides, which are ultimately hydrolyzed to glucose in the
gu t. Maltooligosaccharides are glucose polymers containing
2 !o 10 glucose units, while maltopolysaccharides contain
more than 10 glucose units. For purposes of the present
paper the term "'polysaccharide" will refer to starch-derived
saccharides containing three or more glucose units.
iln commercially prepared starch hydrolysates (e.g., dex-
trims,h maltodextrins, corn syrups) the starch molecules are
partially hydrolyzed with acid and/or enzymes to produce
mixtu~res containing varying amounts of glucose, maltose,
maitooligosaccharides, and maltopolysaccharides; in some
products the glucose is enzymatically converted to fructose,
I.e. I, high-fructose syrups [188]. Most hydrolysate products
hax~e a high sugar content and are used as sweeteners, e.g.,
corn syrup sweeteners, while others are designed to have a
low sugar content and are used as nutritional supplements.
For example, Polycose (Ross Laboratories), which we have
used extensively in our research, is a corn starch hydrolysate
that contains, by weight, approximately 2% glucose, 7% mal-
tose, 55% maltooligosaccharides, and 36% maltopolysac-
CARBOHYDRATE TASTE AND APPETITE 133

TABLE 1
CARBOHYDRATE TASTE TESTS: A LIST OF REPRESENTATIVE STUDIES

Type of Test Tastant Reference

One-Bottle, Brief (1 to 4 min) Sucrose [47,220]

Glucose [47, 48, 162]
Fructose [47]
Maltose [48,162]
Polycose [ 162]
One-Bottle, Short-Term Sucrose [24, 38, 78, 159, 177, 216]
(10 to 60 min) Glucose [24, 116, 159, 177]
Fructose [24]
Maltose [78,159]
Polycose [ 159]
Saccharin [44, 174, 217]
One-Bottle, Long-Term Sucrose [65, 183, 213]
(24 hr) Glucose [291
Saccharin [7.29, 65, 183, 213]
Two-Bottle, Brief Sucrose [6,34]
(Solution vs. Water)
Two-Bottle, Short-Term Glucose [90]
Saccharin [5,129]
Two-Bottle, Long-Term Sucrose [24, 27, 37, 60, 78, 80, 102,
135, 149, 160, 166, 187]
Glucose [24, 60, 90, 135, 149, 178]
Fructose [24]
Maltose [60, 78, 135, 149, 161, 166]
Polycose [60, 160, 161, 166]
Saccharin [41, 80, 160, 174, 189, 196]
Glucose + Saccharin [160, 178, 195]
Sucrose + Saccharin [160]
Polycose + Saccharin [160]
Two-Solution, Brief Sucrose vs. Sucrose [6,217]
Sucrose vs. Saccharin [218]
Sucrose vs. Polycose [161]
Glucose vs. Maltose vs. [162]
Polycose
Two-Solution, Short-Term Sucrose vs. Sucrose [215]
Sucrose vs. Glucose [177]
Sucrose vs. Maltose [78]
Sucrose vs. Polycose [159, 161, 172]
Maltose vs. Polycose [159]
Saccharin vs. Saccharin [217]
Two-Solution, Long-Term Sucrose vs. Sucrose [37, 102, 137]
Glucose vs. Glucose [19, 89, 90, 136, 137, 178]
Sucrose vs. Glucose [24,160]
Sucrose vs. Polycose [160,161]
Sucrose vs. Saccharin [41]
Glucose vs. Fructose [24]
Glucose vs. Saccharin [181, 193, 194, 196]
Saccharin vs. Saccharin [41]
Glucose + Saccharin vs. [27]
Glucose
Multiple-Solution, Long-Term Sucrose [27]
Saccharin [217]
Sham-Feeding, One-Bottle Sucrose [96, 124, 134, 165, 167, 208]
Glucose [96, 124, 165, 167]
Fructose [96]
Maltose [96]
Polycose [134,167]
Sham-Feeding, Two-Solution Sucrose vs. Polycose [134]
Intraoral Infusions, Sucrose [72, 75, 92, 156]
Oral-Facial Response Glucose [ 13]
Saccharin [91,92]
134
TABLE I
IContinued)
Type of Test Tastant
Bar-Pressing, No Choice Sucrose
Glucose
Saccharin
Bar-Pressing, Two-Choice Sucrose vs. Sucrose
Sucrose vs. Maltose
Runway/Maze. No Choice Sucrose
Glucose
Saccharin
Runway/Maze, Two-Choice Sucrose vs. Sucrose
Saccharin vs. Water
One-bottle=only one drinking fluid available; Two-bottle=solution vs. water preference test.
charides (Ross Laboratories, personal communication). De-
spite its low sugar content, Polycose and other starch
hydrolysates are as rapidly digested and absorbed as is free
glucose [152,205].
SWEET TASTE APPETITE
Among nonhuman animals the appetite for sweets is most
extensively documented in the laboratory rat. In short-term
tests nondeprived rats not only avidly consume sweet-tasting
foods and fluids that are readily available, but they will also
run down alleys, negotiate mazes, or press bars to obtain
sweet rewards. A dramatic example of the rat's sweet tooth
is provided by the observation that rats given access to a
warm nest box containing chow and water voluntarily left
the nest to run down a 16 meter long runway kept at - 15C
to obtain a sweet food or drink (shortcake or Coca Cola) [23].
As indicated in Table I, the sweet taste appetite of the rat
has been investigated using a variety of tastants and behav-
ioral methods. Included in the table are the sugars sucrose,
glucose, fructose, and maltose, and the artificial sweetener
saccharin. (Table 1 and subsequent tables also include refer-
ences to Polycose which are discussed latter in this paper.)
Not discussed here are less extensively studied sugars (e.g.,
lactose and galactose, see [48, 92, 135, 149]) and artificial
sweeteners that rats are unresponsive to (e.g., cyclamate,
dulcin, and aspartame, see [62, 158,204]). Most research has
involved solutions, but some studies have measured the rats
hedonic response to sugar in powdered form. For example,
in early studies P. T. Young [214] demonstrated that rats
prefer sucrose powder to flour or wheat powder. As will
become apparent, the physical form of carbohydrate is an
important determinant of appetite and intake.
Methodology
The many behavioral test methods used to study carbo-
hydrate appetite can be classified according to: test duration
(brief, 1-5 min; short-term, 10-60 min: or long-term, 24 hr):
degree and type of choice (one-bottle, two-bottle, multi-
bottle; solution vs. water or solution vs. solution): and re-
sponse type (ingestional or instrumental). Ingestional
methods include measures of intake, lick rates, and orofacial
responses while instrumental methods include performance
SCLAFANI
Reference
[36, 38, 40, 73, 74, 95]
[28]
[28, 35, 551
142, 43, 1551
1781
[39, 103, 182, 209, 219]
[175]
[88, 175, 184]
[106,216]
[174]
in runway, maze, or bar-pressing tasks (see [180]). These
distinctions are important because the various behavioral
measures produce different outcomes. For example, tests
that allow an animal to consume a sugar solution to satiety
produce results different from those that restrict solution
intake. Thus, the rat's response to sucrose in brief intake
tests or in bar-pressing tests increases monotonically as su-
crose concentration increases, whereas in short-term (1 hr)
or long-term (24 hr) intake tests the rat's sucrose consumption
intake increases but then decreases as concentration in-
creases from 1% to 64% [24, 37, 40, 47, 149]. The "'turn-
d o w n " at the higher concentrations is due to the postinges-
tive satiating effects of the solutions [138]. (Note, the turn-
down is not apparent when intake is expressed in terms of
the amount of solute rather than solution consumed [37].)
The influence of postingestive factors on sugar solution in-
take is clearly demonstrated by the results obtained in
sham-feeding tests in which the ingested solution drains out
an esophageal or gastric fistula. In this case, consumption
(sham-feeding) increases monotonically as sugar concentra-
tion increases [124,208].
Choice is another critical factor. For example, in one-
bottle short- or long-term tests rats typically consume more
of a dilute sugar solution (e.g., 4c~ sucrose) than of a concen-
trated solution (e.g., 32% sucrose), yet when given the
choice between the two solutions rats prefer the concen-
trated to the dilute solution [24, 37, 38,216]. Thus it is impor-
tant to distinguish taste "'preference" as measured in choice
tests from taste " a c c e p t a n c e " as measured in one-bottle
tests; the most acceptable solution in terms of amount con-
sumed may not be the most preferred solution. The type of
choice given the animal must also be considered. In the
classic Richter two-bottle test [149] animals are given the
choice between a test solution and water, but with sugar
solutions at above threshold concentrations this is essen-
tially a one-bottle test since the animal drinks little or no
water except at very high sugar concentrations. Thus, al-
though it is a common procedure, judgments of the relative
palatability of different sugars based on the Richter-type test
can be misleading. For example, in solution vs. water tests
rats show a greater preference for glucose over water than
they do for sucrose over water yet in two-solution tests they
strongly prefer sucrose to glucose [24]. As Pfaffmann [138]
has emphasized, "'true preferences" can only be established
when taste stimuli are directly compared in choice tests.
C A R B O H Y D R A T E TASTE A N D APPETITE
Hedonic Response
Taste intensity. With these methodological considerations
in mind, the following conclusions can be made concerning
the rat's response to sweet taste stimuli. With respect to
taste intensity, the rat's hedonic response to sugars (sucrose,
glucose, fructose, maltose) increases as concentration in-
c r e a s e s when postingestive factors are minimized by using
brief intake, sham-feeding, or bar-pressing tests [40, 42, 47,
48, 73, 96, 124,208,220]. However, this is not the case with
saccharin. As saccharin concentration increases, palatabil-
ity, as measured by one or two-solution intake or sham-
feeding tests, increases but then decreases as the concentra-
tion exceeds 0.4 to 0.6% [24, 165,217]. (Note that this turn-
down is not seen in bar-pressing tests that involve small (0.03
ml) reinforcements [35].) The rat's reduced consumption of
the more concentrated saccharin solutions is not due to
postingestive factors [165], but has been attributed to the
bitter taste component of saccharin [138].
Relative palatability. With respect to the relative palata-
bility of the different sugars less definitive conclusions can
be made based on the published data. Despite the extensive
literature on sweet taste, relatively few studies have directly
compared the rat's preference for different sugars. The
available data indicate that the rat's relative preference is:
sucrose > fructose /> glucose [24, 47, 177]. The relative
palatability of maltose is less certain, but some data suggest
that at low concentrations maltose is the most preferred sugar,
while at high concentrations it is less preferred than sucrose
[48, 78, 96, 149]. To clarify this issue, we recently systemati-
cally compared the preference of adult female rats for the
four sugars using brief (3-rain) two-solution preference tests
(Sclafani and Mann, in preparation). At the lowest concen-
tration tested (0.03 M) the order of preference was maltose >
sucrose > glucose = fructose, while at the highest concen-
tration (I M) it was sucrose > maltose > glucose > fructose.
The finding that at low concentrations maltose is the most
preferred sugar is surprising since, of the four sugars, mal-
tose is the least abundant in nature [173]. Furthermore, mal-
tose is the least effective of the four sugars in stimulating the
rat's chorda tympani nerve (CTN) [140]. Pfaffmann [140] has
referred to the discrepancy between the rat's behavioral re-
sponse and its CTN response to maltose as the "'maltose
p a r a d o x " because with the other three sugars the behavioral
and neural response profiles are in agreement. The respon-
sivity of the whole nerve, however, does not necessarily
reflect t ~ t of individual taste fibers, and some single fibers
have I~en found to be more responsive to maltose than to
sucrose [9]. These and other findings indicate that the rat has
two (or more) taste receptors for sugars. The possible func-
tional significance of the rat's " m a l t o s e " receptor is dis-
cussed further below.
The palatability of saccharin relative to the sugars has not
been extensively explored, but the available data indicate
that it is not very palatable. Two-solution preference tests
indicate that the most preferred saccharin solutions (0.2 to
0.4%) are equal in palatability (isohedonic) to sucrose solu-
tions of only 2 to 4% [41,218]/see also [193]). This may be
because of saccharin's bitter after-taste, although it is also
possible that rats simply do not taste saccharin as sweet as
sucrose [165]. The addition of a small amount of glucose to a
saccharin solution substantially improves its palatability
[195]. This synergistic effect has been attributed to the glu-
cose increasing the sweetness of the saccharin solution and
inhibiting its bitter taste [179]. Yet despite the fact that rats
135
consume enormous amounts of a glucose + saccharin (G+S)
solution, some findings indicate that the G + S solution is less
palatable than concentrated glucose solutions [165].
Palatability changes. Rats display a hedonic response to
sucrose and saccharin at a very young age which suggests
that their sweet tooth is innate [75,92]. This does not mean,
however, that sweet preference is unmodifiable. Most nota-
bly, the rat's " g u s t " turns to "'disgust" if a sweet taste is
associated with visceral malaise [13]. More subtle modifica-
tions in sweet appetite may occur as the animal associates
the postingestive nutritive and/or colligative effects of sugar
with its taste [18]. For example, in long-term tests rats ini-
tially prefer concentrated glucose solutions to dilute solu-
tions but they reverse their preference after several days
apparently because of the discomforting postingestive ef-
fects produced by the hypertonic glucose solutions [19]. The
palatability of a saccharin solution also appears to change if
the animal associates the solution with caloric repletion or
depletion [26, 85, 111].
In addition to long-term shifts in palatability, variations in
the animal's nutritive state can produce temporary effects on
its appetite for sweets. Thus, food deprivation or insulin-
induced hypoglycemia enhances, and caloric repletion sup-
presses the hedonic response to sugar [49,70]. In a similar
fashion, experimentally-induced overweight and under-
weight enhances and suppresses, respectively, sweet taste
appetite [128]. The palatability of the animal's maintenance
diet also influences its short-term response to sweet taste
[125,163]. Orosensory stimulation alone may be sufficient to
temporarily satiate the animal's appetite for saccharin solu-
tions or dilute sugar solutions [127,208]. Recent findings
indicate that blood glucose availability affects the rat's re-
sponsiveness to sweet stimuli as measured by both elec-
trophysiological and behavioral tests, and this may be one
mechanism by which changes in nutritive state affects sweet
appetite and satiety [66,67]. However, other findings
demonstrate that the satiety for sugar can be quite specific.
For example, rats that are satiated for a glucose solution will
readily consume glucose powder [126]. Thus, a variety of
factors other than sweet taste influence the rat's appetite for
carbohydrates.
SUGAR-INDUCED HYPERPHAGIAAND OBESITY
Sugar has been extensively studied not only as a taste
stimulus but also as a nutrient. In general, two procedures,
referred to here as the "composite diet" method and the
"'diet option" (or "sugar option") method, have been used
to study the long-term effects of sugar on food intake and
body weight. In the composite diet method the sugar is in-
corporated into a semisynthetic diet whereas in the diet op-
tion method a separate source of sugar is given to the animal
in addition to a nutritionally complete diet. There are advan-
tages to both methods. For example, in the composite diet
method the influence of variations in the concentration or
type of sugar or other nutrients on food intake and body
weight can be assessed since diet composition is determined
by the experimenter. In the diet option method, on the other
hand, the animal selects the amount and proportion of sugar
it prefers. As discussed below, the two procedures yield dif-
ferent results.
Composite Diet Studies
Table 2 summarizes representative studies that have used
the composite diet method; the studies emphasized here in-
136 SCLAFANI

