EXERCESE UFQN H'iE FUUU kUNE‘aUMFHU’N
AND BODY WEIGHTS OF YOUNG
lx’aALE. ALEINQ RATS

Tho-sh for Hm 02ch a? M. A.
MICHMAN STATE UNEVEBSETY
Gunéacs Straufiuiefica
1962

 

THESIS

  
     
   

f" LIBRARY

Michigan Stat:
' University

 

ABSTRACT

THE EFFECTS OF VARIOUS LEVELS OF.EXERCISE UPON
THE FOOD CONSUMPTION AND BODY WEIGHTS
OF YOUNG MAIE ALBINO RATS

by Gundars Strautnieks

The purpose of this experiment was to determine the
effects of various levels of exercise upon the food con~
sumption and body weights of young male albino rats. The
experiment was divided into two major parts. The first
part consisted of thirty-five days and will be referred
to as the treatment phase. The second part consisted of
one hundred and five days and will be referred to as the
post-treatment phase. The experiment ran one hundred
forty consecutive days.

Thirty rats of the Sprague-Dawley strain were used
for the entire experiment. Animals were divided into three
groups, each group consisting of ten rats. The Sedentary
Group was allowed sedentary activity for the treatment
phase but spontaneous activity for the post—treatrsnt
phase. The rats in the Spontaneous Group served as con—
trols and were allowed spontaneous activity for the entire
experiment. I

Individual rat body weight was determined daily for

the first seventy days, and every third day for the last

Gundars Strautnieks
seventy days. Animal food consumption was measured for
each individual rat throughout the entire experiment. The
spontaneous activity was measured daily for each individual

rat by means of revolution counters that were attached to

 

their cages. These data were recorded daily. T

Rotation order was developed so that each rat would .
be in the microswitch—equipped caged once for a 24-hour 9
period every five days. This was done because of another E

study that was run concurrently.

The results indicate that when young male albino rats
are forced to exercise at young age they do not gain as
much weight as those animals that are restricted from ac-
tivity or allowed to exercise at their own desire in the
Spontaneous activity wheels. Even though the activity
was different for each group of animals, there were no
significant differences between the group totals in their
food consumption. Finally, the rats that were forced to
exercise by swimming at an early age ran a significantly
greater number of revolutions than either the postwsaientary

or general group rats.

THE EFFECTS OF VARIOUS LEVELS OF EXERCISE UPON
THE FOOD CONSUMPTION AND BODY WEIGHTS
OF YOUNG MALE ALBINO RATS

By

Gundars Strautnieks

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF ARTS

Department of Health, Physical Education,
and Recreation

1962

ACKNOWLEDGMENTS

I wish to express my sincere gratitude to Dr. Wayne
Van Huss for his generous assistance and supervision
during the course of this study. I am also indebted to
Dale L. Hansen for his assistance in preparing and

carrying out this experiment.

DEDICATION

This thesis is dedicated to my wife, Astrida, for
her love and patience during this period of study and

research.

 

TABLE OF CONTENTS

CHAPTER

I.

II.

III.

IV.

THE P ROBLEM .

Introduction

Statement of the Problem

Justification of the Study . . . . . . . . .

Limitations of the Study

LITERATURE REVIEW

Introduction

0 O O O O O O O 0 O

O O O O 0 O O O

The Effects of Spontaneous Activity on Food

Intake and Growth

The Effects
The Effects
The Effects
The Effects

The Role of

of Animal Handling on Growth. . .

of Exercise on Growth . . . . .

of Diet on Activity and Growth.

of Exercise

Exercise in

METHODS OF PROCEDURE.

Introduction

Subjects and Equipment.

Training Program.

Testing Techniques.

RESULTS . . .

Food Consumption.

Animal Weights.

Spontaneous Activity.

Discussion.

Upon the Food Intake.

Weight Control. . .

O O O O O O O O O O 0

PAGE

[‘0

J—‘T-PTVUFO

\OCDNJ‘:

CHAPTER

v. SUMMARY, CONCLUSIONS,

Summary . . .

Conclusions .

Recommendations
BIBLIOGRAPHY . . .
APPENDICES . . .

AND RECOMMENDATIONS

J

'!"'

 

LIST OF TABLES

Analysis of Variance of Data for Food Con-
sumption During the Treatment Phase . . .

Analysis of Variance of Data for Food Con-
sumption During the Post-Treatment Phase.

Analysis of Variance of Data for Animal
Weights During the Treatment Phase.

Analysis of Variance of Animal Weight Dif-
ferences From the End of Treatment Phase
to the End of the Experiment. . . . . .

Analysis of Variance of Data for Animal
Weight Differences for the Entire Experi—
ment. . . . . . . . . . . . . . . . . . .

PAGE

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(4.)
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LIST OF FIGURES

FIGURE PAGE
1. Average Food Consumption Per Group Per Test
Peri-Od- O O O O I O O 0 O o 6 o o o O o o o o 30
2. Rat Body Weight Per Group. . . . . . . . . . . 3A
3. Average Spontaneous Wheel Revolutions Per
Group 0 O 0 O I O O O O o o o O O O O O O 38

 

'1:
4A..

CHAPTER I

 

THE PROBLEM
Introduction ‘
Overweight is associated with definite health hazards F
in terms of an increased incidence of serious disease and
impaired life expectancy. When all facts are considered, ,
it is apparent that overweight is a major health problem ;

 

of our day. Although the evidence is incomplete in some
respects, it seems quite clear that substantial health
gains would result if overweight could be either prevented
or remedied in an appreciable segment of our population.
Desirable weight is an individual thing. It may be
described as the weight at which a person both looks and
feels his best. A person's age, sex, height, bone struc»
ture, and muscular development all affect his weight.
Life is much easier in many ways for persons who
are not too fat. They usually feel and look better.
They are likely to live longer. They are less likely to
suffer from backaches, foot troubles, fatigue, and other
discomforts. Normal weight is worth any effort it takes
to reach and keep it-—worth in the terms of everyday
comfort and of a healthier, longer life.
This study is not expected to solve any weight con-

trol problems or affect the general health of humans

directly, it may, nevertheless, Shed some light upon the
‘problem that through further research may finally give

some real beneficial results.

Statement of the Problem

 

To study the effects of various levels of exercise
upon the food consumption and body weight of young male I.

albino rats.

Justification of the Study

 

We are starting to realize today that the lean people L’
of the past were so, not because they didn't have the food,
but simply because they led a more active life. The in—
creasing overweight problems and the complications that come
along with obesity are certainly on an upswing in the United
States. '

Even though the methods of controlling obesity are
crude and not universally effective under practical con-
ditions, progress can be made. Research planned or in pro-
gress should do much to improve our understanding of the
role of exercise on weight control as well as the basic
nature and causes of overweight and hence develop better

methods of prevention, diagnosis, and control.

Limitations of the Study
1. This is an animal study and therefore, whatever
results should be obtained, the application to humans

will always be complicated.

2. Some single animal may affect the whole group
average as far as the body weight or food consumption

is concerned.

3. The study was conducted using young male albino
rats.

A. The minimal amounts of food lost through col-
lecting and measuring methods.

5. The room temperature variations may have affected

the food intake.

CHAPTER II

LITERATURE REVIEW

Introduction

 

In recent years tremendous advances have been made T
towards a better understanding of the effects of exercise.

In spite of this advancement in many exercise areas, con-

 

siderably little has been done to understand the effects

of physical work on the food intake and body weights.
Various investigators have studied the influences

of certain diets and how they affect the body weights and

general food consumption. Others have investigated the

effects of spontaneous activity and handling upon growth

of animals.

