Physiology & Behavior, Vol. 23, pp. 241-247. Pergamon Press and Brain Research Publ., 1979.

Printed in the U.S.A.

Exercise and the Development and Persistence

of Dietary Obesity in Male and Female Rats
B A R B A R A J. R O L L S A N D E D W A R D A . R O W E

The Department of Experimental Psychology, University of Oxford, South Parks Road, Oxford OX1 3UD, Great
Britain

R e c e i v e d 22 N o v e m b e r 1978

ROLLS, B. J., AND E. A. ROWE. Exercise and the development and persistence of dietary obesity in male and female
rats. PHYSIOL. BEHAV. 23(2) 241-247, 1979.--Adult male and female hooded rats were housed in sedentary conditions
or were given free access to a running wheel. Exercising and sedentary rats received either a palatable, mixed, high energy
diet with chow (experimental group) or only chow (control group). Exercise reduced the weight gain of the males but
not of the females. All experimental groups preferentially selected the palatable foods. Both exercising and sedentary
females and the sedentary males became obese compared to their controls, but the exercising males did not. The mixed diet
was withdrawn after 10 weeks: thereafter the male and female sedentary experimental groups maintained the elevated body
weight. The exercising experimental females showed significant weight loss. Analysis of x-ray photographs indicated that
elevated body weight in the experimental rats probably reflected increased deposition of fat and not skeletal growth. The
results show that the effect of exercise on the development of dietary obesity is different in males and females, and that
sedentary male and female rats can both show persistent dietary obesity after withdrawal of the palatable foods.

Exercise Feeding Obesity Sex differences

S E D E N T A R Y rats offered a palatable, mixed diet ingest
excess energy and become obese [7, 8, 9, 10, 11, 12]. The
present report is addressed to the question of whether such
rats will develop obesity when given the opportunity to vary
energy expenditure through voluntary exercise. Pitts and
Bull [6] found that male C F E albino rats did not become
obese on a high fat diet when forced to exercise on a tread-
mill. On the other hand Sclafani and Springer [12] found that
female C F E rats eating a high energy supermarket diet be-
came obese when given free access to a running wheel.
These females did not, however, gain as much weight as
sedentary rats eating a similar diet2 It is not clear whether
these two studies indicate that there is sex difference in the
effect of exercise on the development of obesity or whether
the results simply reflect some difference in experimental
method.
It has been suggested that the sexes show different re-
sponses to dietary changes a a d to the effect_ of exercise on
feeding. Females respond to dietary dilution by adjusting
food intake, and show compensation for increased energy
expenditure during enforced exercise by hyperphagia. Males
are less responsive to dietary dilution and do not show com-
pensation for increased energy expenditure during enforced
exercise. Thus females appear to regulate body weight more
precisely than males in the face of changing energy intakes
and energy expenditure [4,5]. One aim of the present study is
to determine whether males and females differ in their re-
sponse to a high energy diet and to the effect of voluntary
exercise on the development of dietary obesity.
In some studies male rats not only become obese while
c o n s u m i n g a high e n e r g y diet, b u t r e t a i n m o s t o f the ex-
c e s s w e i g h t w h e n the c a l o r i f i c diet is w i t h d r a w n [7, 8, 9].
In another study female rats have been found to return to
control weights when the fattening diet is withdrawn [12]. It
is not clear whether this difference reflects sexual dimorph-
ism or a difference in experimental procedure. The present
study is designed to determine whether obesity will persist
after withdrawal of the mixed diet in both sexes, and also to
determine how voluntary exercise affects the persistence of
obesity.
GENERAL METHOD
Animals
Hooded Lister rats 9 to 12 weeks of age, were used. The
animals were maintained in conditions of controlled tem-
perature (23°C __. 1°) and lighting (12-hr light, 12-hr dark),
and water and laboratory chow were freely available at all
times.
Procedure
The rats were matched for body weight, and allocated to
two equal size groups: the sedentary group, or the exercise
group. Animals in the sedentary condition were individually
housed in small metal cages (34 x 15 x 13.5 cm), animals
in the exercise condition were individually housed in small
cages of similar size with a running wheel (diameter 32 cm)
attached. After a two week period of adaptation to the hous-
ing conditions, the animals within the exercise and sedentary

1We gratefully acknowledge the advice given by E. T. Rolls and the assistance of N. McNaughton and G. C. Preston with the statistical
analysis. This Research was supported by the Medical Research Council of Great Britain. We thank Symbol Biscuits Ltd for supplying the
Maryland Cookies, and United Biscuits for supplying the Cheddars.