TABLE 2
EFFECTS OF HIGH-SUGAR COMPOSITE DIETS ON FOOD INTAKE, BODY WEIGHT, AND BODY FAT:
A SUMMARY OF REPRESENTATIVE STUDIES

Authors Sex Diet Food Body Body

Reference Age Type Intake Weight Fat

Allen and Leahy [1] Male 81% Sucrose [ -- T

Adult 81% Fructose J, -- T
81% Glucose ~ ~, --
81% Polysaccharide ~ -- --
Berdanier [111 Male 65% Sucrose 9 I" --
Weanling
Berdanier and Burrell [12] Male 65% Sucrose -- ~ ?
Weanling
Dulloo et al. [56] Both 66% Sucrose -- -- --
Weanling
Hallfrisch et al. [77] Male 54% Sucrose ? T 1'
Weanling
Hallfrisch et al. [76] Male 30% Sucrose 9 I" T
Weanling + 40% Fat
Lakshmanan et al. [109] Male 33% Sucrose __ __ 9
Weanling + 45% Protein
Laube eta/. [l IO] Male 68% Sucrose ? -- 1'
Weanling
Marshall et al. [I 18] Male 35-70% Sucrose-HP T T T
Adult 35-70% Sucrose-LP -- -- --
McCusker et al. [I 17] Male 65% Sucrose ~__ __ 9
Weanling
Michaelis et al. [122] Male 54% Sucrose '? q' ~'
Weanling 54% Glucose ? -- --
54% Invert Sugar 9 __ __
Reiser et al. [146] Male 54% Sucrose -- T T
Weanling
Williams and Szepesi [210] Male 56% Sucrose -- T --
Adult 56% Maltose T T 1'
56% Glucose -- ~ --
56% Invert Sugar -- -- --

Age=age of rats at start of experiment. T=increase, - - = n o change, ~=decrease, ?=data not reported or
statistical significance of reported change uncertain. HP=high protein diet, LP=low protein diet.
See references for additional details concerning diet composition, strain of rats, and duration of experiment.

v o l v e d m o d e r a t e to high protein c o m p o s i t e diets, see [143]
for d i s c u s s i o n of low protein diets. T h e typical p r o c e d u r e
used in t h e s e studies i n v o l v e s replacing some or all of the
s t a r c h in the c o m p o s i t e diet with sugar, usually s u c r o s e , for a
w e e k to several m o n t h s . T h e effects of the h i g h - s u g a r diet o n
food intake, b o d y weight a n d / o r b o d y fat, as well as on o t h e r
p a r a m e t e r s (e.g., p l a s m a insulin levels) are c o m p a r e d to
t h o s e o b t a i n e d with the c o n t r o l diet. I n s p e c t i o n of T a b l e 2
reveals c o n s i d e r a b l e variability in the feeding a n d b o d y
weight effects o b t a i n e d with high-sugar c o m p o s i t e diets.
S o m e studies report that h i g h - s u c r o s e c o m p o s i t e diets
p r o d u c e o v e r e a t i n g , o v e r w e i g h t , a n d obesity [118] but this is
not the typical effect. M o r e often, h i g h - s u c r o s e diets h a v e
b e e n f o u n d not to alter, or in s o m e cases, to r e d u c e caloric
intake [1, 12, 56, 109, 117, 146, 210]. N e v e r t h e l e s s , e v e n if
they do not elevate caloric intake, high-sugar diets m a y in-
c r e a s e b o d y weight a n d / o r b o d y fat; this indicates t h a t the
diets e n h a n c e feed efficiency [I, 12, 146, 210]. Finally, the
effects of high-sugar diets on b o d y weight a n d b o d y fat are
not always c o r r e l a t e d : in some c a s e s b o d y fat but not b o d y
weight is i n c r e a s e d , or less often, b o d y weight but not b o d y
fat is e l e v a t e d [1, 11, 110, 210].
C o m p a r i s o n s b e t w e e n different sugars indicate that in
s o m e , but not all studies, s u c r o s e diets h a v e a g r e a t e r effect
on b o d y fat, as well as on a variety o f o t h e r m e t a b o l i c pa-
r a m e t e r s , than do glucose, f r u c t o s e , or i n v e r t (glucose +
fructose) diets [1, 122, 210]. M a l t o s e diets h a v e also b e e n
f o u n d to i n c r e a s e b o d y fat more t h a n glucose diets [210]. T h e
CARBOHYDRATE TASTE AND APPETITE 137
TABLE 3
EFFECTS OF SUGAR OPTION DIETS ON FOOD INTAKE, BODY WEIGHT, AND BODY FAT:
A SUMMARY OF REPRESENTATIVE STUDIES

Authors Sex Diet Food Body Body

Reference Age Option Intake Weight Fat

Bukowiecki et al. [22] Female 12% S u c r o s e -- -- 1`

Adult 32% S u c r o s e 1" --
Coca-Cola t -- T
C a s t o n g u a y et al. [31] Male 32% Sucrose 1' 1"
Adult 62% Sucrose T T ?
16% Glucose 1' 1' ?
32% Glucose 1" ~' ?
D e s h a i e s et al. [50] Female 32% S u c r o s e 1` -- ?
Adult

F a u s t et al. [59] Male 16% S u c r o s e ? 1'

Adult
G r a n n e m a n and Wade [68] Male 32% S u c r o s e 1" T
Adult

H i r s c h et al. [82] Both 32% S u c r o s e I" T

Adult
H i r s c h et al. [83] Male 32% S u c r o s e T --
Weanling

Hirsch and Walsh [84] Male 32% S u c r o s e I" 1`

Adult
Hill et al. [81] Male 32% S u c r o s e -- 1'
Adult Starch P o w d e r -- --
K a n a r e k and M a r k s - K a u f m a n Male 32% S u c r o s e ~' --
[98] " Weanling
K a n a r e k and O r t h e n - Male 32% S u c r o s e 1" -- 1'
Gambill [99] Adult Sucrose Powder -- -- --
32% G l u c o s e T -- T
32% F r u c t o s e 1' -- T
K e n n e y and Collier [I01] Male 32% S u c r o s e -- -- ?
Adult
Rattigan and Clark [144] Male 37.5% S u c r o s e -- T 1'
Adult

M u t o and Miyahara [130] Male 40% S u c r o s e / H P I" )'? T?