The Effects of Spontaneous Activity on Food Intake and Growth
Slonaker (40) suggested that the fluctuation in spon~
taneous activity of the albino rat appear to be of normal
occurrence and that therefore must be considered in experi_
ments dealing with activity. In his animal work he found
that the female albino rat not only exhibits variations in
daily activity, known as the oestrual rhythm, but also shows
gradual fluctuation in average daily activity which extend
over long periods of time. Slonaker also found that fluc-

tuations in average daily activity are common to both sexes.

\fl

He further states that the lack of synchronism of these
fluctuations in a group of animals indicates that changes
in environment such as moisture, temperature or barometric
pressure, are not the cause. Nevertheless, that a greater

tendency toward synchronous fluctuation existed in the

 

males than in the females. t
Hetherington and Hanson (24) investigated spontaneous

running activity both before and after Operation of 10

rats with hypothalamic lesions frequently causing adiposity

and of 6 normal litter-mate controls. It was observed that i

the animals having bilateral lesions in the hypothalamus
tended to indulge in much less spontaneous running than

did the majority of normal controls. The food consumption
of the obese Operated animals greatly exceeded the intake
of normal controls or did not exceed at all, depending upon
the nature of the food supplied. A soft palatable diet
encouraged maximum consumption, whereas the hard dry pellet
diet apparently discouraged higher intake.

Shirley (37) Slonaker (Al) and Slonaker (42) after con—
ducting extensive experiments on the activity of rats
throughout the major part of the life period, Showed that
young and very old animals are quite inactive, but that
there is little change in activity level from puberty to
the age of one hundred days. They also demonstrated that
the activity of each animal tends to a relatively constant

value during the major part of the life cycle, with the

female rats being much more active than the male. In
addition, Slonaker Showed that during the prime of life
the activity is confined principally to night, and rest
to day time. Of the whole amount of work done during the
life time, almost three—fourths is done before reaching
middle age. The males far surpassed the females in
weight. The control males not only reached their maxi~
mum weight at an earlier age than the exercised males,
but also greatly excelled them. The exercised rats
were more active, more alert and brighter in appearance
than the non-exercised ones, nevertheless, the control
rats lived longer than the exercised rats.

Brody (9) did a study on the effects of selective
breeding on activity and found that selective breeding
had no effect upon increasing the mean activity or raw
ducing the variability within the active strain of the
albino rat.

Browman (lO) controlled temperature and observed
the effects upon the spontaneous activity rhythms of the
albino rat. He succeeded in modifying the daily sponm
taneous activity rhythm and onset of oestrous of normal
animals by daily variation of light and dark regardless
of environmental temperature and by daily variation in
temperature keeping the light conditions constant. And
Brobeck (2) observed, that when temperature was low the

average activity was high, but when the temperature was

female rats being much more active than the male. In
addition, Slonaker showed that during the prime of life
the activity is confined principally to night, and rest
to day time. Of the whole amount of work done during the

life time, almost three-fourths is done before reaching

middle age. The males far surpassed the females in TE
weight. The control males not only reached their maxi- E
mum weight at an earlier age than the exercised males, 5

but also greatly excelled them. The exercised rats

 

were more active, more alert and brighter in appearance is
than the non—exercised ones, nevertheless, the control
rats lived longer than the exercised rats.

Brody (9) did a study on the effects of selective
breeding on activity and found that selective breeding
had no effect upon increasing the mean activity or re—
ducing the variability within the active strain of the
albino rat.

Browman (lO) controlled temperature and observed
the effects upon the spontaneous activity rhythms of the
albino rat. He succeeded in modifying the daily sponw
taneous activity rhythm and onset of oestrous of normal
animals by daily variation of light and dark regardless
of environmental temperature and by daily variation in
temperature keeping the light conditions constant. And
Brobeck (2) observed, that when temperature was low the

average activity was high, but when the temperature was

“J.

raised, the average daily spontaneous activity decreased.
And, after bilateral destruction of the lateral group of

amygdaloid nuclei in rats showed that the activity of all
the operated animals, whether normally active or inactive,
showed a marked decrease, but that there were no signifi~
cant changes in food intake and body weight in comparison

with those of controls.

The Effects of Animal Handling on Growth

 

 

McClelland (33) and Weininger (A?) and others, in :
their animal work, support the theory that tactile stimuw
lation during handling in the form of gentling enhances
the vitality of the albino rat in the following specific
ways: a) increasing body weight and skeletal length under
ad libitum feeding conditions, b) increasing ambulatory
activity and thereby decreasing emotionality, and c) de~
creasing organic damage under severe stress in the form
of immobilization and food and water deprivation.

Greenman and Duhring (18) observed that handling of
animals reduces emotionality and that the animals become
tame and respond to the experimental procedures instead
of "freezing." Gertz, (16) on the other hand, showed
that there was no significant differences in emotionality
from formal handling between the groups when tested as
adults. Gertz concluded that there was no evidence for
a "critical age” at which handling is effective in terms

of later behavior.

Hall (22) from studying the relationships between
external stimulation, food deprivation, and activity con-
cludes that increased environmental stimulation results
in an increase in activity, and the addition of the food-

deprived state increases it still more.

The Effects of Exercise on Growth
There appears to be some controversy about the effects
of exercise on growth. Donaldson (15) states that "exercise

improves growth" when allowed as voluntary exercise, but

 

that the over-exercising of rats at young age will retard
growth. Price and Jones (35) report the effects of forced
exercise on the growth of young rats exercised daily in
drums turned by a motor. No special diet was administered.
In results they found clear variation of growth between

the resting rats and the exercised. The resting group
showed greater rate of growth when expressed in the average
percentage weightS--whereas the exercised rats lagged be~
hind. Measurements of body weights, body length, weight
of humerus, length of humerus, weight of femur, and length
of femur were given for 6 animals from controls (resting)
and exercised animals--on every single count the controls
were larger, longer and heavier. ”All the weights and
measurements of the exercised rats are less than the cor~
responding weights and measurements of the control rats."
Price and Jones concluded that exercise retards the in—

crease in weight and length of the body and of the skeleton.

Kimeldorf and Baum (27) report on alternations in
organ and body weight of rats following daily exhaustive
exercise, and exposure to x-rays. In their experiment
they observed that body weight and weight of the gluteus
maximus, kidney, and thymus were reduced while adrenal
weight was increased in animals subjected to daily ex-
haustive exercise or exposed to x-rays. Spleen and heart
were enlarged on particular days of sacrifice following
exercise, whereas they were reduced with most doses of
x-ray. Testis and pituitary weights were decreased by

irradiation but not by exhaustive exercise.

The Effects of Diet on Activity and Growth

There appears to be a lot of experimental work done
on various kinds of diets and how they affect the general
health and well-being of laboratory animals., The refer-
ences selected here were just the ones dealing with growth
and exercise and how they are affected by diet, vitamin
content in diet, and the denial of food or water.

Deuel and others (14) found that ad libitum feeding
to rats of diets varying in fat content from 5 to 50 per
cent resulted in better growth, greater physical capacity,
and better reproductive and lactation performance. Opu
timum growth was observed on diets containing 20 to 40
per cent fat. Carpenter and Mayer (12) observed that

the yellow mice gained the most weight on high fat diets,

10

but if the animals were given a possibility to exercise
in activity cages, they lost weight.

Slonaker (39) studied the effects of vegetable diet
on the spontaneous activity, rate of growth, and longevity
of albino rats. The rats fed on vegetable diet plus aniw
mal food were much more active and did more work in sponm
taneous activity cages than the vegetarians during their
lifetime. He further observed that the vegetarian rats
aged much earlier in life, and that their total growth
was greatly retarded.