C o p y r i g h t © 1979 B r a i n R e s e a r c h P u b l i c a t i o n s Inc.--0031-9384/79/080241-07501.20/0
 242 ROLLS AND ROWE

groups were separately matched for body weight and allo-

cated to the two dietary conditions: experimental which re-
ceived the mixed diet and chow, or control which received • mixed d i e t . exercise end mixed diet
only chow. The mixed diet contained three supermarket • mixed diet
foods: plain salted potato chips (Golden Wonder crisps), 500 z~ chow + exercise
chocolate chip cookies (Maryland Cookies, Lyons), and o chow
cheese crackers (Cheddars, Crawfords) energy value of the
palatable, mixed diet: 20.3 to 23.4 KJ/g: energy value of
chow: 16.3 KJ/g from data supplied by the manufacturers.
See [8] for composition of foods. The mixed diet was offered 450
for 10 weeks, was then withdrawn, and the rats continued to
receive free access to chow. Body weights, water intakes,
and activity (number of revolutions of the activity wheel)
were recorded daily throughout the experiment, and food start ""
mixed diet A" "A" ~lk zr A
intakes were recorded at the beginning and end of the wt. 400 , .~t.j.
fattening period, the day after the mixed diet was withdrawn (g) .A'"• .~.~-"
and thereafter at weekly intervals for seven weeks. Food &.A ~ ..,ff .Z~ .zy
intakes during the fattening period were determined by
measuring separately the amounts of the mixed diet and 350 )i" b..t Y
chow consumed over twenty-four hours. The measurements
were corrected for spillage which was collected in a paper
trap suspended beneath each cage.
The experiment was conducted in three successive runs,
and at the end of runs two and three the rats were anes- 3oo r/~ I
thetized with chloro-nembutal (3 ml/kg) and x-ray photo-
graphs were taken with the animals positioned as described
by Hughes and Tanner [2]. Pelvis and femur lengths, and the -2 0 2 4 6 8 10 12 ¼ 16
greatest width of the animals were then measured from the Weeks
x-ray photographs.
FIG. 1. The mean body weights for the male exercisingexperimental
(N=8) and control (N=8) groups, and sedentary experimental
Statistics (N=8) and control (N=8) groups during the period of adaptation
Weekly measurements of body weights were examined by while the mixed diet was offered, and alter withdrawal of the mixed
analysis of variance with exercise × diet x time (weeks) as diet.
factors, separately for the fattening period, and for the
period after the withdrawal of the mixed diet. Comparisons
of group means were made using the appropriate error term Influence of exercise. The animals in the sedentary con-
from the analysis of variance. Food intakes, water intakes dition became significantly heavier than the animals with
and activity were similarly analyzed. To reduce variability in access to the running wheels after the two-week period of
activity levels due to the estrus cycle, daily activity was adaptation to the housing conditions (t=2.9, 30, p<O.Ol).
averaged over four consecutive days each week in all exer- During the fattening period both the sedentary animals re-
cising groups. Rates of weight gain were examined by ceiving the mixed diet, and the sedentary animals receiving
analysis of variance of the regression coefficients (with only chow, exhibited greater rates of weight gain than the
time). Comparisons of group means were made using the experimental and control exercising groups (F=216.6, 1,279,
Student's t test (two-tailed). p<0.001; for the mixed diet t=20.5, 14, p<0.001; for chow
t=8.9, 14, p<0.001).
Influence of diet. Both the sedentary and exercising
EXPERIMENT 1 groups receiving the mixed diet exhibited greater rates of
Experiment 1 determines whether male rats given the weight gain than their respective controls receiving chow
opportunity to exercise freely in a running wheel become (F=170.4, 1,279, p<0.01; for the sedentary groups t=18.8,
obese while consuming a palatable, mixed energy dense diet. 14, p<0.001; for the exercising groups t=7.3, 14, p<0.001)
The effect of withdrawal of the palatable diet on body hut the mixed diet had a greater effect on rate of weight gain
weight maintenance, chow intake and amount of exercise relative to controls in the sedentary animals than in the
was also determined. exercising animals (diet × exercise interaction: F=33.1,
1,279, p <0.01).
METHOD Ten weeks after the mixed diet was introduced the seden-
The animals were thirty-two male rats, 259 g to 335 g. tary experimental rats were significantly heavier than seden-
Two animals per group were used in the first run, and three tary control rats (t =4.8, 14, p<0.01), and the exercising ex-
animals per group were used in the second and third runs. perimental rats were not significantly heavier than the exer-
cising control rats (t=l.7, 14, N.S.), despite exhibiting a
RESULTS greater rate of weight gain.
Body Weights After withdrawal of the mixed diet the sedentary and
exercising experimental groups exhibited a small loss of
The mean body weights of the exercising and sedentary weight which rapidly stabilized, and after four weeks the
experimental and control groups are shown in Fig. 1. experimental animals began to increase in weight, maintain-
E X E R C I S E A N D D I E T A R Y OBESITY 243