Weanling 40% S u c r o s e / L P ,[ ~ --
Sclafani and X e n a k i s [1721 Male 32% S u c r o s e -- 1` 1"
Adult 32% Polycose T T 1`
Sclafani [157] Female 32% S u c r o s e 1` T T
Adult Sucrose Powder -- -- --
32% G l u c o s e ~ T 1'
Glucose Powder -- -- --
32% Polycose T T I'
Polycose P o w d e r -- -- --
T e a g u e et al. [190] Male 16% S u c r o s e -- -- '~
Adult
T r o u t et al. [192] Male 20% S u c r o s e T -- T
Adult 20% Glucose T -- T
Vasselli et al. [199] Female 40% S u c r o s e 1' 1' T
Adult

A g e = a g e of rats at start o f e x p e r i m e n t . 1`=increase, - - = n o change, $ = d e c r e a s e , ? = d a t a not reported or

statistical significance of reported change uncertain. H P = h i g h protein diet, L P = l o w protein diet.
See references for additional details concerning diet composition, strain of rats, and duration of experiment.
138
differential effects produced by sucrose and maltose as com-
pared to their component sugars (glucose and fructose) has
been referred to as the "disaccharide effect" [145]. The di-
saccharide effect is not always obtained, though, and it has
been described as a "'fragile" phenomenon [210].
Many factors are thought to contribute to the inconsistent
results obtained with the composite diet method. Differences
in the sugar and protein content of the diets, length of time on
the diet, age, sex, and strain of animals have all been men-
tioned as important variables [10,143]. One potentially im-
portant factor that has received little attention is the avail-
ability of the test diet to the animals; most studies do not
even describe the method of diet presentation. We recently
observed that simply using an antiscatter plate to minimize
food spillage from a standard food cup (Wahmann LC-306A)
was sufficient to block the rat's hyperphagic response to a
high-carbohydrate diet (Sclafani, Vigorito, and Pfieffer, un-
published findings). Another factor that has received little
attention in composite diet studies is palatability. Few
studies, for example, have measured the rat's preference for
the high-sugar diet over the control diet using two-diet
choice tests.
Diet Option Studies
Compared to the findings obtained with the composite
diet method, the results obtained with the sugar option
method have been more consistent. Table 3 summarizes the
results of experiments in which rats were fed, in addition to a
complete diet (e.g., chow) a separate source of sugar, usually
in the form of a sucrose solution. In the majority of the
studies surveyed, rats offered a sugar solution option con-
sumed about 60%, of their calories as sugar, and increased
their total caloric intake by 10 to 20% compared to control
rats fed only chow [22, 31, 50, 68, 82-84, 98, 99]. Fur-
thermore, compared to controls, the sugar solution rats usu-
ally gained more weight [31, 59, 68, 81, 82, 84, 144, 172] and
deposited more body fat [22, 59, 68, 82, 98, 99, 144]. In the
few studies that compared the effects of different sugars (su-
crose, glucose, fructose) no reliable differences were ob-
tained in caloric intake, body weight or body fat [31,99] (see
also [157]). Some of the variability in the food intake and
body weight results can be attributed to age and sex differ-
ences; young growing animals do not gain weight with a
sugar option [83, 98, 130] (see also [172]), and male rats are
less likely to overeat than are female rats when offered a
sugar solution (see Table 2).
There is one inconsistency in the data, however, that re-
mains to be explained. That is, in several studies involving
adult male rats, sucrose-induced overweight and/or obesity
was associated with increased caloric intake [31, 68, 82, 84,
99, 192], while in other studies [30, 81, 144, 172] overweight
occurred in the absence of hyperphagia. Composite diet
studies also demonstrate that sucrose-induced obesity can
occur in adult male rats with or without overeating [1, 118,
210]. Compared to starch, sucrose is digested and absorbed
faster, and the metabolic sequelae of this rapid absorption
(e.g, hyperinsulinemia) may explain how sucrose can in-
crease body fat without increasing caloric intake [64,145].
On the other hand, under some conditions sucrose-feeding
has been found to increase energy expenditure, perhaps due
to its effects on brown adipose tissue size and activity [22,
68, 99, 153, 190], and this could explain why rats sometimes
gain less weight than expected on high-sucrose diets, Why
sucrose increases feed efficiency under some conditions but
not others is not known.
SCLAFANI
Composite diet vs. diet option: diet,[i~rm. Not only is the
incidence of overeating, overweight, and obesity higher with
the diet option method than with the composite diet method,
but the magnitude of the response is also greater. In general,
rats increase their food intake, body weight, and body fat
more when given a sugar option than when fed a high-sugar
composite diet [143]. Why sugar option diets promote more
overeating and obesity than do high-sugar composite diets
has not been investigated but several factors are implicated.
Diet form, in particular, is likely to be an important variable:
in the composite diet experiments the high-sugar foods are
typically presented in a dry form, whereas in the diet option
experiments the sugar option is usually in liquid form. The
importance of diet form is indicated by the finding [99] that
rats given a sucrose solution option consumed more sucrose
and total calories than did rats given a sucrose powder option
(see below and [157]). Furthermore, rats fed a high-sucrose
composite diet in semiliquid form have been found to eat
more food and gain more weight than do rats fed the same
diet in dry form [143]. Possible reasons why liquid diets pro-
mote more overeating and obesity than do dry diets are dis-
cussed below and in latter sections of this paper.
Diet variety. Another factor that may contribute to the
hyperphagia promoting effect of a sugar option is variety.
Variety in the diet has been found to increase food intake and
weight gain [ 151 ], and in the sugar option condition, unlike in
the composite diet condition, the animal has two foods to
choose from. Alternatively, Kratz and Levitsky [107] have
argued that the hyperphagia and overweight obtained with a
sugar option diet is an artifact of limiting the animal's choice
of food to pure sugar and a complete diet. These inves-
tigators reported that rats allowed to self-select their diet
from separate sources of protein, fat, and carbohydrate
(starch) showed less of a hyperphagic response to a sucrose
option than did rats fed a single, nutritionally complete diet.
However, Kratz and Levitsky used sucrose powder rather
than a sucrose solution and even the complete diet group did
not overeat very much when offered the sucrose powder. In
another study, rats that were offered a 32% sucrose solution
in addition to separate macronutrient sources displayed a
substantial increase in their total caloric intake [57]. Thus,
diet form is again implicated as an important determinant of
the rat's feeding response to sugar.
Sweetness vs. nutritional balance. Still another factor that
would seem to be important is diet sweetness: presenting the
sugar in the form of a sugar solution provides the animal with
a sweeter tasting food than does mixing powdered sugar into
a dry semisynthetic diet. However, the role of sweet taste in
the rat's response to sugar option diets has been questioned.
In one study, for example, rats offered sugar in the form of
16% or 32% glucose solutions, or 32% or 62% sucrose solu-
tions consumed nearly identical amounts of sugar and total
calories despite the differences in the sweetness of the four
sugar solutions [31]. The authors of this study concluded that
the nutritional consequences of sugar are more important
than sweet taste in determining long-term intake. In particu-
lar, they suggested that the rats selected amounts of sugar
and chow diet to maintain some "'privileged" combination of
protein and carbohydrate. A second study from the same
laboratory led to a similar conclusion [81]. In this experiment
adult male rats were offered either a sweet 32% sucrose
solution or a "'bland" dextrinized starch powder in addition
to their chow diet. Again, despite the taste difference the rats
consumed nearly identical amounts of the two carbohydrate
options as well as of total calories (carbohydrate + chow).
The sucrose group, however, gained more weight than did
CARBOHYDRATE TASTE AND APPETITE
the starch group and chow-only group. Based on these find-
ings the authors concluded that taste is neither a critical
nor a necessary factor in the regulation of carbohydrate in-
take, and they questioned the "conventional wisdom about
the rat's sweet tooth" in determining long-term food intake.
Other investigators have also observed that rats offered the
choice between high and low carbohydrate diets consume
the same amount of carbohydrate irrespective of carbo-
hydrate type (sucrose or starch) [212].
Consistent with the nutritional balance hypothesis, some
studies report that variations in the protein or fat content of
the maintenance diet influence the rat's consumption of a
sucrose solution option [130,144]. However, other findings
indicate that macronutritional balance is not the sole deter-
minant of sugar intake. For example, in a recent study rats
fed a glucose-chow diet (50% glucose powder + 50% chow
powder) consumed as much of a 32% glucose solution as did
rats fed a standard chow diet [163]. As a result the glucose-
chow rats consumed significantly more carbohydrate, and
less protein than did the chow rats (see also [57]). In another
study [33], varying the protein content of the maintenance
diet from 5% to 20% failed to influence the rats' intake of a
32% sucrose solution. Thus, in these experiments the rats
defended their sugar solution intake rather than a particular
balance of macronutrients. This is perhaps not surprising
given the palatability of the sucrose solution and the fact that
adult rats can tolerate rather wide variations in their mac-
ronutrient intakes [97].
Based on the evidence reviewed above, what can be con-
cluded concerning the influence of dietary sugar on food
intake and obesity in the rat? Clearly high-sugar diets do not
always lead to overeating and obesity. Rather the response
to sugar varies as a function of the type and form of diet used
as well as the age, sex, and strain of animal involved. Never-
theless, rats are strongly disposed to overeat and become
obese when offered a sugar solution option. However,
whether it is the sweet taste of sugar or some other property
that is responsible for this effect is open to question. As
discussed next, to answer this question we have examined
the t a r s feeding response to sweet and nonsweet carbo-
hydrates.
POLYSACCHARIDE-INDUCED HYPERPHAGIA AND OBESITY
Polycose solution. The findings of Hill et al. [81] that rats
consumed as much starch as sucrose would appear to pro-
vide strong evidence that sweet taste does not influence
long-term carbohydrate intake. However, the interpretation
of this study is complicated by the fact that the sugar and
starch options differed not only in sweetness but also in
physical form (liquid vs. powder) and molecular structure
(disaccharide vs. polysaccharide). We (Sclafani and Xenakis
[172]) therefore repeated the Hill et al. experiment but with
two modifications: the sucrose and polysaccharide options
were presented in solution form (32% concentration) and
sweet (saccharin-Polycose) as well as " b l a n d " polysac-
charide (Polycose) solutions were used. (Short-term prefer-
ence tests revealed that the rats preferred the sucrose solu-
tion to the saccharin-Polycose solution, and the saccharin-
Polycose solution to the Polycose solution [172].)
As illustrated in Fig. 2, the groups given the Polycose
solutions consumed more saccharide and total calories than
did the sucrose group. The three experimental groups gained
comparable amounts of weight, though, and their weight
gains exceeded that of the chow-fed control group. Thus,
while the Polycose solutions induced overeating and over-
139
,~ ~5o A 200 B
g
i J_
i -- ~ 160
,~ lOO
Q
0
~ so ~ 12o
o
o
o
GROUP: GROUPS
125 I C 140 D
I
"~
~ ~o0 [ v~ too
2o
~ 75 ~ 60
~ 0
~o~
o
GROUPS GROUPS
FIG. 2. Mean (+SE) 30-day body weight gain (A), total daily food
intake (B), saccharide intake (C), and chow intake (D). The four
groups of male rats were fed 0.2%, saccharin + 32% Polycose solu-
tion, 32% Polycose solution, 32% sucrose solution, or no solution
(Control group) in addition to Purina chow and water for 30 days.
Data from Sclafani and Xenakis 1172].
weight, the sucrose solution as in the Hill et al. experiment
[81], induced overweight in the absence of hyperphagia.
Sweetening the Polycose with saccharin did not reliably in-
crease total caloric intake or weight gain, although it in-
creased Polycose intake. This latter finding demonstrates
that sweet taste can influence carbohydrate intake. Never-
theless, the fact that the bland Polycose solution induced as
much overeating and overweight as did the saccharin-
Polycose solution, and increased food intake more than did
the sucrose solution indicated that sweet taste is not a critical
determinant of long-term food intake and body weight.
Polycose powder. The hyperphagia and obesity obtained
with the Polycose solution contrasts with previous reports of
normal food intake and body weight in rats fed
polysaccharide-based diets [1, 15, 81]. However, in previous
experiments the polysaccharides were presented in dry
form, whereas in our study the Polycose was presented as a
solution. To determine if diet form is the critical factor we
conducted a second study [171] which compared the feeding
response of rats to Polycose in solution form or powder
form. The Polycose powder was either mixed into the pow-
dered chow diet or presented as an option to the chow. As
illustrated in Fig. 3, the group given the Polycose solution,
but not the groups given Polycose powder, consumed signif-
icantly more food and gained more weight than did the
chow-fed control group. Note, though, that the Polycose
powder groups deposited more body fat than did the control
group, although less than did the Polycose solution group.
Furthermore, the Polycose intake of the solution group ex-
ceeded that of the Polycose powder groups; the chow intakes
of the three groups were comparable. The findings of this
study clearly demonstrate that diet form is an important
determiT ant of the rat's feeding response to Polycose. Prior
140 SCLAFANI