Mann and others (28) related their experimental
findings to the develOpment of obesity. Their study
showed that the doubling of the caloric supply of young
men did not increase the serum lipoproteins and choles~
terol levels so long as the surplus energy was expended
as heat and muscular energy. These findings probably
reflect the benefits of physical activity.

Smith and Conger (43) suggested that differences in
spontaneous activity of albino rats are due to d_etary
differences. In their experiment they fed one group a
diet high in fat, the other group high in protein, and
the third group a well-balanced diet. All the animals
used in the study were litter-mates. The results showed
that as much as 56 per cent of calorific value of the
food may come from fat, and spontaneous activity be main«

tained at a normal level, but if the diet is 72 per cent

ll

of fat calories it depresses the activity a little.
However, if the diet is 50 per cent animal protein,
it induces a marked increase in activity.

Slanetz (38) evaluated diets for.laboratory animals
in respect to protein, fat, certain minerals, vitamin A

and a number of the vitamin B fractions. He found

I .I

variations between manufacturers claims and his findings
on vitamin A content in diets. He states that the diets

investigated are apparently adequate in respect to vitamin

 

B fractions, but feels that rat diets may be somewhat low
in fat and too low in vitamin A for good liver storage of
that vitamin.

Keith and Mitchell (26) tried the effects of muscular
work with rats on vitamin requirements. The results in~
dicated that the amount of exercise did not uniformly
affect the well-being of the rats on complete rations or
of those on rations lacking in vitamin B. They indicated
that exercise did distinctly hasten the appearance of the
symptoms of vitamin A deficiency, and hasten the death of
young animals receiving insufficient amount of this vita-
min. This study indicates a rather clear evidence that
muscular exercise increases the demand for vitamin A.

Guerrant and others (20) using young rats showed that
less food was consumed, smaller increases in body weight
were made, and less severe symptoms of vitamin A deficiency

develOped when the animals were forced to exercise in a

12

rotating wheel, than when permitted to exercise volun—
tarily, or when confined in the usual type of cage. This
study showed that animals which were forced to exercise
exhibited the greatest efficiency of food utilization,
they voided the greatest number of fecal pellets, indi-
cating a beneficial effect of exercise on intestinal
mobility. Furthermore, it was shown that animals, which
were exercised voluntarily, and which received daily al-
lotments of vitamin A, were more active physically than
were the litter-mates which did not receive the vitamin
supplement.

Beznak and others (7) and Beznak (6) in their studies
on growth and food consumption of albino rats exercised
in revolving drums found that the growth of inactive rats
on a low vitamin B1 diet was proportional to the amount
of fat in the diet. When the latter was reduCed to 3 per
cent, the growth becomes stunted. Forced exercise caused
resumption of growth in the low fat diet group and slowing
of growth in the high fat diet groups. On return to in—
active life growth in all groups quickened. It was also
observed in this study that water consumption during exer—
cise period was inversely proportional to the fat content
in diet and that it increased in later stages of exercise.

Guerrant and Dutcher (19) in their conducted experi-
ments with carefully selected groups of young rats, yielded

data which demonstrate that vitamin B1 requirement of these

f-l
I
Rik,

animals, as indicated by the time required for the ces_
sation of growth and for the development of paralytic
symtoms, is increased by physical exercise. In the ani-
mals receiving no vitamin B1, increased exercise resulted
in less food consumption, less growth, early develOpment
of paralytic symptoms, and the elimination of a greater
number of fecal particles. During the first 4 weeks of
the test period, those animals which were permitted to
exercise voluntarily were more active physically while
not receiving vitamin B1 than were the animals receiving
the vitamin B1. However, the physical activity declined
during the latter stages of vitamin B1 depletion.

Sheer and others (36) studied the physical capacity
of rats in relation to energy and fat content of the diet.
In their work they describe methods for the determination
of body specific gravity, and capacity for exhaustive work
in rats. The specific gravity determination was based
upon determination of body volume by displacement. They
determined the physical capacity by measuring the duration
of a swim to exhaustion with regularly increased work load.
No significant correlation between specific gravity and
fat content from O to AC per cent were fed ad libitum,
physical capacity of the animals increased. When caloric
intake was severely restricted, physical capacity decreased

markedly.

14

ThewEffects of Exercise Upon the Food Intake

This has been the most controversial issue in the
past, and for that matter, still is. The general belief
is that exercise directly affects the food consumption.
Most research and experimental work show just the Opposite,
Simply, that there is no direct relationship between in-
crease in food intake and mild activity.

Thomas and Miller (44) studied the influence of
treadmill running on food intake, body weight and spon—
taneous activity in male albino rats. The initial re-
Sponse to gradually increasing exercise load was a de-
crease infood intake and body weight on exercise days,
with compensatory increase in both factors on week-ends
(rest days). By the time the exercise was increased to
one mile per day, food intake on exercise days had re-
turned to normal or slightly above, and food intake on
rest days was significantly elevated. During the re-
covery period following the exercise, food intake re-
mained elevated despite the absence of a significant
weight deficit. Overnight Spontaneous activity was
strikingly reduced in exercised rats, with only partial
restoration on rest days, and with gradual return to
normal during the post—exercise recovery period.

Andik and others (3) studied the effects of exercise
on food consumption and selection. Animals were offered

free choice of 3 foods containing 1/3 standard mixture

15

and 2/3 starch, casein and lard, respectively. The rats
ran from 8 to 9 hours daily at 750 meters per hour and

it was observed that the consumption of the starch food
diminished, intake of fat rich, and especially of the
casein rich food, increased markedly. The bulk of the
food consumed remained practically unchanged, the change
over from starch to fat furnishing the surplus calories.
The termination of exercise was followed by return to pram
exercise levels of intake of all 3 foods. -

Mayer and others (31) observed, that when mature
rats accustomed to a sedentary existence were exercised
in a treadmill for increasing daily periods, no increase
in food intake was noted for low duration of exercise,

20 minutes to 1 hour, but the food intake actually de—
creased slightly but significantly, as did body weight
also. With longer durations of exercise (1 to 6 hours)
food intake increased linearlly and weight was maintained.
For very long duration, animals lost weight, food intake
decreased and the animals’appearance deteriorated.

Tribe (45) maintained 3 groups of 8 hooded Lister
rats on a self-selection system of feeding and each group
was submitted to 3 different levels of exercise. It was
observed that at no time did the voluntary caloric intake
vary with the amount of exercise undergone. Instead, the
rate of body-weight increase slowed down as the amount

of exercise increased.

l6

Mayer and others (32) extended their experimental
studies on work and food intake relationships to man.
They established relation between caloric intake, body
weight, and physical work in a group of 213 mill workers
in West Bengal, India. The workers fit into a wide range
of physical activity, from sedentary to very hard work.
It was found that caloric intake increases with activity
only within a certain zone. Below that range, a decrease
in activity is not followed by a decrease in foOd intake,
but rather by an increase. Body weight is also increased'
in that range. The picture presented here on humans is

similar to that found in experimental animals.

ThefiRole of Exercise in Weight Control

Mayer and others (31) in order to gain better insight
on the effects of exercise on body weight studied non-
obese and genetically obese mice. When the sedentary
animals were exercised on a treadmill for 1 hour a day,
‘the exercise did not affect the weight of the normal mice
but considerably decreased the weight gain of the obese
mice, even though the latter responded to exercise with
increased food intake. The importance of considering
exercise, as well as food intake, in the weight control,
is well illustrated by these results.