TABLE 1
DAILY FOOD INTAKES (g) OF CHOW AND THE MIXED DIET (MEAN _.+ SE OF MEAN)
AT THE BEGINNING (WEEK 1) AND THE END (WEEK 10) OF THE FAITI'~NIIKG
PERIOD FOR MALES

Week
1 10
Condition Chow Mixed Chow Mixed

Exercise/Mixed 3.9 --- 1.6 16.4 _ 1.6 4.1 ___ 1.1 18.2 --- 1.5
Exercise/Chow 23.6 --- 1.0 -- 22.0 --- 1.5 --
Sedentary/Mixed 7.1 - 1.2 15.6 --- 1.4 7.7 +- 1.0 13.3 - 1.4
Sedentary/Chow 23.9 _+ 1.4 -- 24.5 - 0.8 --

Food Intake
I • mixeddiet c¢'
& chow The mean food intakes at the beginning and end of the
fattening period are shown in Table 1. At these times the
mixed diet was selected in preference to chow. There were
no consistent differences in the total amount of food eaten at
the beginning and end of the fattening period. The exercising
experimental rats consumed proportionately more of the
0
g
A / ~ end mixeddiet mixed diet than the sedentary experimental rats and propor-
tionately less chow (t=5.0, 14, p<0.001).
Immediately after withdrawal of the mixed diet the exer-
cising and sedentary experimental rats consumed slightly but
t~ not significantly less than the control rats. Mean intakes of
C
9 chow ( _ S.E.) on the day after withdrawing the mixed diet
were: exercising experimental 19.4 _ 0.7 g; exercising con-
>
0 trol 21.0 _ 0.9 g: sedentary experimental 18.5 _ 0.5 g:
Ill
n- sedentary control 20.4 _ 0.7 g. Subsequently the food in-
takes of both experimental groups increased while there
were no consistent changes in the food intakes of the control
I I I , I I I I [ l * l i I
groups (for comparison of the linear regression co-efficients
0 2 t, 6 8 10 12 14 16 between the experimental and control animals: F = 10.8 1,192
Weeks p<0.01: diet x exercise interaction: N.S.). Seven weeks
FIG. 2. The voluntary activity averaged over four days each week after withdrawal of the mixed diet the mean intakes of chow
for the male experimental and control groups while the mixed diet (___ S.E.) were: exercising experimental 24.1 _ 1.2 g:
was offered and after withdrawal of the mixed diet. The mixed diet exercising control 21.7 _ 1.2 g: sedentary, experimental
was introduced after week 0. 25.5 ___ 2.3 g: sedentary control 22.3 _ 1.2 g.