studies have used only dry polysaccharide diets which ex-

plains why polysaccharide-induced hyperphagia has not 75 A ~ 110
been previously reported.
The results of our Polycose experiments demonstrate that
sugars are not the only carbohydrates that promote hyper-
z_ 50 .~ 90
phagia and obesity. However, rather than explaining why
sugar promotes overeating, the Polycose findings raise new
questions concerning the carbohydrate appetite of the rat.
That is, why do rats consume so much of a Polycose solution
which to the human palate is minimally sweet and not very
palatable; why do they only overeat when the Polycose is
presented in liquid form; why do they consume more Poly-
cose than sucrose but gain equivalent amounts of weight? In GROUPS GROUPS
an attempt to answer these and other questions several ex-
periments were conducted which further investigated the 80 I C 75 D
A
rat's feeding response to polysaccharides and sugars. The
findings of these studies, which are described in detail in the
following papers of this issue, are summarized below. 60 ' -~ 50
,.=

POLYSACCHARIDE TASTE AND APPETITE

Effects of Saccharide Type. Form and Taste [ 157]

The first study in this series was an extensive follow-up to o

our initial two Polycose experiments and compared the ef-

fects of carbohydrate options that differed in saccharide type
(sugar or polysaccharide), form (liquid, powder or gel), and GROUPS GROUP