Johnson and others (25) compared two groups of high
school girls, 28 obese subjects and 28 non-obese individuals

of similar age, grade and height in regard to maturation,

17

food intake and activity. The groups differed in matura—
tion the obese girls showing advanced development, that
is, earlier deceleration of growth in height. Both groups
were found to be relatively inactive, but the obese girls
being significantly more so. When caloric intakes and

activity indices were compared to determine the energy

___E- .33"

factors in the develOpment and maintenance of obesity,
on statistical basis, it was observed, inactivity was

much more important than "overeating". In fact, the

 

caloric intake of the obese group was significantly lower J
than that of the non—obese girls, with the relatively

greater energy balance being consequently supplied by

inactivity.

Passmore (34) theorizes, that "there is no doubt
that in physically active persons appetite balances with
remarkable precision, the energy values of the food eaten
and the work performed.” He performed a study of food
intake on a dog. This animal within a short period of
exercise did not increase the appetite above the seden-
tary level, but some slight loss of weight occurred.

The results are thus in line with a number of other
studies and support their suggestion that a certain mini-
mal level of physical activity is necessary for the appe—
tite to function with precision and to adjust food intake

to the requirements.

18

"I am convinced that inactivity is the most impor-
tant factor explaining the frequency of 'creeping‘ over-
weight in modern western societies." These words belong
to Jean Mayer, who has done considerable experimental

work to support his statement. Mayer (29) studied in

 

detail the food intake and the daily schedule of over- ‘E
weight children. He fOund that they differed from the if
normal, not in that they ate more but that they were
extraordinary inactive. It was noted that the obese

"children just sat all their waking time. The role of A

physical inactivity in the development of the overweight
was indicated by the fact that the overweight children
who attended camps, where physical exercise was compul-
sory, lost weight during their summers. Mayer concludes
that "the lack of physical activity is a major factor

in the develOpment of obesity in the adult as well as

in the field-—moderate exercise does not increase food
intake, but does reduce body weight." Green (17) also
traced the cause of obesity in human adults to a sudden
decrease in activity.

Cecil and Loch (13) have this to say about over-
weight problems, "weight reduction through caloric re—
striction alone is rarely successful, additional measures
such as psychotherapy as well as physiological therapy are
needed." Guyton (21) states that forced exercise is often

an essential part of the treatment of obesity. Along with

the previous thoughts Bard (4) suggests that emphasis
should be placed on both exercise and diet-efithe store
of body fat can be increased or decreased only if the
energy intake is greater or less respectively, than the
energy expenditure. The fact that foOd intake is not
the only factor involved in overweight is indicated by Hi
Beaudoin and Mayer (5) in their work, where they found i
that the average caloric intake of obese women does not

contain significantly higher proportion of either car—

"in

 

' bohydrate or fat than does that of their normal weight
controls.

Whalley and others (48) in their experiments forced
animals to swim with additional weights amounting to 2.5
per dent, at other times forced them to swim without the
weights. This study shows that rats offered a diet of
fixed composition ingested less food and water during
periods of enforced daily exercise than during periods
of relative_rest. It was conducted that physical stress
(such as employed in this study) depressed in rats the
desire to eat and drink.

Brobeck's (8) experimental work suggests that food
intake, activity and temperature regulation in the rat
are in a certain sense independent variables upon which
weight gain depends.' Brobeck further theorizes that since
the regulation of body temperature, of activity, and of

food intake is disturbed in animals with hypothalamic

lesions, the hypOthalamuS may be the level of the central
nervous system responsible for the control of every exchange.
Concerning the origin and mechanism of obesity in con-

trast to the over-simplified overeating theory, Mayer (30)
theorizes that a number of different genes, probably working
by different mechanisms, lead to obesity, that hormonal fac- 7
tors are implicated, and that exercise plays a role of dif—
ferent importance in different obesity syndromes.

Finally, the role of exercise in the control of body

 

weight should not be overlooked. The teachers of physical I
education should constantly keep the importance of their

job in this area in their minds. The possibility that the

importance of physical activity has not been stressed enough

in the past will always exist. Burt and Blyth (11) state

3 implications on weight control:

1. Body weight can be maintained or reduced by a
considerable reduction in caloric intake.

2. Body weight can be maintained or reduced by
vigorous daily exercise, with no reduction in caloric
intake.

3. Body weight can be maintained or reduced by
only a small reduction in caloric intake and a corresponding
amount of daily exercise.

Thus, while diet may do the job temporarily, a more
satisfactory way of weight control is through regular

activity combined with sensible eating habits.

CHAPTER III
METHODS OF PROCEDURE

Introduction

 

The present study was undertaken to determine the
effects of forced exercise, Spontaneous activity, and
sedentary activity upon the food intake and body weights
of young male albino rats. More specifically,this ex—
periment was undertaken to determine 3 things:

1. The effects of forced exercise, spontaneous
activity and sedentary activity on food consumption and
body weights of young male albino rats.

2. The effects of spontaneous activity on food con-
sumption and body weight of young male albino rats fol-
lowing a period of forced exercise and sedentary activity.

3. To relate individual rat's spontaneous activity
level with its food consumption and body weight following
a period of forced exercise and sedentary activity.

It is necessary to state here that these same animals
were used for another study that was being run concurrently.
Therefore, a lot of information reported here will also
reflect on the other study, nevertheless, it is felt that
in order to present the picture in the clearest possible
way, it is necessary to bring out that information. The

other experiment was concerned about the effects of forced

[‘0
h)

exercise upon the spontaneous activity. The length of
this study was 140 days. The first 35 days will be re—
ferred to as the Treatment Phase while the remaining
105 days as Post-Treatment Phase.

The animals assigned to Group I were sedentary for
the first 35 days but were allowed spontaneous activity
for the remaining 105 days. Those animals which were
assigned to Group II were allowed spontaneous activity
during the entire experiment. The animals assigned to
Group III in addition to spontaneous activity were forced
to swim daily with the overload method for the first 35
days, for the final 105 days these animals were allowed
spontaneous activity only. The animals from Group II
were kept as Controls, whereas the animals from Groups
I and III were held as Experimental Groups.

The food consumption was measured for each animal
for the entire experiment. The average daily food con-
sumption per animals per group was determined 11 times
through the Treatment Phase and 24 times throughout the

Post-Treatment Phase.

Subjects and Equipment

Thirty young male albino rats of the Sprague—Dawley
strain were used for the study. These animals were 29
days old when they were received in the laboratory, 33

days old at the start of the experiment, 68 days old at

IO
U.)

the end of the Treatment Phase (first 35 days) and 173
days old at the termination of the experiment. These
animals were all born on the same day but were not
litter-mates.

The animals were assigned to groups by means of
random selection. Fifty-four animals were received and
were placed in individual sedentary cages. Each cage
was numbered. Using a random number table, selection
into groups of 10 rats each was made, and the animals
were placed into the proper cage for their respective
group. Each cage was numbered for the group and the
individual. The animals assigned to Group I were housed
in individual rectangular cages that measured approxi-
mately 10 inches long, 8 inches wide, and 7 inces high.
Those animals which were assigned to Groups II and III
were placed into individual spontaneous activity cages.
These cages consisted of a 4 x 4 hardware cloth rectan—
gular enclosement that was approximately 12 inches long,
5 1/2 inches wide, and 5 inches high. These cages were
attached to a partition wall that measured 18 inches
high and 18 inches wide with a 1/16 of one inch thick-
ness. On the reverse Side of the partition was a Spon-
taneous wheel that measured approximately 45 inches in
circumference and was 5 inches wide. An animal would
have to run 1408 revolutions to cover a mile. The

outer surface of this wheel was made of plexiglass while

 

the running surface was made of 4 x 4 hardware cloth.
The spontaneous wheel was connected to the partition
wall by means of a bicycle axle which was free wheeling.
A hole that measured 3-3/4 inches x 3—1/4 inches was cut
in the partition wall between the rectangular cage and
the wheel giving the animal free access to either por~
tion of the cage.