ing the elevated weights above the weight of the control
animals. At the end of the experiment, seven weeks after
withdrawal of the mixed diet, the exercising experimental
rats were twenty-five g heavier than the exercising control
rats ( t = l . 4 , 14; N.S.) and the sedentary experimental rats
were forty g heavier than the sedentary control rats (t =2.4,
14, p <0.05).
Activity
The mean daily, levels of activity are plotted weekly for
the experimental and control exercising groups in Fig. 2.
Both groups exhibited an increase in activity during the
period of adaptation and .the beginning of the fattening
period, but after approximately four weeks activity declined
in both groups to the end of the experiment. Although there
were no significant differences in activity between groups
during the fattening period or after withdrawal of the mixed
diet, the animals receiving the mixed diet took consistently
more exercise than the control rats during the fattening
period, and less exercise than the control rats for most of the
period after the mixed diet was withdrawn.
Water Intake
Water intake was significantly greater in the exercising
males than the sedentary males throughout the experiment
(F=61.5 1,28 p<0.001).The control males receiving chow
consumed significantly more water than the experimental
males receiving the mixed diet during the fattening period
(F=27.8 1,28p<0.001), the effect of the diet being greater in
the exercising groups than the sedentary groups (interaction
between diet and exercise: F = 9 . 0 1,28 p<0.01). After with-
drawal of the mixed diet the water intakes of the experi-
mental males increased to control levels.
Body Weight and Activity
The exercising experimental rats were slightly heavier
and were taking more exercise than the exercising control
rats ten weeks after the introduction of the mixed diet, al-
though these differences were not statistically significant. It
is possible that the exercising experimental rats were ex-
pending more energy during the fattening period than the
exercising control rats, and this may have reduced the differ-
ence between their weight gains during the fattening period.
244
TABLE 2
BODY MEASUREMENTS (MEAN - SE OF MEAN) FOR MALES (N=5 PER GROUP) AND FEMALES (N=6 PER GROUP) TAKEN
FROM X-RAY PHOTOGRAPHS SEVEN WEEKS AFTER WITHDRAWAL OF THE MIXED DIET
Males
Condition Pelvis Length F e m u r Length
(cm) (cm)
Exercise/Mixed 45.2 ± 0.5 35.7 -+ 0.2
Exercise/Chow 45.3 ± 0.6 36.4 _+ 0.2
Sedentary/Mixed 46.0 ± 0.7 36.0 +_ 0.3
Sedentary/Control 45.4 ± 0.8 36.6 _+ 0.4
This is indicated by the finding that although the difference
between absolute body weights was not significant, when an
analysis of covariance of these body weights was performed
to take into account the differences in exercise, the adjusted
body weights of the exercising experimental rats were sig-
nificantly greater than the adjusted body weights of the
exercising control rats (t=2.2, 14, p<0.05).
After the withdrawal of the mixed diet the exercising ex-
perimental rats maintained the same degree of elevated
weight above the control rats; and the exercising groups con-
sumed equal amounts of food. The activity levels of the
exercising experimental rats appeared to fall below the levels
of the exercising control rats when the mixed diet was with-
drawn, although this difference was not significant, which
suggests that some conservation of energy expenditure may
have taken place.
Skeletal Size
The measurements of skeletal size are shown in Table 2.
There were no significant differences in lengths of the femur
or pelvis between any of the groups. However, the sedentary
experimental rats had significantly greater body widths than
the sedentary control rats (t=2.4, 14, p<0.05), and the