taste (sweet, bland, bitter) on the food intake, body weight,
and adiposity of female rats. Among its findings, the study
revealed the following: (1) Polycose is not unique in its
hyperphagia-obesity promoting effect. Comparable effects
were obtained with other polysaccharide solutions that dif-
fered from the Polycose solution in degree of hydrolysis,
osmolarity, viscosity, and taste. (2) The polysaccharide so-
lutions induced as much overeating, overweight, and obesity
as did equicaloric glucose and sucrose solutions. The sac-
charides differed, however, in that the rats consumed more
of the polysaccharide solutions than of the sugar solutions.
This confirms our earlier finding that male rats consumed
more of a Polycose solution than of a sucrose solution [ 172].
Note, though, that the sucrose solution induced hyperphagia
in the female rats (present study) but not in the male rats
[172]. (3) As in our previous study [171], Polycose did not
reliably increase food intake or body weight when presented
in powdered rather than solution form, and similar effects
were obtained with glucose and sucrose. (4) Presenting the
Polycose solution as a solid gel rather than as a liquid did not
reduce its hyperphagia-obesity promoting effect. This indi-
cated that it is the water of hydration, not liquidity per se that
is responsible for the hyperphagia inducing effects of carbo-
hydrate solutions. Taken together, the results indicate that
physical form is more important than carbohydrate type in
determining long-term food intake and body weight (but see
below).
(5) The influence of taste on carbohydrate-induced over-
eating and obesity was further examined by comparing the
rats' feeding response to Polycose solutions made sweet with
saccharin or bitter with sucrose octa acetate (SOA). Al-
though saccharin was found not to enhance the
hyperphagia-promoting effect of Polycose in male rats [172],
perhaps it would in female rats since females are reported to
be more responsive than males to sweet taste [203]. The
purpose of the SOA adulteration was to determine if masking
the natural flavor of Polycose with a bitter taste would re-
duce the hyperphagia-promoting effect of Polycose. (Poly-
cose, due to the presence of small amounts of glucose and
maltose, has a mildly sweet taste and perhaps this minimal
FIG. 3. Mean (+SE) 30-day body weight gain (A), total daily food
intake (B), Polycose intake (C), and chow intake (D). The four
groups of female rats were fed 32% Polycose solution (Solution
group) or Polycose powder (Powder group) plus chow and water; or
Polycose powder mixed into the chow diet (Mixed group); or only
chow and water (Control group) for 30 days. Data from Sclafani and
Xenakis [171].
sweetness is sufficient to cause rats to overeat the Polycose
solution.) These taste manipulations were found not to reli-
ably affect the rats" feeding response to Polycose. That is,
the animals consumed comparable amounts of the Polycose,
saccharin-Polycose, and SOA-Polycose solutions, and dis-
played similar increases in total caloric intake, body weight,
and body fat. These results were quite surprising since to the
human palate the Polycose, saccharin-Polycose, and SOA-
Polycose solutions differed greatly in palatability. However,
perhaps to rats saccharin and SOA have little effect on the
palatability of a Polycose solution, particularly when the
solution is available 24 hr/day. This possibility was tested in
the next study.
Effects of Taste and Form on Polycose Preference [1691
The first part of this study determined whether saccharin
or SOA adulteration affects the rat's long-term preference
for 32% Polycose solutions. This was accomplished by giving
rats 24-hr/day two-solution preference tests; chow was
available ad lib. The animals significantly preferred the
saccharin-Polycose solution to the plain Polycose solution,
the plain Polycose solution to the SOA-Polycose solution,
and strongly preferred the saccharin-Polycose solution to the
SOA-Polycose solution. Yet, in confirmation of the previous
study 1157], rats given only the SOA-Polycose solution, in
addition to chow and water, consumed as much solution as
did rats given the Polycose or saccharin-Polycose solution.
Therefore, while the addition of saccharin or SOA to a Poly-
cose solution does alter its palatability this alteration does
not affect Polycose intake in the no-choice condition. (Note
that quinine adulteration significantly reduces Polycose
CARBOHYDRATE TASTE AND APPETITE
solution intake [169] but we avoided the use of quinine be-
cause of its possible toxic postingestive effects [3].)
In the second part of this study we examined the role of
palatability in the rat's differential feeding response to sac-
charide solutions and powders. In both short-term (30-
rain/day) and long-term (24-hr/day) two-choice tests the rats
preferred a 32% Polycose solution to pure Polycose powder
and, to a lesser extent, SOA-Polycose solution to Polycose
powder. Based on these results it could be argued that rats
consume more saccharide solution than powder because the
solution is more palatable. This interpretation is not sup-
ported, however, by the results obtained with a third group
of rats. This group was given the choice between the SOA-
Polycose solution and sucrose powder. (This particular
choice was tested because in the previous study [157] the
rats fed the SOA-Polycose solution consumed more sac-
charide than did the rats fed the sucrose powder.) The rats in
this third group displayed a strong preference for the sucrose
powder over the SOA-Polycose solution during the short-
term test and the initial days of the 24-hr/day test. However,
with continued testing (24 hr/day) the rats switched their
preference and consumed more SOA-Polycose solution than
sucrose powder. These results clearly indicate that rats do
not initially prefer the flavor of the bitter SOA-Polycose
solution to the sweet sucrose powder. Thus, the fact that rats
consume more SOA-Polycose solution than sucrose powder
in long-term tests would seem to be due to the postingestive
rather than the orosensory properties of the carbohydrate
sources. The nature of the postingestive effects involved re-
mains to be identified, but it may be related to differences in
the rate of gastric emptying and/or intestinal absorption of
carbohydrate in liquid and powder form (see below).
Origin of Polycose Appetite [167]
At this point in our research it appeared that postingestive
nutritional factors far outweighed taste factors in determin-
ing long-term carbohydrate intake. This was most evident in
the case of the SOA-Polycose solution in which the data
suggested that rats consumed the solution for its nutritional
rather than its taste properties. We evaluated this interpreta-
tion by measuring the rat's sham-feeding response to SOA-
Polycose and Polycose solutions. With postingestive factors
minimized with a gastric fistula it was predicted that the rat
would consume little of these solutions.
In the first experiment rats were first trained to sham-feed
a 32% glucose solution and were then tested with a 32%
Polycose solution adulterated with SOA. As expected, they
sham-fed significantly less of the SOA-Polycose solution
than of the glucose solution. However, their sham-intake of
the SOA-Polycose solution was substantial (45 ml/hr) which,
given the gastric fistula, could not be attributed to postinges-
tive factors. Additional tests indicated that it was not the
small amounts of glucose and maltose in the SOA-Polycose
solution, nor the solution's viscosity, and certainly not its
bitter taste that caused the animals to sham-feed the SOA-
Polycose solution. Thus, it appeared that the rats were at-
tracted to the taste of the polysaccharides in the solution.
Consistent with this interpretation, in a second experiment
rats were observed to sham-feed as much of an unadulter-
ated 32% Polycose solution as they did of a 32% sucrose
solution. These findings, while they do not argue against the
importance of postingestive factors in the long-term control
of polysaccharide intake, seriously questioned the assump-
tion that polysaccharides are tasteless to rats. We therefore
conducted several additional experiments to further assess
the rat's orosensory response to polysaccharides.
141
Hedonic Response to Polyeose and Sugar Solutions [159]
In this study we compared the rat's hedonic response to
solutions of Polycose, glucose, maltose, and sucrose using
short-term, one- and two-solution intake tests. Maltose was
of particular interest because it is a highly preferred sugar,
and, along with glucose, is a constituent of Polycose. The
rats were neither food nor water deprived and thus were
presumably drinking the saccharide solutions primarily for
taste.
In the one-bottle tests the rats consumed more Polycose
than any of the sugars at low molar concentrations (0.0125
and 0.025 M); at higher concentrations (0.2 or 0.4 M) sac-
charide intakes did not differ. In the two-solution tests the
rats preferred Polycose to sucrose at low molar concentra-
tions but sucrose to Polycose at high concentrations; they
preferred Polycose to maltose at all concentrations. (Poly-
cose vs. glucose preferences were not tested but in an un-
published study Polycose was preferred to glucose at low to
high molar concentrations.) Thus, rather than being bland
tasting the present results indicate that rats find polysac-
charide solutions to be more palatable than maltose, sucrose,
and glucose solutions at low concentrations, and second only
to sucrose solutions at high concentrations.
Poly('ose and Sugar Taste Pr~:/?rence Thresholds [166]
Since t.he preceding study indicated that rats are strongly
attracted to Polycose solutions at low concentrations, we
next compared their preference threshold for Polycose with
that of sucrose and maltose using the 24-hr two-bottle
method of Richter. The preference threshold for Polycose
(0.0001 M) was found to be 25 to 26 times lower, on a molar
basis, than the preference thresholds for maltose (0.0025 M)
and sucrose (0.0026 M). Thus, not only do rats like the taste
of polysaccharides, they can detect polysaccharides at rather
low concentrations. The rat's low taste threshold for
polysaccharides would appear to be an adaptive response to
the low solubility of the naturally occurring polysaccharides.
Qualitative Differences in Polycose and Sugar Tastes [133]
In this study we asked the question what does Polycose
taste like to rats? Does it have a sweet, i.e., sucrose-like
taste, or does it have a qualitatively different taste? In par-
ticular, does Polycose taste like maltose? (There were rea-
sons to suspect that both Polycose and maltose share a non-
sucrose like taste (see [133].) This question was addressed by
using a conditioned taste aversion paradigm. Rats were
conditioned to avoid one saccharide by pairing its ingestion
with visceral malaise, and the degree to which they general-
ized this aversion to other saccharides was assessed.
The results showed that rats generalized between Poly-
cose and maltose taste aversions but showed little gener-
alization between Polycose-maltose aversions and a sucrose
aversion. Thus, rats appear to taste Polycose as similar to
maltose but different from sucrose. Additional tests revealed
that only the sucrose aversion generalized to fructose or sac-
charin, while both Polycose and sucrose aversions general-
ized to glucose. Based on these and other findings we hy-
pothesized that rats have two types of carbohydrate taste
receptors, one being a sucrose-type or "sweet" receptor,
and the other being a receptor for polysaccharides. We
further speculated that maltose, since it has a molecular
configuration similar to that of polysaccharides, stimulates
the rat's polysaccharide taste receptor and this accounts for
 142
the palatability of maltose. Glucose may stimulate both re-
ceptor types because glucose is a component of sucrose and
polysaccharides.
h~lTttence of Saceharide Length on Polysaccharide Appetite
[/621
Polycose contains saccharides of varying lengths and thus
it was uncertain as to what the optimal stimulus is for the
rat's postulated polysaccharide taste receptor. To provide
some information concerning this question we examined the
rat's hedonic response to saccharides of different chain
lengths using brief one- and two-solution intake tests. The
results revealed that as saccharide length increased from one
(glucose) to two (maltose) to three (maltotriose) to 4--8 (mal-
tooligosaccharides) glucose units the rat's preference for the
solution increased, but with longer saccharides (mal-
topolysaccharides of 20 or more glucose units) their solution
preference declined somewhat. Thus, of the saccharide so-
lutions tested, the most preferred was one that contained
maltooligosaccharides of 4 to 8 glucose units in length. The