Animals housed in the spontaneous activity cages
entered the spontaneous wheel at will and made it turn
by running. Many of the animals jumped high on the wheel
to start it in motion and then they ran on it much like
running on a tread mill. The wheels were designed so that
the animal could run in either direction with the same
effort and the revolution counters that were attached
to each cage counted each complete revolution. A mechani-
cal revolution counter was mounted on the rectangular
cage side of the partition and its arm operated from an
extension rod which attaches to a circular disc that was
screwed onto the bicycle axle. This counter was cali~
brated to measure each revolution that the wheel makes
and works equally well in either direction.

Water was supplied to the animals in all three groups
by means of an inverted bottle which was fastened to the
outside of the rectangular cage. A metal tube extended
from the bottle into the cage and the animals received

water by placing their mouth on the end of the tube.

A marking code was designed for the identification
of individual rats. The code consisted of various pat-
terns of circular and triangular holes made in the ears
of the rats. The circles were made with a punch and the
triangles with scissors. There was very little bleeding
and no infection from this marking.

The temperature in the laboratory was not controlled,
but was recorded daily with a Daystrom—Weston temperature
recorder. The relative humidity was kept low by the use
of a dehumidifier.

Small plastic containers were used for collection
and transfer of food. The containers were specially
marked for each animal. All food was weighed by the use
of the Cent 0 Gram Triple Beam Balance. Fine screen
plates were used to collect all the food that fell out
of the spontaneous wheel or from the rectangular cage.
These screen plates were placed over the trays containing
the woodshavings and allowed all the urine and water to
be absorved by the woodshavings but leaving the food on
top of the plate covering the tray.

All animals were fed Wayne Lab Blox for Rats, manu-
factured by the Allied Mills of Chicago and were allowed

to consume all the food they desired.

Training Program
The entire experiment took 140 consecutive days.

These 140 days were divided into two parts, the Treatment

26

Phase (first 35 days) and the Post—Treatment Phase (last
105 days). Within the Treatment Phase the individual
daily average food consumption was determined 11 times.
During the Post-Treatment Phase of the experiment the
food intake of each animal was obtained 24 times. During
the third and fourth phases, 8 times. Throughout the
Treatment Phase, the animals from Group I were kept in
individual sedentary cages while the animals from Group
II and III were kept in spontaneous activity cages. The
animals from Group III were forced to swim for 30 minutes
daily using the overload method. Two per cent of the
animafs weight was applied to his tail by water-proof
adhesive tape. These animals swam in a sink which was
lined with plexiglass. The water used for swimming was.
kept between 35 and 37 degrees Centigrade.

Since only six of the spontaneous exercise cages
contained microswitches to measure the intensity of
Spontaneous activity for the other study, a rotation
order was developed whereby each rat from the control
group and forced exercise group was placed in these
cages for 24 hours every 5 days. At the end of the 24
hour period, the rats were removed from these cages
and were placed in the cage of the rat which was to take
his place in the microswitch-equipped spontaneous ac—
tivity cage. Prior to this move, all of the food was

removed from the cages of the animals involved and each

animal from all three groups was weighed. The animal's
water bottle was moved with him as was his identification
tag that was taped to the outside of his cage. This
moving procedure shuffled the rats about the room so that
they would be exposed to the varying environments in all
of the locations within the room. The sedentary cages
were shuffled around every second day so that the animals
would not be on the same shelf for three days in a row.
During the Post—Treatment Phase of the experiment,
all animals from each group were housed in the spontaneous
activity cages. The ten sedentary animals were marked
for identification purposes. A new rotation order was
established to use the spontaneous activity cages equipped
with the microswitches. Each animal was placed in one of
these special Spontaneous exercise intensity cages for
24 hours every five days. Tflmztotal.spontaneous wheel re—

volutions were recorded each day.

Testing Techniques

 

Each day starting at 2 p.m. all the animals were
weighed. The food was then collected from the animal‘s
cages that were scheduled to be tested for that day.
Actually, the food was picked up from the six spontaneous
activity cages equipped with microswitches as well as
from the cages from which the animals were transferred
to these specially equipped cages. Since there were

three cages used for both, the spontaneous activity group

28

and the forced exercise group, to measure the intensity
of the running, the food was measured for six animals
from each group. The food was collected in small
plastic containers marked for each animal. The same
procedure in the food collection was followed with the
Sedentary Group animals.

The new food was weighed and prepared so that it
could be placed in the cages right after the animals
were rotated according to their rotation order for
that day. Each animal was given from 30 to 40 grams
extra food to avoid any possible shortages.

Specially built fine, screen plates were placed
over the trays containing the woodshavings so that no
food would be lost. Each smallest piece of leftover
food was always collected from the animal's cage and
the screen plates.

The weighing of the animals, collecting the food
from the cages, weighing the collected food, weighing
the new food, placing the new food in the cages and
filling the water bottles took approximately from 2-1/4
to 2-3/4 hours. All this was done in one block of time.
On the days when the waste trays were cleaned, it took
an additional hour of time.

The same procedures were followed for the Post-
Treatment Phase of the experiment. Instead of weighing
the food for 18 animals each day as in the first half,

now the food was handled only for 12 animals daily.

 

CHAPTER IV
RESULTS

This experiment was undertaken to determine the
effects of various levels of exercise upon the food
consumption and body weight of young male albino rats.
The experiment consisted of two parts, a treatment phase
(35 days in length) and a post-treatment phase (105
days in length). The various treatments applied to
each group were described earlier. Individual rat
number nine of the control group was eliminated because
of extreme deviate activity and a large daily food con—
sumption in the post-treatment phase. This animal's
activity accounted for almost 50 per cent of the total
spontaneous activity revolutions of the control group.
This animal was also considerably lighter than any other

animal in that group.

Food Consumption

 

Figure 1 shows the animal daily average food con-
sumption. The sedentary and the forced exercise rats
at the start of the experiment averaged approximately
the same amount of food per day. The Controls were
slightly below the other two groups in their food in—

take per day. The average daily food intake for the

3O

   
 

 

 

 

 

July ll-IZ- I
“J I3‘I5" ‘s‘\
2 16484 '\
35 l9-2lr-
.. 22'24' ‘ ........ _ I SEDENTARY
5 252;- ‘ ‘\ —--. CONTROL
2 28- -
.— = ........ ‘ FORCED EXERCISE
<1 3I-Aug.2~ A
uJ ‘
E Augusi 3-5~ A
6-8-
940" a END OF TREATMENT PHASE
ll-I4 /,t
I5-l8- ‘ .
~ I....“-..
19-23- ‘\ “,,.:::.-A
24-27_ _, J
28-Septl+ ,fl’
September 2-5- Q
e-Io~ R
II-l4- ’Jfi
l5-I9'- 0(\
\ .
20-24- 3‘
3’1 25-29— :A
<1! .30-0c1.4 - ’)-‘
’ .-'
0- October 5-9- .C; A .
,_ Io-I3— ‘0 A
5 I4-I7- ‘\ A.
E Ia-ZI- ,3". Sis-A
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m 22-25- Q’;\
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*.‘ 26-29— A?”
(*3 30— Nov. 2 - 0:5...
8 November 3'6 - :‘2‘
7-10L ,0’ K
”-14- U:
548— Q "A
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l l l l I I I I J
l8 IS 20 an 22 23 24 25 26
GRAMS
Figure 1. Average FQfid Tonsun;tl-n Per Grggp
Per Test Ieriud

31

treatment phase came out about equal for all three
groups. The Sedentary Group averaged daily 21.290
grams per rat, the Controls 20.202 grams, and the
Forced Exercise Group 20.466 grams per rat.
Statistically there were not any significant
differences between the groups in animal food consump-
tion for the Treatment Phase. Based on twenty-nine
rats and three treatments, the allocation of degree of
freedom were the following for food consumption during
the treatment phase: Total~e28, Treatments--2, and
Error-e26. The variance due to treatments was not
significant at the 5 per cent level (F = 1.59). The
results of the analysis of variance are shown in

Table I.