sedentary experimental and control rats had significantly
greater body widths than the respective exercising groups
(for experimental rats t=6.6, 14, p<0.001; for the control
rats t=3.9, 14, p<0.01).
DISCUSSION
The sedentary males receiving the mixed diet preferen-
tially selected the energy-dense foods and became signifi-
cantly obese compared to the control rats. When the mixed
diet was withdrawn, the experimental rats maintained the
elevated body weight and ate the same amount of chow as
the sedentary control rats. The body measurements
suggested that the elevated weight of the experimental rats
did not represent greater growth in the animals, and there-
fore probably represents a permanent accumulation of fat.
These results confkrm the results of a previous study [8].
The exercising males exhibited greatly reduced weight
gains compared to the corresponding sedentary males. The
food intake measurements indicated that the exercising con-
trol rats consumed less food than the sedentary control rats,
and this, with the greater energy expenditure of voluntary
exercise, is consistent with the low weight gain. In the fatten-
ing pet'iod the exercising experimental rats preferentially
selected the palatable food(Table 1). It is possible that they
were consuming more energy than the sedentary experi-
ROLLS AND ROWE
Females
Width Pelvis Length F e m u r Length Width
(cm) (cm) (cm) (cm)
75.0 ± 1.7 42.3 ___ 0.5 33.0 -+ 0.4 73.5 ± 3.7
72.0 ± 1.3 42.5 ± 0.6 32.4 ± 0.4 67.8 ± 3.2
92.1 -+ 1.9 38.5 ± 0.8 32.2 ± 0.5 78.6 ± 2.4
84.0 ± 2.8 39.7 ± 0.8 31.4 _+ 0.6 70.8 _+ 2.0
mental rats, because while taking approximately the same
quantity of food, the exercising experimental rats took a
greater proportion of the palatable, high energy foods than
the sedentary experimental rats. This is consistent with the
results of Collier, Leshner and Squibb [1] which show that
when given a choice between a high carbohydrate or high
protein diet, active rats select a higher proportion of carbo-
hydrate than inactive rats.
Although the rate of weight gain of the exercising experi-
mental animals was significantly greater than the exercising
control animals, it was insufficient to induce s i ~ a n t l y
elevated body weights above the exercising control animals
during the ten-week fattening period. This may he explained
partly by a tendency for the experimental males to display
greater activity than the control males, and suggests that the
exercising experimental males may have compensated for
the surplus energy intake by expending more energy than the
exercising control males.
EXPERIMENT 2
In Experiment 1, sedentary males receiving a mixed
energy-dense diet developed obesity, which persisted when
the diet was withdrawn and replaced by a uniform, calori-
cally less dense diet (chow). Exercising males receiving the
mixed diet failed to develop obesity during the fattening
period. Because females are more responsive m dietary di-
lution than males and show little weight change in response
to enforced exercise [5], the responses of female rats to ac-
cess to and subsequent withdrawal of the mixed diet under
sedentary and exercising conditions were examined.
METHOD
The animals were thirty-six female rats, 174 g to 225 g.
The experiment was conducted as three successive runs,
with three animals per group in each run.
RESULTS
Body Weights
The mean body weights of the exercising and sedentary
experimental and control groups are shown in Fig. 3.
Influence of exercise. Exercise had n o s ~ i f l c a n t effect
on body weight during the fattening period (F=I,0,: 1.32
N.S.) and there was no significant difference between the
rate of weight gain of the exercising and sedentary animals
(F=3.7, 1,320, N.S.).
ROLLS AND ROWE 245