rats also preferred the maltooligosaccharide solution to a
Polycose solution which is consistent with the fact that 60%
of the saccharides in Polycose are smaller than four or larger
than eight glucose units in length. Our previous findings ob-
tained with Polycose, therefore, underestimate the strength
of the rat's polysaccharide, or more precisely, mal-
tooligosaccharide appetite. (Note that because of the limited
availability of pure maltooligosaccharide preparations we
have continued to use Polycose in our research.)
E/.fects of Gustatoo' Det/ferentution on Sac('haride Appetite
[2021
Our behavioral findings indicated that rats taste polysac-
charides and taste them as different from sucrose, but there
is no published data on the response of the rat's gustatory
nerves to polysaccharides (such studies are now in prog-
ress). Based on indirect evidence (see [202]), however, it
seemed likely that different gustatory nerves mediate the
rat's appetite for polysaccharide and sucrose. This
possibility was explored in the present study by comparing
the effects of selective gustatory nerve transections on the
intake of sucrose and Polycose solutions in short-term tests.
The results of this study revealed that selective transec-
tions of the glossopharyngeal nerve, superficial petrosal
nerve, and the pharyngeal branch of the vagus nerve differ-
entially altered the intake of sucrose and Polycose solutions.
On the other hand, selective transection of the chorda tym-
pani nerve (CTN) produced comparable effects on sucrose
and Polycose intake. In addition to the groups given selected
nerve transections, one group of rats received combined
bilateral transections of all four gustatory nerves. These
animals displayed comparable reductions in Polycose and
sucrose intake. Taken together, the results indicate that
polysaccharide and sucrose appetites are mediated to the
same degree by the gustatory system, although the gustatory
pathways involved appear to differ. This latter finding is
consistent with the hypothesis that rats have different taste
receptors for sucrose and polysaccharides.
Effects oJ'Anosmia on Polycose Appetite [2011
The near-total gustatory denervation produced in the
previous study did not completely block the rat's Polycose
intake. Taste fibers spared by the surgery may have been
SCLAFANI
responsible for the animal's residual appetite but it is also
possible that other orosensory systems are involved in
polysaccharide appetite. The olfactory system appears to
have little if any influence on the rat's ingestive response to
sucrose [198], but its role in polysaccharide appetite is uncer-
tain. We therefore investigated whether anosmia alters the
rat's ingestive response to Polycose solutions. Animals
treated with intranasal zinc sulphate, which produces tempo-
rary anosmia, were found not to differ from control rats in
their Polycose solution intake. Thus, olfaction is not a major
factor in the rat's appetite for polysaccharide solutions.
However, the present findings do not exclude a role for ol-
faction in the animal's response to "'real" foods as opposed
to pure carbohydrate solutions. The possible involvement of
trigeminal sensory system in polysaccharide appetite re-
mains to be investigated (see [14]).
Sham-Feeding Response to Polycose and Sucrose [134]
In this experiment we further investigated the sham-
feeding response of rats to Polycose and sucrose solutions.
In our initial sham-feeding study [167] rats sham-fed com-
parable amounts of 32% Polycose and 32% sucrose solutions.
Yet, other findings indicated that rats prefer sucrose to Poly-
cose at this concentration [161,172]. This apparent discrep-
ancy was resolved in the present study: in two-solution,
sham-feeding tests the rats strongly preferred the 32% su-
crose solution to the 32% Polycose solution, although in
one-bottle tests they sham-fed similar amounts of the two
solutions. The failure to obtain a difference in Polycose and
sucrose solution intakes in the one-bottle tests was attributed
to a ceiling effect, i.e., the rats were sham-feeding at near-
maximum rates throughout the 30-min test periods.
The sham-feeding response to less concentrated solutions
was also measured and an unexpected result emerged from
these tests. That is, whereas in one-solution tests the rats
sham-fed significantly more 4% Polycose than 4% sucrose, in
the two-solution test they preferred the 4% sucrose to the 4%
Polycose. Analysis of the rat's drinking pattern during the
one-bottle tests suggested that the animals satiated to the
taste of sucrose faster than to the taste of the Polycose. (This
satiety effect was presumably orosensory in origin since the
gastric fistula eliminated postingestive satiety (see [127].)
The mechanism responsible for the differential "oral sati-
ety" to sucrose and Polycose remains to be determined, but
the results are compatible with the view that different gus-
tatory systems mediate the taste and appetite for these
carbohydrates.
Saccharin-Carbohydrate Taste Interactions [160]
Rats consume enormous amounts of solutions containing
glucose (3%) and saccharin (0. I-0.2%) [195]. This polydipsic
response has been attributed to the synergistic taste effects
of the glucose + saccharin (G+S) mixture (see [179]). In this
study we explored the possibility that the rat's polysac-
charide taste may be involved in the overconsumption of
G + S solutions. Our previous findings [133] suggest that glu-
cose stimulates both polysaccharide and sucrose taste recep-
tors whereas saccharin stimulates only sucrose receptors,
and perhaps the combined stimulation of both receptor types
by the G + S mixture contributes to the polydipsia.
The results showed that a sucrose + saccharin solution,
which presumably stimulates only sucrose type receptors,
induced as much polydipsia as did a Polycose + saccharin
solution, which presumably stimulates both receptor types.
CARBOHYDRATE TASTE AND APPETITE
Based on these findings we concluded that the combined
activation of sucrose-type and polysaccharide-type taste re-
ceptors is a sufficient but not a necessary condition to obtain
taste-induced polydipsia.
Species D(fferences in Saccharide Taste Preferences [60]
In this study we turned our attention to the species speci-
ficity of polysaccharide taste. Informal taste tests indicated
that Polycose is minimally sweet and not very palatable to
humans and this was confirmed in a formal experiment.
Human subjects rated Polycose solutions to be less sweet
and less palatable than equimolar solutions of sucrose or
maltose solutions. In fact, at the higher concentrations, the
subjects reported the Polycose solutions to be unpleasant.
Thus humans and rats respond quite differently to polysac-
charide solutions.
To determine if other species share the rat's appetite for
polysaccharides a second experiment compared the taste
preferences of rats, hamsters, gerbils, and spiny mice using
24-hr/day two-bottle tests (solution vs. water). In general,
the rats displayed stronger preferences for Polycose and mal-
tose (over water) than did the other three species. The spe-
cies did not differ in their sucrose preferences except that the
spiny mice preferred sucrose less than did the other three
species. Within-species comparisons revealed that for all the
rodents the preference for Polycose was as strong or
stronger than their preference for sucrose or maltose. There-
fore, while less responsive than the laboratory rat, the gerbil,
hamster, and spiny mouse all share with the rat an appetite
for polysaccharides. In contrast with these rodent species, in
another study [79] we observed that cats failed to prefer
either Polycose or sucrose solutions to water. These species
differences in carbohydrate taste preference are presumably
related to the natural dietary habits of the animals, but this
relationship requires much further study.
Sex Differences in Saccharide Taste Preferences [161]
In the preceding study both male and female rodents were
tested, but few sex differences in taste preference were ob-
tained. This was surprising since previous studies indicate
that male and female rodents differ in their sweet taste pref-
erence [203]. In this study, therefore, we investigated the
taste preferences of male and female rats in greater detail.
While 24-hr/day two-bottle (solution vs. water) tests failed to
reveal sex differences, 3-rain and 30-rain/day Polycose vs.
sucrose tests indicated that male rats have a greater prefer-
ence for Polycose, and female rats a greater preference for
sucrose. Somewhat different results were obtained with 24-
hr/day Polycose vs. sucrose tests. In this case, both sexes
initially displayed a preference for 32% sucrose over 32%
Polycose, but then shifted their preference over days to the
Polycose solution. The female rats, though, displayed a more
pronounced preference shift than did the male rats. The
cause of the preference reversal appears to be related to the
postingestive rather than the orosensory properties of the
Polycose and sucrose solutions. Taken together, the results
indicate that there are differences in the carbohydrate taste
preferences of male and female rats as well as in their re-
sponse to the postingestive effects of carbohydrates.
Starch Preference in Rats [168]
The presumed function of the polysaccharide taste sen-
sitivity revealed in our studies is to allow rodents to readily
143
identify starch-rich foods. Consistent with this interpreta-
tion, in this study rats displayed an immediate preference for
powdered corn starch over powdered cellulose in short-
term, two-choice tests. The rats also preferred the corn
starch to Polycose powder. This latter finding was unex-
pected. We had assumed that because Polycose contains
smaller and more soluble polysaccharides than does corn
starch, Polycose would be a better stimulus for the rat's
polysaccharide taste receptors than corn starch.
Since corn starch consists mostly of branched-chain
polysaccharides (amylopectin) whereas Polycose contains
predominantly straight-chain polysacchardes we determined
the rat's preference for these two types of polysacchardes.
In short-term tests using powdered saccharides the rats reli-
ably preferred amylopectin (branched chain) starch to
amylose (straight-chain) starch as well as to Polycose. How-
ever, when tested with the saccharides in a hydrated form
(solutions, suspensions or gels) the rats strongly preferred
Polycose to amylopectin starch. These latter findings
suggested that the preferences observed with the powdered
saccharides was due to nongustatory factors, i.e., texture or
odor. The results of additional tests, though, failed to iden-
tify these factors. That is, texture, as manipulated by particle
size, was found not to influence the rat's preferences for
powdered saccharides. Also, the noncarbohydrate con-
stituents of corn starch, which give it its "cereal" flavor,
were found not to be a factor in the rat's preference for
amylopectin powder over Polycose powder. Thus, the
orosensory determinants of starch preference remain to be
fully identified.
The present study also examined the rat's long-term pref-
erence for starch and Polycose powder. Despite their strong
preference for powdered starch over powdered Polycose in
short-term tests, the rats quickly reversed this preference
during 24-hr/day tests and consumed more Polycose than
starch. In a subsequent experiment (Sclafani, Vigorito and
Pfeiffer, in preparation) we observed that rats consumed
more Polycose than amylopectin in long-term tests whether
the saccharides were presented in powder or gel form. Fur-
thermore, the Polycose gel produced greater increases in
food intake and body weight than did the amylopectin gel.
This latter finding indicates that food intake is influenced by
saccharide type as well as form. Rats may switch their pref-
erence from amylopectin to Polycose and consume more
Polycose than starch in long-term tests because the Polycose
is digested and absorbed at a faster rate than is starch.
INTEGRATION
The findings reviewed above and detailed in the subse-
quent papers in this issue provide a considerable amount of
new information concerning carbohydrate taste and appetite
in animals. These new findings are integrated with previous
research in the following "working" models of carbohydrate
taste and appetite in the rat.
Carbohy~h'ate Taste
With respect to carbohydrate taste, Fig. 4 presents a
two-carbohydrate taste model that postulates that rats have
two types of taste receptors for carbohydrates: a sucrose-
type or "sweet" receptor and a polysaccharide-type recep-
tor. The sucrose-type receptor is thought to consist of glu-
cose and fructose subsites [8], and is responsive to sucrose,
fructose, glucose, and maltose, in that order, as well as to
saccharin. The polysaccharide-type receptor is postulated to
144 SCLAFANI