.TABLE I

ANALYSIS OF VARIANCE OF DATA FOR FOOD CONSUMPTION
DURING THE TREATMENT PHASE

vhf

 

Source of Sums of Mean

Variation D.F. Squares Square F F.05
Total 28 57.236

Treatments ”'2 6.227 3.11354 1.59 3.39
Error 26 51.009 1.9619

'7 w

The daily average food consumption for the post-
treatment phase for the three groups came out as follows:

Sedentary Group 23.321 grams per rat, Controls 22.934

grams per rat, and Forced Exercise 22.934 grams per rat.
When the daily averages of the post-treatment phase are
compared with those of the treatment phase, the daily
food consumption per rat has increased by all three groups.
Statistically there were not any significant dif-
ferences between the groups in animal food consumption
during the postetreatment phase. Based on twenty—nine
rats and three treatments, the allocation of degrees of
freedom were the following for the food consumption for
the post-treatment phase: Tota1—-28, Treatments~—2, and
Error-~26. The variance due to treatments was not sig-
nificant at the 5 per cent level (F = 0.70). The results
of the analysis of variance are shown in Table II. The
means of the three groups were tested at the 5 per cent

level through the use of the Duncan Test for Mean Values.

TABLE II

ANALYSIS OF VARIANCE OF DATA FOR FOOD CONSUMPTION
DURING THE POST-TREATMENT PHASE

 

 

Source of Sum of Mean

Variation D.F. Squares Square F F.05
Total 28 108.207

Treatments 2 5.584 2.782 0.70 3.39

Error 26 102.623 3.9470

 

 

 

33

Animal Weights

At the start of the experiment the Sedentary and
the Control Groups were averaging 107 grams per rat. The
Forced Exercise Group averaged 105 grams per rat. Figure
2 shows the weight gain patterns for the entire experiment.

At the end of the Treatment Phase the Sedentary
Group averaged 320 grams per rat, the Control Group 296
grams, and the Forced Exercise Group 280 grams per rat.
From this it can be seen that the Sedentary Group gained
approximately 213 grams per rat in the 35 days of seden-
tary activity. The Control Group gained approximately
189 grams per rat while running at their own desire in
the spontaneous activity wheel for 35 days. The Forced
Exercise Group, while swimming daily for 30 minutes with
the overload method in addition to the running in the
spontaneous activity wheels, gained 174 grams per rat
for the same 35 days.

Statistically, there was a significant difference
between the groups in the animal weights at the termina-
tion point of the Treatment Phase. Based on twenty-nine
rats and three treatments, the allocation of degrees of
freedom were the following for the animal weights for
the Treatment Phase: Total--28, Treatments--2, and Error--
26. The variance due to treatments was found to be signi-
ficant both, at the 5 per cent level (F.05 = 3.37) and at
the 1 per cent level (F.01 = 5.53). The results of the

analysis of variance are shown in Table III.

GRAMS

460

440

420

400

380

360

340

320

280

260

240

I40

I20

I00

 

34

 

 

 

 

f
P
)-
r-
h.-
.. -—-I SEDENTARY
, f: w ---0 CONTROL
L - g;
4‘ I ...... ‘ FORCED EXERCISE
. ,- at
— .—
Z
UJ
2
'—
— <1
LU
" (I
'—
- u.
. o
_ D
2
.° LIJ
I- t.
P
l n l L l l l 1 l L L 4 l J l l l .L L n l A J l l 1 J
S IO 20 30 4O 50 60 7O 80 90 IOO IIO IZO I30 I40
DAYS
ulé‘UI‘E 2 dy Irieiqiit Par ~f_}rf?u

35

TABLE III

ANALYSIS OF VARIANCE OF DATA FOR THE ANIMAL WEIGHTS
DURING THE TREATMENT PHASE

 

 

-.———

 

Source of Sum of Mean

Variation D.F. Squares Squares F F.05 F.01
Total 28 16,217.95

Treatments 2 7,563.99 3782.00 11.36 3.37 5.53
Error 26 8,653.96 332.84

 

 

To Test Group Means at the 5 Per Cent Level:

 

 

174.30 188.80 212.80
Forced Control Sedentary
Exercise Group Group
Group

Using Range Table with 26 D.F.

 

2 3
2.91 3.06 s; = 332.8h l.+.l +.;
(2) (3) 3 10 9 10

17.07 17.95

To Compare Sedentary Group with the other two groups:
212.80 194.85 is greater than 188.80 so, therefore,
~1£.35 the sedentary group mean is significantly
19 . 5 greater than the other two groups at the
5 per cent level.
To Compare the Control Group with the Forced Exercise Group:

188.80 171.73 is less than 174.30 so, therefore,
-lZ.07 the Control Group mean is not significantly
171.73 greater than the Forced Exercise Group.
At the termination of the experiment (140 days) the
Sedentary Group was averaging 465 grams per rat, the Control

Group 458 grams per rat. Thus, since the end of the

Treatment Phase the Sedentary Group had gained about
145 grams per rat, the Controls 162 grams per rat, and
the Forced Exercise Group about 159 grams per rat.
Statistically there were not any significant dif;
ferences in animal weights between the groups from the
end of the treatment phase to the end of the post—
treatment phase. Based on twenty-nine rats and three
treatments, the allocation of degrees of freedom were
the following for animal weights during the post-
treatment phase: Total—-28, Treatments--2, and Error--
26. The variance due to treatments was not found to
be significant at the 5 per cent level (F = .92). The

results of the analysis of variance are shown in Table IV.

TABLE IV

ANALYSIS OF VARIANCE OF ANIMAL WEIGHT DIFFERENCES
FROM THE END OF TREATMENT PHASE TO
THE END OF THE EXPERIMENT

 

 

Source of Sum of Mean

Variation D.F. Squares Square F F.05
Total 28 24,842.98

Treatments 2 1,642.52 821.26 .92 3.37
Error 26 23,200.46 892.32

 

 

Over the entire experiment (140 days) the Sedentary
Group gained 357 grams per rat, the Control Group gained
351 grams per rat, and the Forced Exercise Group gained

334 grams per rat.

 

Statistically there were not any significant dif~
ferences between the groups on the total weights gained
for the entire experiment. Based on twenty—nine rats
and three treatments, the allocation of degrees of free~
dom were the following for the animal weights for the
entire experiment: Tota1--28, Treatments—;2, Error—~26.
The variance due to treatments was not found to be sig-
nificant at the 5 per cent level (F = 1.04). The results

of the analysis of variance are shown in Table V.

TABLE V

ANALYSIS OF VARIANCE OF DATA FOR ANIMAL WEIGHT
DIFFERENCES FOR THE ENTIRE EXPERIMENT

 

 

W T

 

Source of Sum of Mean

Variance D.F. Squares Square F F.05
Total 28 41,113.55

Treatments 2 3,032.97 1516.49 1.04 3.37
Error 26 38,080.58 1464.64

 

Spontaneous Activity

Along with this experiment, the amount and intensity
of spontaneous activity of the same rats was tested through
another study by Dale L. Hanson. (23) Spontaneous activity
was measured by the total number of wheel revolutions made
by the individual rats. The mean values for spontaneous

wheel reVolutions are given in Figure 3.