350
• mixed diet • exercise
• mixed diet
A c h o w . exercise end mixed diet
@
&. mixed
& chow
diet] ¢~)
o chow

8
300
% .

Body 7
F .A""
wt. 0
mixed diet ~ ~ Jr 0
(g) O 6
250
x
5
tn
¢.
o
m,P 4
200
0 end mixed diet
> 3
.t t i t . . . . , I ! I ! ! t ! . , J .
,,y
-2 0 2 4 6 8 10 12 14 16
Weeks
2
FIG. 3. The mean body weights for the female exercising experi-
mental (N=9) and control (N=9) groups, and the sedentary experi- I
mental (N =9) and control fN =9) groups during the period of adap-
tation, while the mixed diet was offered, and after withdrawal of the t | i ! t t i i t | J I i I | t |

mixed diet. 0
0 2 4 6 8 10 12 14 16
Weeks
FIG. 4. The voluntary activity averaged over four days each week
Influence of diet. Both the sedentary and exercising for the female experimental and control groups while the mixed diet
groups receiving the mixed diet exhibited greater rates of was offered and after withdrawal of the mixed diet. The mixed diet
weight gain than their respective controls receiving chow was introduced after week 0.
(F=194, 1,320, p<0.001; for the sedentary groups t=15.4,
16, p <0.001; for the exercising groups t = 12.4, 16, p <0.001),
and the mixed diet had a similar effect on rate of weight gain
in the exercising and sedentary groups (diet × exercise in-
teraction: F=2.3, 1,320 N.S.). Food Intake
Ten weeks after the introduction of the mixed diet the
exercising experimental rats and the sedentary experimental The mean food intakes at the beginning and the end of the
rats were signifmantly heavier than their respective control fatting period are shown in Table 3. At these times the mixed
groups (for the exercising groups t--4.9, 16, p<0.001; for the diet was selected in preference to chow, and the exercising
sedentary groups t=5.0, 16, p<0.001). experimental rats consumed proportionately more of the
After withdrawal of the mixed diet the sedentary experi- mixed diet than the sedentary experimental rats, and pro-
mental rats exhibited a small loss of weight which stabilized portionately less chow (t=7.5, 16, p<0.01).The exercising
after two weeks. At the end of the experiment, seven weeks control rats consumed more chow than the sedentary control
after withdrawal of the mixed diet the sedentary experi- rats in the fattening period (Table 3) and after withdrawal of
mental rats remained signifcanfly heavier than the sedentary the mixed diet (F=14.7, 1,32, p<0.001).
control rats (t=2.2, 16, p<0.05). The exercising experi- Immediately after withdrawal of the mixed diet the seden-
mental animals lost more weight than the sedentary experi- taw experimental animals consumed slightly, but not signifi-
mental animals over the fast week after withdrawal of the cantly less than the control animals; the exercising experi-
mixed diet (t =2.6, 16, p<0.02). Subsequently the weights of mental rats consumed si£mificantly less than the exercising
the exercising experimental rats stabilized, but at the end of control rats for one week after the withdrawal of the mixed
the experiment the body weights of the exercising experi- diet (t=2.8, 16,p<0.02). Mean intakes of chow ( - S.E.) on
mental rats were not significantly greater than exercising the day after withdrawing the mixed diet were: exercising
control rats (t=2.0, 16, N.S.). experimental I2.1 _+ 0.9g; exercising control 17.8 +_ 1.0
g; sedentary experimental 13.4 _+ 0.5 g; sedentary control
Activity 16.4 + 1.0 g. Subsequently the food intakes of both exper-
imental groups increased, while there were no consistent
The mean daily levels of activity are plotted weekly for changes in food intakes of the control groups (for compari-
the experimental and control groups in Fig. 4. son of the linear regression coefficients between the experi-
During the experiment the levels of exercise of the exper- mental and control groups F--14.6 1,222 p<0.001; diet ×
imental and control groups were not significantly different. exercise interaction N.S.). Seven weeks after withdrawal of
Activity increased in both groups for two weeks after the the mixed diet the mean intakes of chow ( _ S.E.) were:
mixed diet was introduced and thereafter declined. exercising experimental 21.7 +_ 1.9 g; exercising control
Throughout the experiment the females took consistently 18.7 _ 2.1 g; sedentary experimental 17.8 +_ 1.3 g; seden-
more exercise than the males (F--17.6 1,33 p<0.001). tary control 14.9 _+ 1.3 g.
246
TABLE 3
DALLY FOOD INTAKES (g) OF CHOW AND THE MIXED DIET (MEAN --. SE OF MEAN)
AT THE BEGINNING (WEEK 1) AND END (WEEK 10) OF THE FATTENING PERIOD
FOR FEMALES
Condition Chow
Exercise/Mixed 3.7 ± 1.2
Exercise/Chow 19.2 + 1.0
Sedentary/Mixed 3.6 ± 0.6
Sedentary/Chow 18.3 ± 1.3
Water Intake
Water intake was significantly greater in the exercising
females than the sedentary females throughout the experi-
ment (F=30.9 1,32 p<0.001), and the control females con-
sumed significantly more water during the fattening period
than the experimental females receiving the mixed diet
(F=31.9 1,32 p<0.001). After withdrawal of the mixed diet
the water intakes of the experimental females increased to
control levels.
Skeletal Size
The measurements of skeletal size are shown in Table 2.
The pelvis lengths of the exercising groups were significantly
greater than their respective sedentary groups (for the exper-
imental rats t =2.5, 10, p <0.05; for the control rats t =4.1, 10,
p<0.01), but there were no other significant differences in
skeletal lengths between any of the groups. The sedentary
experimental rats had significantly greater body widths than
the sedentary control rats (t=2.5, 10, p<0.05), but there
were no other significant differences between the groups in
body width.
DISCUSSION
Both the sedentary and exercising females receiving the
mixed diet preferentially selected the palatable foods and
became significantly obese compared to their respective con-
trol groups. Exercise had little effect on the weight gains of
the experimental or control groups: the exercising control
group appeared to adjust to the greater energy expenditure of
exercise by greater energy consumption throughout the ex-
periment compared to the sedentary control group. The