Two C a r b o h y d r a t e
T a s t e Model

Polysaccharide Receptor Sucrose R ecept or

-V-1V
- = -
Glucose Subslte ~ = Fructose Subsite

R e c e p t o r S t i m u l a t e d bY: Receptor S t i m u l a t e d bF:

N a l t o o l Igosaccharldes Sucrose
Maltopolysacchar Ides Fructose
Maltose Glucose
Glucose Maltose
Saccharin

FIG. 4. Schematic representation of the two carbohydrate taste model. The model
postulates that rats have a polysaccharide taste receptor that consists of several
glucose subsites and that is stimulated by maltooligosaccharides, maltopolysac-
charides, maltose, and glucose. The sucrose receptor consists of a glucose subsite
and a fructose subsite and is stimulated by sucrose, fructose, glucose, maltose,
and saccharin.

consist of several glucose subsites and to be responsive to
maltooligosaccharides, maltopolysaccharides, maltose, and
glucose, in that order. The model further postulates that the
two receptors activate separate gustatory systems (or sub-
systems) such that rats perceive the taste of sucrose and
polysaccharides as qualitatively different. This two-taste
model is based on the findings that (a) rats display an avid
appetite for Polycose and, in particular, for maltooligosac-
charide solutions; (b) the relative preference for Polycose
and sucrose varies as a function of concentration, i.e., Poly-
cose is preferred at low concentrations, sucrose is preferred
at high concentrations; (c) the preference threshold for Poly-
cose is much lower than that for sucrose; (d) conditioned
taste aversions to sucrose and Polycose do not cross-
generalize; (e) Polycose and sucrose differ in their "'oral"
satiety effects; and (f) Polycose and sucrose intake are dif-
ferentially affected by selective gustatory deafferentation.
Electrophysiological evidence. The above findings are
based on behavioral studies and obviously electrophysiolog-
ical data are needed to fully evaluate the two-carbohydrate
taste model. Preliminary results obtained with whole nerve
recordings indicate that the rat's chorda tympani nerve is
equally responsive to Polycose and sucrose, whereas the
greater superficial nerve is more responsive to sucrose than
to Polycose (Beidler and Nejad, unpublished observations).
In contrast to the rat, in the gerbil initial results indicate that
the corda tympani nerve is less responsive to Polycose than
to s,fcrose (Beider and Nejad, unpublished observations;
Jakinovich, unpublished observations). These observations
are compatible with our findings that selective deafferenta-
tion of the chorda tympani and superficial nerves differen-
tially affect Polycose and sucrose intake, and that rats and
gerbils differ in the strength of their preference for Polycose.
Thus, although further electrophysiological evidence is re-
quired, particularly single fiber data, the preliminary results
provide direct evidence that the gustatory system is respon-
sive to polysaccharides.
Maltose receptors? It could be argued that since the rat
appears to have multiple sugar receptor subtypes (see [9])
their polysaccharide taste sensitivity may be mediated by
one of these sugar receptor subtypes, in particular by a
"'maltose" receptor, rather than by a polysaccharide recep-
tor. After all, maltose-sensitive gustatory single fibers have
been identified [8], and the conditioned taste aversion data
indicate that maltose and Polycose produce a similar taste
sensation [133]. Furthermore, although Polycose contains
only small amounts of free maltose, hydrolysis of Polycose
in the mouth by salivary amylase would increase its maltose
content. According to this argument, though, rats should
prefer maltose solutions to Polycose solutions, and their mal-
tose preference threshold should be lower than their Poly-
cose preference threshold but this is not the case. Note in
particular, that at the Polycose preference threshold even if
all the Polycose in solution was converted to maltose the
concentration would be below the rat's maltose preference
threshold. In fact, given the rapidity of the rat's drinking
response probably only a small amount of Polycose in solu-
tion is hydrolyzed in the mouth. Empirical evidence that
salivary amylase plays little role in Polycose appetite is pro-
vided by the findings that inhibiting salivation with atropine
methyl nitrate does not reduce Polycose solution intake
(Sclafani and Nissenbaum, unpublished observations). Fi-
nally, preliminary observations indicate that the rat's chorda
tympani nerve is equally responsive to Polycose and sucrose
(Beidler and Nejad, unpublished observations) whereas
previous reports indicate that the CTN is less responsive to
maltose than to sucrose [140]. Thus, while further elec-
trophysiological evidence is required, the available data do
not support the idea that maltose taste receptors mediate
polysaccharide appetite in the rat. Rather it may be that the
rat's responsivity to maltose is mediated by polysaccharide
receptors.
Salivary amylase, while it may not be involved in the taste
response to polysaccharide solutions, may play an important
role in the taste response to naturally occurring polysac-
charides [150]. That is, as the rat masticates starch foods
sufficient amounts of soluble maltopolysaccharides and mal-
tooligosaccharides may be produced by the action of salivary
C A R B O H Y D R A T E TASTE AND APPETITE
Multifactor
C a r b o h y d r a t e Appetite Model
DIET AVAILABILITY
FLAVOR ~ be produced by a food low in sugar. Thus, rats may prefer
Taste
Texture~ ) APPETITE ) HYPERPHAGIA ~ OBESITY
Smell /
POSTINGESTIVE FACTORS
Rate of Absorption
Osmolarlty
Nutrient Interactions
Others
FIG. 5. Multifactor model of carbohydrate appetite. See text for
explanation.
amylase to stimulate the animal's polysaccharide taste re-
ceptors. The fact that the rat's preference threshold for
polysaccharides is very low [166] is consistent with this in-
terpretation. Thus, salivary amylase, in addition to its role in
the digestion of carbohydrates in the rat's stomach [108],
may facilitate the gustatory detection of carbohydrates. In
view of the species differences in polysaccharide taste pref-
erences it is interesting to note that rat salivary amylase is
100 times more active than is human salivary amylase [108],
and that many mammalian species lack salivary amylase
[150].
Ftmctional significance. If, according to the two-
carbohydrate taste model, rats can taste polysaccharides as
well as sugars, and taste them as qualitatively different from
sugars (i.e., sucrose and fructose), what is the functional
significance of this dual taste sensitivity? It has long been
assumed that sweet taste sensitivity is adaptive because it
allows animals to readily identify sugar-rich foods. Similarly,
polysaccharide taste may be adaptive because it allows
animals to readily identify starch-rich foods. Less obvious is
why polysaccharides and sugars should produce different
taste sensations. Only speculations can be offered at this
point. Perhaps having two rather than one carbohydrate
taste improves the animal's ability to adjust its food selection
according to its nutritional needs and variations in food
availability and abundance. For example, animals in need of
immediate energy may preferentially select sweet foods since
sugar is more rapidly digested than is starch. This may ex-
plain the interesting finding [185] that insulin injections in-
crease glucose and sucrose intake, but not maltose intake in
mice--assuming that maltose has a polysaccharide-like taste
to mice. On the other hand, animals in need of protein may
preferentially select polysaccharide-tasting foods since
starch-rich plants (e.g., vegetables) have a higher protein
content than do sugar-rich plants (fruits). Previous studies in
fact demonstrate that pregnant rats display a reduced sweet
taste preference, as measured by two-bottle preference tests,
and this has been attributed to their increased protein re-
quirement [203]. However, whether pregnancy or other
states of increased protein need alter polysaccharide taste
preferences remains to be determined.
Another interesting question is why should rats prefer
Polycose solutions to sucrose solutions at low concentra-
145
tions, but sucrose to Polycose at high concentrations [134,
159, 161]? Perhaps this preference behavior represents an
adaptation related to the different solubilities of starch and
sugar. That is, since starch is poorly soluble, a starch-rich
food would produce a " w e a k " polysaccharide taste. In con-
trast, sugar is very soluble and a " w e a k " sugar taste would
Polycose to sucrose at low concentrations because the Poly-
cose taste signifies a high-starch (and high-calorie) food
whereas the sucrose taste signifies a low-sugar (and low-
calorie) food. At high concentrations, on the other hand, the
sucrose taste would signify a sugar-rich food which rats
may ordinarily prefer to a starch-rich food. Recall, however
that with experience with concentrated solutions rats alter
their preferences for sucrose and Polycose (see below).
Carbohydrate Appetite
Flavor. The above discussion has focused on taste but
clearly other factors contribute to carbohydrate appetite.
This is illustrated in Fig. 5 which presents a multifactor
model of carbohydrate appetite. As the model indicates,
taste is only one component, albeit the most extensively
studied one, of flavor. Recent studies demonstrate that the
orosomatosensory system plays an important role in the con-
trol of food intake and macronutrient selection [221]. Of par-
ticular interest is the finding that trigeminal deafferentation
reduces the rat's hedonic response to sucrose solutions [14].
Note that food technologists have long recognized that sugar
adds texture ( " b o d y " or "mouth feel") as well as sweet
taste to foods and fluids [131]. Mouth feel is probably a fac-
tor in the preference rats display for carbohydrate liquids
and gels over carbohydrate powders in short-term tests
[ 168,169]. However, whether orosomatosensory stimuli con-
tribute to the rat's differential hedonic response to sugars
and polysaccharides is less certain. Olfactory stimuli do not
appear to be involved in the rat's response to carbohydrates
in pure form [168, 198,201], but olfaction presumably plays a
crucial role in the location and identification of carbo-
hydrate-rich foods in the animal's natural environment. Fur-
thermore, olfactory cues are important in the conditioning of
food preferences [111] (see below). Much remains to be
learned about the contribution of the different orosensory
systems to carbohydrate appetite and preference. In particu-
lar, as previously mentioned, the sensory basis of the rat's
preference for starch powders is not fully understood (see
[ 1681).
Cephalic phase responses. Carbohydrate taste (flavor), in
addition to directly stimulating appetite via central nervous
system mechanisms, may have an indirect influence through
the activation of cephalic-phase digestive responses
mediated by the autonomic nervous system. It is well estab-
lished that the taste, texture, and smell of food can elicit
reflexive changes in pancreatic and gastrointestinal secre-
tions and motility which facilitate the postingestive disposi-
tion of food [21,141]. Some investigators hypothesize that
these cephalic reflexes, the cephalic insulin response (CIR)
in particular, may also modulate the animal's hedonic and
ingestive responses to food [114,115]. It has been reported,
for example, that the size of the CIR is related to diet
palatability, and that manipulations that block this cephalic
reflex (e.g., vagotomy) abolish the influence of diet palata-
bility on food intake [114,115]. Sweet taste is a potent
stimulus of the CIR [141] and therefore this may be one way
146
in which sweet taste influences carbohydrate appetite and
intake.
The relationship between sweet taste, the CIR, and sugar
intake remains controversial, however. For example, Grill et
al. [71] reported that of several sweeteners tested, including
seven sugars and saccharin, only glucose produced a reliable
CIR in rats. In contrast, other investigators obtained a CIR
to saccharin although there was considerable interanimal
variability in the size of the response [141]. Furthermore,
manipulations that block the CIR do not consistently inhibit
sugar intake. For example, atropine methyl nitrate injections
reduce sucrose intake in sham-feeding tests but not in real-
feeding tests [132, 170, 207]. Also, while subdiaphragmatic
vagotomy reduces the intake of sucrose solutions it does not
suppress the intake of sucrose powder [ 164,200]. In addition,
gastric vagotomy, which is reported to spare the CIR, in-
hibits sucrose solution intake to the same extent as does
subdiaphragmatic wlgotomy [200]. Thus, no firm conclu-
sions can be reached concerning the role of cephalic reflexes
in the rat's appetite for sugar. Even less is known about the
possible involvement of cephalic reflexes in starch and Poly-
cose appetite.
Postingestive inflttences. In addition to orosensory fac-
tors, postingestive factors have a major influence on carbo-
hydrate appetite and intake. Feeding inhibitory effects, i.e.,
postingestive satiety, are the most extensively documented
and these limit the amount of carbohydrate animals can con-
sume [87]. Thus, variations in carbohydrate taste (or flavor)
may not affect intake because of the limits set by postinges-
live satiety. On the other hand, variations in the form or type
of carbohydrate may alter intake because they affect the
postingestive disposition of the carbohydrate [186]. For
example, rats may consume more of carbohydrate solutions
or gels than of carbohydrate powders [157] because carbo-
hydrates in hyd,ated forms are more rapidly digested and
absorbed than are carbohydrates in a dehydrated form. Simi-
larly, rats may consume more Polycose than corn starch in
long-term tests [168] because the Polycose is more rapidly
digested than is the corn starch, it is now clear that a number
of factors affect the digestibility and absorption of carbo-
hydrates [94,1471.
Pret'erence reversals. The postingestive actions of carbo-
hydrates not only have an inhibitory effect on food intake but
they can also modulate food preferences and, under certain
conditions, actually stimulate food intake. The reversals in
saccharide preference observed in our studies and in previ-
ous experiments appea. to be due to postingestive actions of
the carbohydrates. Recall that in long-term tests rats reverse
their initial preference for starch powder over Polycose
powder [ 168], 329,; sucrose solution over 32% Polycose solu-
tion [161], 32CA glucose solution over 32% Polycose (unpub-
lished findings), sucrose powder over SOA-Polycose solu-
tion [169], and concentrated glucose solutions over dilute
glucose solutions [19,1371. As previously mentioned, the
preference reversal obtained with concentrated and dilute
glucose solutions has been attributed to the hypertonicity of
the concentrated solutions [19]. Osmotic factors may also
contribute to the preference reversals obtained with Poly-
cose, sucrose, and glucose solutions since the osmolarity of
Polycose is less than that of the sucrose and glucose at
isocaloric concentrations 1152]. It is less certain that osmo-
larity contributes to the preference reversal obtained with
sucrose powder and SOA-Polycose solution, and starch and
Polycose powders. In these cases rate of gastric emptying
and/or intestinal absorption may be the critical factor. That
SCLAFANI
is, rats may develop a preference for the saccharide that is
more rapidly absorbed.
Conditio:u,d prefi'rences. Other studies report that rats
develop preferences for flavors that are added to polysac-
charide solutions [19,121]. In these studies it was assumed
that rats do not taste polysaccharides and that postingestive
stimuli were responsible for the flavor conditioning. How-
ever, the discovery that rats have a very sensitive polysac-
charide taste complicates the interpretation of these studies.
That is, it is possible that the taste of the polysaccharides in
addition to their postingestive actions may have served as
the unconditioned stimulus in the conditioning of the flavor
preference (see [58]).
The most direct evidence for the postingestive modula-
tion of appetite is provided by experiments in which flavor
preferences were conditioned by carbohydrate loads that
bypassed the oral cavity. Although negative results have
been obtained [53, 104, 142, 148], intragastric and intrave-
nous infusions of glucose have been found to condition
flavor preferences [I 19. 176, 191]. (The failure to obtain pos-
itive conditioning in some experiments may have been due to
the use of hypertonic glucose solutions which have aversive
consequences.) We have recently observed that rats also
rapidly develop preferences for flavors associated with
intragastric infusions of Polycose (Nissenbaum and Sclafani,