,I..l/

;:w;. peg acoahzeo>mm H6663 mdomdeCO mm mm:.
EmEEmF mfluloxm .085“. 2: 0532.0“. 230

mNN .NN 2N mON hm. mm. .0. nb. mm. hm. m3 :1 mm. MN. b: mo. —00_ 0m 0m ON Om Om O¢
_ _ _ . _ _ _ _ _ _ a _ _ _ a a 1 a a _ 4 _ q 00.

0‘. \.
......ICWHHHI. ."JU’I‘ I. {JO/Xi

000.000 o
oo’o‘coo'ooo. '

 

ICON

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...(si «......»

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... .... .. / 5

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I

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100m—

 

T 3265 638a

>50’COUmm o 0.....-
100V.

... Don.

 

 

_ b _ — P _ — p — _ _ _ b P p — r _ 00w—

 

IaauM

Suoummaa

39

The results showed that rats which were forced to
exercise by swimming with the overload method, increased
their total spontaneous activity significantly as measured'
by wheel revolutions during the post:treatment phase but
not during the treatment phase. The post:sedentary rats
had comparable total spontaneous activity with that of
control rats, but the total spontaneous activity of both
of these two groups was significantly less than that of

the forced exercise rats.

Discussion
The evidence presented indicated that in rats,
under the conditions of this experiment, the voluntary
food intake was not affected by forced exercise, nor
was it decreased when the forced exercise treatment
was terminated. Altogether, there were no significant
Idifferences between the groups in the average food con-
sumption throughout the entire experiment. Nevertheless,
the mean food consumption of the sedentary rats was
slightly higher than that of the forced exercise rats
during the treatment phase and also during the post-
treatment phase. The food consumption of the control
group was less than that of any of the other two groups.
The obvious drop in food consumption in the forced
exercise group as well as in the control group during
the treatment phase could probably be due to an increase

in room temperature. It is very interesting to note,

40

however, that if the temperature really was the deciding
factor here, it affected the food intake of the seden—
tary rats very little.

The daily food consumption of the sedentary rats
fell off quite considerably during the first stages of
the post-treatment phase when these animals were placed
on spontaneous activity for the first time. This sudden
drop was probably due to the opportunity to exercise and
also due to the new environmental conditions. Thus, it
could be assumed that the opportunity to run in the
spontaneous activity wheel, along with the new environ-
ment, temporarily depressed the rat's desire for the
food offered. This also was the only time in the en-
tire experiment when the sedentary rats did not gain
any weight for five consecutive days, and even when
they started to gain again, the rate of weight gained
per day was not as great as previously during the
treatment phase. The forced exercise group and the
control group were averaging approximately the same
weight until the twentieth day of the experiment. From
then on the rate of the daily weight gain of the forced
exercise rats started to decrease and continued so to
the end of the treatment phase. During the first part
of the post-treatment phase the rate of the daily weight
gain of the forced exercise rats was slightly increased,

but then, as these rats started to increase their

I:
I...)

spontaneous activity, the rate of the weight gain became
slower again. Nevertheless, the rate of the weight gain
during the post-treatment phase was slightly higher in
the control and forced exercise groups than in the
sedentary group. In the case of the forced exercise
rats, this could be an attempt to restore the body weight
deficit incurred during the forced swimming period, why
this is so with the controls, remains to be investigated.

The fact that spentaneous activity was depressed
in the forced exercise rats during the treatment phase,
but was greatly elevated following the termination of
the treatment phase, indicates that forced exercise at
early age does affect the degree of spontaneous activity
in adult male albino rats.

Finally, considering voluntary food consumption a
measure of desire for food, it can be conCluded that the
levels of exercise designed for this experiment did not
affect the rats food intake. The exercise did, however,

affect the rats body weight.

CHAPTER'V

SUMMARY, CONCLUSIONS, AND RECOMMENDATIONS

Summary

The purpose of this experiment was to determine
the effects of various levels of exercise upon the food
consumption and body weights of young male albino rats.
The experiment was divided into two major parts. The
first part consisted of thirty—five days and will be
referred to as the treatment phase. The second part
consisted of one hundred and five days and will be re-
ferred to as the post-treatment phase. The experiment
ran one hundred forty consecutive days.

Thirty rats of the Sprague-Dawley strain were
used for the entire experiment. Animals were divided
into three groups, each group consisting of ten rats.
The Sedentary Group was allowed sedentary activity for
the treatment phase but spontaneous activity for the
post-treatment phase. The rats in the Spontaneous
Group served as controls and were allowed spontaneous
activity throughout the entire experiment. The Forced
Exercise Group was forced to swim with the overload
method for the treatment phase and were allowed spon-

taneous activity for the entire experiment.

43

Individual rat body weight was determined daily for
the first seventy days, and every third day for the last
seventy days. Animal food consumption was measured for
each individual rat throughout the entire experiment.
The Spontaneous activity was measured daily for each
individual rat by means of revolution counters that
were attached to their cages. These data were recorded
daily.

Rotation order was developed so that each rat
would be in the microswitch-equipped cage once for a
24-hour period every five days. This was done because
of another study that was run concurrently.

The results indicate that when young male albino
rats are forced to exercise at young age they do not
gain as much weight as those animals that are restricted
from activity or allowed to exercise at their own desire
in the Spontaneous activity wheels. Even though the
activity was different for each group of animals, there
were no significant differences between the groups total
in their food consumption. Finally, the rats that were
forced to exercise by swimming at an early age ran a
significantly greater number of revolutions than either

the post-sedentary or control group rats.

Conclusions

 

1. Exercise does Significantly affect body weight.

2. Mild exercise does not increase food intake.

3. Forced exercise rats participate significantly
more in Spontaneous running.

4. Independent from the group, those animals
which do the least work gain the most weight and also
on the average eat more.

5. Independent from the group, those animals
which do the most work gain the least weight and Also

on the average-eat less.

Recommendations

 

1. A similar study Should be done over a longer
period of time to determine whether the animal weights
for all three groups would ever come together.

2. A Similar study should be done over a longer
period of time to determine whether the spontaneous
activity level of the forced exercise rats would ever
drop below that of the other two groups.

3. A similar study should be done with an ad—
ditional treatment phase in later stages of life to
determine the effects upon food consumption, body
weight, and Spontaneous activity.

4. A study should be done varying the overload
weights to determine the effects upon food consumption,
body weight, and spontaneous activity during the treatu
ment phase and during the post-treatment phase.

5. A similar study should be done on a carefully

selected active strain of rats.

lo.

BIBLIOGRAPHY

American Public Health Association, Inc. Care of
laboratory Animals. New York: American
Public Health Association, Inc., 1954.

Anand, Bal K., and John R. Brobeck. "Food Intake
and Spontaneous Activity of Rats with Lesions
in the Amygdaloid Nuclei," Journal of Neuro-
physiology, 15:421—430, September, 1952.

Andik, I., J. Bank, I. Moring, and Gy Szegvari.
"The Effect of Exercise on Food Intake and
Selection," Acta Physiologica Academiae
Scientiarum Hungaricae, 5:457-461, 1954.

Bard, Philip. Medical Physiology. St. Louis:
C. V. Mosby Company, 1956.

Beaudoin, R., and J. Mayer. "Food Intakes of Obese
and Non—obese Women," Journal of American
Dietetic Association, 29:29-33, January, 1953.

Beznak, A. B. L. ”Dietary Fat, Work, and Growth,"
Experientia, pp. 236-238, 1953.