exercising and sedentary experimental groups consumed
similar quantities of food in the fattening period (Table 3.),
but the exercising experimental rats consumed a greater pro-
portion of the energy-dense foods, suggesting that the energy
expenditure of exercise was compensated for by greater en-
ergy intake in the experimental rats. The activity levels of
the experimenatal and control groups were similar, thus vol-
untary energy expenditure did not appear to influence the
development of obesity in the exercising females.
After withdrawal of the mixed diet the sedentary experi-
mental rats exhibited a small loss of weight and modest but
not significant hypophagia, but after two w e ~ s the body
weights stabilized, elevated above the weights of the seden-
tary control rats, and the food intakes of the sedentary
groups became similar. The exe_r_cising experimental rats
exhibited greater weight loss than the sedentary experi-
ROLLS AND ROWE
Week
10
Mixed Chow Mixed
18.6 ± 1.6 2.6 __+-1.0 14.9 ± 1.9
-- 20.6 --- 1.1 --
13.8 ± 0.7 5.9 -+ 1.I 9.5 ± 1.2
-- 18.3 -+ 1.2 --
mental rats, and significant hypophagia compared to the
exercising control rats after withdrawal of the mixed diet.
After two weeks the body weights of the exe~ising experi-
mental rats stabilized but they were no longer s i ~ i f i c ~ y
heavier than the exercising control rats. The food intakes of
the exercising groups became similar one week after the
withdrawal of the mixed diet.
GENERAL DISCUSSION
Obesity in Male and Female Sedentary Rats
Both male and female sedentary rats become obese when
offered a variety of palatable, high energy foods. The ohesity
persists when the mixed diet is withdrawn, at which time
chow intakes are similar to the intakes by the control ani-
mals. The two sexes displayed similar weight p i n s relative to
controls during the fattening period (the experimental males
gained 69 g more than the control males, the experimental
females gained 63 g more than the control females).
Moreover, the two sexes achieved similar percentage weight
gains relative to the weights at the start of the fattening
period (the obese males gained 54% of their initial weight; the
obese females gained 55% of their initial weight). This result
differs from a previous report in which females achieved greater
absolute weight gains compared to the controls than the
males when they were offered the mixed diet [11]. In that
study a wider range of foods and different strain of rat (CFE
albino) were used.
Sedentary males and females exhibit some differences in
response to withdrawal of the mixed diet: the obese females
appear to lose more weight than the males (in the first two
weeks after withdrawing the mixed diet the females showed
changes of + 11 to -32 g in weight, males showed changes of
+ 10 to -12 g). After a period of stable body weight, the
obese males increase at a comparable rate to the control rats,
while the females do not show a pronounced weight in-
crease. This may be related to the sexual dimorphism in
growth pattern of the rat [3]. Males grow throughout adult
life, females reach an asymptotic weight. When account is
taken of the weight changes of the control rats, both obese
males and females appear to maintain the elzvated body
weight after withdrawal of the mixed diet. The relative
change between the weights of the experimental and control
rats after withdrawal of the mixed diet is the same in males
and females. Therefore it appears that in both sexes the pat-
tern of weight change after removal of the mixed diet is
appropriate to the maintenance of body weight above control
level~.
E X E R C I S E A N D DIETARY OBESITY
Effect o f Exercise on Body Weight in Males and Females
Sexual dimorphism is further expressed in the response to
exercise shown by rats which are eating chow [4]. Exercise
markedly reduces the weight gains of males, but has little
effect on females. Energy expenditure incurred during exer-
cise will include the work required to rotate the activity
wheels and metabolic energy expenditure, which may be in-
creased during exercise. Males do not appear to compensate
for the increased energy expenditure during exercise by ad-
justment of energy intake; females do adjust energy intake to
compensate for energy expenditure. It appears that there is a
greater potential for the energy demands of exercise to effect
body weight in males than in females. Sedentary males prog-
ressively accumulate fat and it is likely that the main effect of
exercise on body composition is to reduce body fat [6].
Sedentary females which are eating chow show a smaller
tendency to accumulate fat. They reach asymptotic weights
[4].
Obesity in Male and Female Exercising Rats
The sexes also respond differently to the influence of vol-
untary exercise upon the development of obesity. Exercising
males do not become significantly obese when offered palat-
able, high energy diet. The failure to develop obesity may be
caused by insufficient energy intake which does not compen-
sate for the increased energy expenditure incurred in volun-
tary exercise. It appears that a further factor may be an
additional voluntary increase in energy expenditure by the
exercising males receiving the mixed diet, since they dis-
played a tendency to take more exercise. The patterns of
weight gain in response to exercise and high energy diet were
consistent with the results of Pitts and Bull [6], although in
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247
that study the forced exercise regime presumably minimized
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the sedentary females offered the mixed diet.
In the experimental exercising males the level of food
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was insufficient to maintain the elevated body weights, and
weight loss to control levels occurred.
A hormonal basis for sexual dimorphism in the growth of
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androgen potentiates the male pattern of growth. The female
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absence of androgen in the neonatal period [4]. The sexual
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