unpublished findings). We further observed that retarding
the rate of digestion and absorption of the Polycose with the
glucosidase inhibitor acarbose [46] blocked flavor condition-
ing. Much more remains to be learned about the physiolog-
ical mechanisms involved in the postingestive conditioning
of food preferences although recent reports indicate that
both preabsorptive (i.e., gastric) and postabsorptive (i.e..
hepatic) nutrient "'sensors'" are involved [52,191].
The evidence that the postingestive effects of carbo-
hydrates can condition flavor preferences, or in some cases
aversions [18], raises the question as to the degree to which
the rat's hedonic response to carbohydrate taste is learned or
unlearned. As already noted, behavioral findings indicate
that rats have an innate preference for sweet taste [75,92] but
whether they also have an innate preference for polysac-
charide taste remains to be determined. Even if rats have
innate preferences for sweet and polysaccharide tastes their
preferences can be modulated by the postingestive feedback
provided by the saccharides. Nevertheless, there may be
limits to the modifiability of carbohydrate taste preferences
by postingestive nutritive feedback. This is suggested by
preliminary results obtained in a study in which rats were
trained to sham-feed one flavored Polycose solution and
really-feed a different flavored Polycose solution (Nissen-
baum and Sclafani, unpublished findings). With 8% Polycose
solutions the rats developed a preference for the flavor that
they really fed (and that was associated with nutritive ef-
fects) over the flavor that they sham-fed, but with 32% Poly-
cose solutions the rats failed to display such a preference
(see also [197]). Thus a strong carbohydrate taste may limit
the ability of postingestive nutritional cues to modify food
preferences. Another issue concerns whether or not rats in-
nately associate sweet and/or polysaccharide taste with the
specific nutritive effects of carbohydrates. Booth [18] has
argued that in this respect carbohydrate appetite (or hunger
to use his term) is learned. For example, Booth maintains
that the sweet taste preference elicited by insulin-induced
hypoglycemia is a conditioned response.
Feeding stimuhm,3' effect. In addition to their effects on
food preferences, the postingestive actions of carbohydrates
C A R B O H Y D R A T E TASTE AND APPETITE
may directly stimulate food intake under some conditions.
This is indicated by the findings that intragastric,
intraduodenal, or hepatic-portal infusions of sugar solutions
increase the amount of chow consumed by rabbits in their
next meal [63]. Note that the feeding stimulatory effect was
dependent upon the rate of infusion; at a high rate of infusion
(3 ml/min) subsequent food intake was increased, but at a
low infusion rate (1 ml/min) food intake was decreased. The
feeding stimulation obtained at the high infusion rate was
hypothesized to be a consequence of the hyperinsulinemia
and subsequent events triggered by the rapid rise in blood
glucose [63,64].
Rate ~/'absorption. The findings obtained with low and
high infusion rates compliment other results cited in this
paper that implicate an important role of absorption rate in
the feeding effects of carbohydrates (see also [186]). This is
further indicated by the results of a study in which the
glucosidase inhibitor acarbose [46] was used to slow down
the rate of digestion and absorption of Polycose (Sclafani and
Mann, unpublished observations). The rats in this experi-
ment were fed, in addition to chow and water, a 32% Poly-
cose solution to which was added 0, 20, 40, or 60 mg/100 ml
of acarbose. The acarbose produced a dose dependent de-
crease in absolute and percent Polycose intake, as well as in
total caloric intake. (Note that Vasselli et al. [199] reported
that acarbose did not reduce 32% sucrose intake in rats, but
in their study the acarbose was added to the chow diet rather
than the sucrose solution.)
Variations in carbohydrate absorption may affect food
intake via several different mechanisms. The rapid rise in
blood glucose is thought to trigger a sequence of hormonal
and metabolic events that promotes lipogenesis and overeat-
ing [63,64]. Also, rats may develop preferences for carbo-
hydrate foods that are rapidly absorbed (Nissenbaum and
Sclafani, unpublished observations; [191]). On the other
hand, slowly absorbed carbohydrates ("lente carbo-
hydrates" [94]) may prolong satiety by their action in the
gastrointestinal tract. For example, slowly absorbed carbo-
hydrates travel further down the intestinal tract than do
rapidly absorbed carbohydrates [94] where they might ac-
tivate lower gut satiety mechanisms (see [105]).
Diet ft,-re. Although it has been discussed in previous
sections, it is worth emphasizing here the importance of
carbohydrate form as a determinant of food preference and
intake. The findings reviewed in this paper demonstrate that
rats respond quite differently to carbohydrate diets in hy-
drated (solutions, gels, semiliquids) and dehydrated forms
(powders). Both orosensory and postingestive factors anpear
to contribute to the greater hyperphagic response produced
by hydrated diets compared to dehydrated diets. That is, rats
prefer the flavor of carbohydrate solutions and gels to the
flavor of carbohydrate powders, and hydrated carbohydrates
appear to produce postingestive effects that enhance appe-
tite and food intake more than those produced by dehydrated
carbohydrates. The influence of carbohydrate form on appe-
tite and food intake has until recently received relatively
little attention. Dry diets are the norm in laboratory feeding
studies since they are more convenient to use and more re-
sistant to spoilage than are hydrated diets. Yet, animals and
people rarely eat dehydrated foods in the real world. Thus,
greater emphasis should be given in future research to the
study of hydrated foods.
Diet availability. Another factor that can influence carbo-
hydrate appetite, intake, and obesity is diet availability (Fig.
5). This is illustrated by the results of studies in which rats
147
were required to bar press for sucrose solution " m e a l s " on
fixed ratio schedules of 1 to 200 or more presses/meal
[30,32]. Increasing the bar-pressing requirement or " c o s t " to
obtain sucrose increased sucrose meal size but decreased the
number of meals such that daily sucrose intake was reduced.
Making the sucrose more costly also blocked its weight-
promoting effect, but apparently not its effect on feed effi-
ciency. That is, rats that worked for sucrose meals gained
more weight per calorie consumed than did control rats that
were not given access to the sucrose solution. As previously
mentioned, we have also observed that reducing diet avail-
ability by placing an antiscatter plate in the food cup was
sufficient to block the rat's hyperphagic response to the diet,
i.e., Polycose gel (Sclafani et al., unpublished findings). In
one experiment we observed that when the antiscatter plate
was removed the animals did not recover their hyperphagia
but nevertheless continued to gain more weight than chow-
fed controls. This finding raises the intriguing possibility that
reductions in diet availability may actually increase feed ef-
ficiency under certain conditions.
Limiting the availability of a carbohydrate source also
appears to increase the animal's carbohydrate appetite. This
is suggested by the results of a study [112] in which rats were
maintained in a complex environment in which they had to
bar press for all their food and water. Both sucrose and
regular food pellets were available from separate feeders.
When the daily supply of sucrose pellets was limited the rats
increased their rate of bar pressing for sucrose even though
the regular food pellets remained available and the animals
were never calorically deprived. Thus, rats seem to behave
like humans: when the supply of sugar is reduced, the de-
mand increases. Conversely, other studies show that when
rats are fed a sweet, palatable maintenance diet their appetite
for a sugar meal is reduced [125,163]. Since in the real world
animals must forage for food that varies in abundance and
availability, as well as in caloric value and nutrient composi-
tion, the influence of diet availability on appetite, intake and
feed efficiency is of considerable interest and requires much
further study.
A related issue concerns the availability of nutrients other
than carbohydrates. As discussed in a previous section,
under some conditions at least, the protein and fat content of
the diet has a significant impact on the rat's preference and
intake of carbohydrate-rich foods [69, 113, 118, 130, 211]
(see also [30]). A considerable amount of research has focused
on the factors involved in carbohydrate and protein selec-
tion, particularly the involvement of specific brain neuro-
transmitter systems, but a discussion of this work is beyond
the scope of the present paper (see [2, 4, 61]).
Importance of Carbohydrate Taste
Given the multiplicity of factors reviewed above, what
can be concluded concerning the relative importance of
carbohydrate taste in determining carbohydrate appetite and
intake'? With respect to the original question addressed by
our research--the role of sweet taste in carbohydrate-
induced overeating--it can be concluded that sweet taste is
not essential since rats overeat and become obese when of-
fered nonsweet polysaccharide solutions. However, since it
is now apparent that rats are very attracted to the taste of
polysaccharides, the question must be rephrased: what is the
role of carbohydrate taste, sweet or polysaccharide, in
carbohydrate-induced overeating and obesity? Clearly,
orosensory stimulation facilitates the feeding response of
148
rats [20,85], and the tastes of sugar and polysaccharides
provide a strong incentive to feed, but are these specific
tastes necessary to obtain the hyperphagic response to
carbohydrates? Or are the postingestive effects of carbo-
hydrates sufficient to induce and/or maintain carbohydrate
appetite and carbohydrate-induced hyperphagia in the ab-
sence of a sugar or polysaccharide taste? The answer to this
question is not known, but it appears likely that it is the
combination of taste and postingestive effects that make
carbohydrate foods so attractive to animals.
Other Nutrient Receptors?
The discovery of a heretofore unknown taste sensitivity
to starch-derived polysaccharides expands our understand-
ing of the "taste worlds" of animals. It also raises the
possibility that other taste sensitivities remain to be dis-
covered. Is it the case, as the quotation at the beginning of
this paper indicates, that animals have orosensory receptors
for carbohydrates only or is it possible that receptors for fats
and proteins exist in some species (see [51]). Much also re-
mains to be learned about the location, nutrient specificity,
and function of visceral receptors responsible for the
postingestive modulation of food preferences.
Relevance to Human Feeding and Obesity
The role of dietary sugar, sucrose in particular, in human
feeding behavior and obesity has been the subject of consid-
erable concern and controversy. While it is a popular belief
that sweet taste promotes overeating and sugar promotes
obesity in humans, the data to support this belief are lacking
[54,210]. The results of animals studies have been cited as
evidence both for and against the view that sugar causes
overeating and obesity [64, 143, 206]. There are of course
many differences between rats and humans as well as be-
tween the environmental conditions of the laboratory and the
real world. These differences limit the inferences that can be
made from animal studies concerning human behavior and
nutrition. Nevertheless, the findings obtained with labora-
tory rats suggest possibilities that should be considered and
variables that should be examined in the human feeding re-
sponse to carbohydrates.
Much of the discussion in the literature has focused on the
question: Is sucrose more obesity-promoting than other
sugars and starch?. The research reviewed above indicates
that sucrose is more obesity-promoting than other carbo-
hydrates but only under certain dietary conditions and the
relevance of these conditions to the human situation has
been questioned [143]. The rat studies also demonstrate,
however, that monosaccharides and polysaccharides can
produce overeating and obesity, and that the physical form
of the carbohydrate is at least as important as carbohydrate
type in determining food intake and adiposity. Rate of ab-
sorption, as influenced by diet form, composition, and other
factors, appears to be a critical determinant of the feeding
and body weight response to carbohydrates. Perhaps less
emphasis should be placed on the sugar vs. starch
dichotomy, and more attention should be directed to the
influence of rapidly absorbed vs. slowly absorbed carbo-
hydrate foods on human adiposity. This latter classification
scheme has received considerable attention recently in the
dietary treatment of diabetes mellitus [45,93].
The role of sweet taste in human obesity has also received
much attention. The rat studies reviewed here indicate that
sweet taste is not essential to obtain carbohydrate-induced
SCLAFANI
obesity since rats are also very attracted to the taste of
polysaccharides. Although humans lack this appetite for
polysaccharide taste, we are attracted to starch-rich foods
appropriately flavored with sauces and spices (see [60]). The
source of this attraction is not entirely understood [154].
Findings obtained with rats demonstrate that postingestive
factors have an important modulatory influence on food
preferences. In some cases postingestive factors may re-
verse initial food choices and the animal's hedonic response
in short-term tests do not always predict its long-term pref-
erence and intake. Human feeding studies indicate that the
hedonic response to taste does not predict long-term food
selection and intake [120]. Thus, it may be that in humans, as
in rats, the postingestive effects of foods condition our appe-
tites and flavor preferences, although clearly many other fac-
tors are involved in human food selection [154].
CONCLUSIONS
I. Rats and presumably other rodents have two different
taste (sub)systems for carbohydrates. The sweet taste sysem
is stimulated by sucrose, other sugars, and saccharin, while
the polysaccharide taste system is stimulated by starch-
derived polysaccharides, oligosaccharides, maltose, and
glucose. Rats display an avid appetite for both carbohydrate
tastes and their relative preference varies as a function of
carbohydrate type and concentration. The presumed func-
tion of the polysaccharide taste system is to facilitate the
detection and recognition of starch-rich foods but the
orosensory determinants of starch preference are not fully
understood.
2. Postingestive factors have an important modulatory ef-
fect on carbohydrate appetite and preference. The
postingestive actions of carbohydrates can condition flavor
preferences and can in some cases reverse the animal's ini-
tial food choices. The postingestive mechanisms responsible
for this appetite modulatory effect remain to be identified.
3. Other factors that influence carbohydrate appetite and
intake include the diet availability and macronutrient com-
position. Cephalic-phase digestive responses have also been
implicated but their role in carbohydrate appetite is ques-
tionable.
4. The long-term feeding and body weight response to
carbohydrate diets depends upon a number of factors. Diet
form is a particularly important variable; carbohydrates in
hydrated form (liquid or gel) promote more overeating and
obesity than do carbohydrates in dehydrated form (powder).
This may be related to the differential rate of absorption of
hydrated and dehydrated carbohydrates. Less important is
the specific type of carbohydrate since monosaccharides
(glucose, fructose), disaccharides (sucrose, maltose), and
polysaccharides (Polycose) all promote obesity when offered
m solution form. However. carbohydrate type is a significant
factor under some dietary conditions. For example, with
composite diets dissacharides are often more effective than
monosaccharides or polysaccharides in promoting obesity.
5. Results obtained with rats suggest that research on the
role of carbohydrates in human feeding and obesity should
not be limited to sugars and sweet taste. The influence of
other carbohydrates, rate of carbohydrate digestion and ab-
sorption, and the postingestive modulation of appetite
should be considered.
6. The discovery of a polysaccharide taste sensitivity in
rodents raises the question as to what other nutrient-specific
sensory receptors may exist in animals.
CARBOHYDRATE TASTE AND APPETITE
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