Beznak, A. B. L., M. Beznak, and J. Hajdu. ”The
Growth and the Food and Water-Consumption of
the Testing or Exercising Albino Rat on Diets
Containing Various Amounts of Fat and Reduced
Quantity of Vitamin B1," Hungarica Acta Phy—
siologica et Biologica, l (2737:35, 1947.

Brobeck, John R. ”Effects of Variations in Activity,
Food Intake and Environmental Temperature on
Weight Gain in the Albino Rat," The American
Journal of Physiology, 143:1-5, January, 1945.

Brody, Elizabeth Graves. Genetic Basis of Spontaneous
Activity in the Albino Rat. Baltimore, Maryland;
The Williams & Wilkins Company, 1942.

Browman, Ludvig G. "Effects of Controlled Temperature
Upon the Spontaneous Activity Rhythms of the
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94:477-489, December, 1943.

11.

12.

13.

14.

15.

16.

17.

18.

19.

20.

21.

46

Burt, J. J., and c. S. Blyth. "Role of Exercise
in Weight Control," Journal of Health,
Physical Educationy_and Recreation, 32:23-24,
January, 1961.

 

 

Carpenter, K. J., and Jean Mayer. "Physiologic
Observations on Yellow Obesity in the Mouse,"
American Journal of Physiology, 193:499-504,
June, 1958.

 

Cecil, Russell L., and Robert F. Loeb. A Textbook
of Medicine. Philadelphia: W. B. Saunders

Company, 1955.

 

 

Deuel, Harry J., Emily R. Merserve, Evelyn Straub,
Cornelia Hendrick, and Bradley T. Scheer.
"The Effect of Fat Level of the Diet on
General Nutrition," Journal of Nutrition,

33:569'582: May, 1947'

Donaldson, H. H. The Rat--Data and Reference
Tables, 2nd ed., p. 179, 1924.

Gertz, Boris. I'The Effect of Handling at Various
Age Levels on Emotional Behavior of Adult
Rats," Journal of Comparative Physiological
Psychology, 50:613-616, December, 1957.

Greene, James A. "Clinical Study of the Etiology
of Obesity,: Annals of Internal Medicine,

12:1797-1803, May, 1939.

Greenman, M. J., and F. L. Duhring. Breeding and
Care of the Albino Rat For Research Purposes,
2nd ed. Philadelphia: The Wistar Institute
of Anatomy and Biology, 1931.

 

Guerrant, N. B., and R. A. Dutcher. ”The Influence
of Exercise on the Growing Rat in the Presence
and Absence of Vitamin B1," Journal of
Nutrition, 20:589-598, December, 1940.

 

 

Guerrant, N.B., R. A. Dutcher, and F. Chornock.
”The Influence of Exercise on the Growing
Rat in the Presence and Absence of Vitamin
A," Journal of Nutrition, 172473—484, May,

1939-

Guyton, Arthur C. Textbook of Medical Physiology.
Philadelphia: W. B. Saunders Company, 1956.

 

11.

l2.

l3.

14.

15.

16.

17.

18.

19.

20.

21.

46

Burt, J. J., and 0. s. Blyth. "Role of Exercise
in Weight Control," Journal of Health,
Physical Education, and Recreation, 32:23—24,
January, 1961.

 

 

Carpenter, K. J., and Jean Mayer. "Physiologic
Observations on Yellow Obesity in the Mouse,"
American Journal of Physiology, 193:499—504,

June, 1958.

Cecil, Russell L., and Robert F. Loeb. A Textbook
of Medicine. Philadelphia: W. B. Saunders

Company, 1955.

 

 

Deuel, Harry J., Emily R. Merserve, Evelyn Straub,
Cornelia Hendrick, and Bradley T. Scheer.
"The Effect of Fat Level of the Diet on
General Nutrition," Journal of Nutrition,

33:559-582. May. 1947.

Donaldson, H. H. The_Rat--Data and Reference
Tables, 2nd ed., p. 179, 1924.

Gertz, Boris. "The Effect of Handling at Various
Age Levels on Emotional Behavior of Adult
Rats," Journal of Comparative Physiological
Psychology, 50?613-616: December, 1957.

 

Greene, James A. "Clinical Study of the Etiology
of Obesity,: Annals of Internal Medicine,

12:1797-1803, May, 1939.

Greenman, M. J., and F. L. Duhring. Breeding and
Care of the Albino Rat For Research Purposes,
2nd ed. Philadelphia: The Wistar Institute
of Anatomy and Biology, 1931.

 

Guerrant, N. B., and R. A. Dutcher. ”The Influence
of Exercise on the Growing Rat in the Presence
and Absence of Vitamin B1," Journal of
Nutrition, 20:589-598, December, 1940.

 

 

Guerrant, N.B., R. A. Dutcher, and F. Chornock.
”The Influence of Exercise on the Growing
Rat in the Presence and Absence of Vitamin
A," Journal of Nutrition, 17 473—484, May,

1939.

Guyton, Arthur C. Textbook of Medical Physiology.
Philadelphia: W. B. Saunders Company, 1956.

 

22.

23.

24.

25.

26.

27.

28.

29.

30.

31.

Hall, John F. "The Relationship Between External
Stimulation, Food Deprivation, and Activity,"
Journal of Comparative and Physiological
Psychology,449:339-34l, March, 1955.

 

Hanson, Dale L. "The Effects of Forced Exercise
Upon the Amount and Intensity of Spontaneous
Activity of Young Male Albino Rats." Unpublished
Ph. D. Thesis, Michigan State University, East
Lansing, 1962.

Hetherington, A. W., and S. W. Ranson. "The Spon-
taneous Activity and Food Intake of Rats with
Hypothalamic Lesions," American Journal of
Physiology, 136:609-617, March, 1942.

Johnson, Mary Louise, Bertha S. Bruke, and Jean
Mayer. "Relative Importance of Inactivity
and Overeating in the Energy Balance of Obese
High School Girls," American Journal of
Clinical Nutrition, 4:37-44, January—February,

1956.

Keith, M. H., and H. H. Mitchell. HThe Effect of
Exercise on Vitamin Requirement," American
Journal of Physiology, 65:128-138, June, 1923.

Kimeldorf, D. J., and S. J. Baum. "Alternations in
Organ and Body Growth of Rats Following Daily
Exhaustive Exercise, X-Irradiation, and Post—
Irradiation Exercise," Growth, 18:79-96,
May, 1954.

Mann, George V., Katherine Teel, Olive Hayes, Ann
McNally, and Dorothy Bruno. "Exercise in the
Disposition of Dietary Calories," The New
England Journal of Medicine, 253:349-355,
September, 1955.

Mayer, Jean. "Exercise Can Keep Us Fit,” Journal
of HealthnghySical Education & Recreation,
27:12+, September, 1956:

Mayer, Jean. "An Experimentalist's Approach to the
Problem of Obesity," Journal of American
Dietetic Association, 31:230-235, March, 1955.

Mayer, Jean, M. Marshall, J. J. Vitale, J. Christen—
sen, M. Mashayekli, and F. J. Stare. ”Exercise,
Food Intake and Body Weight, in Normal Rats and
Genetically Obese Adult Mice," American Journal
of Physiology, 177:544-548, June, 1954.

32.

33.

34.

35.

36.

37.

38.

39-

40.

41.

48

Mayer, Jean, P. Roy, and K. P- Mitra. "Relation
Between Caloric Intake, Body Weight and
Physical Work in an Industrial Male P0pula-
tion in West Bengel," American Journal of
Cligical Nutrition, 4:169-175, March-April,
195 .

 

 

McClelland, W. J. "Differential Handling and Weight
Gain in the Albino Rat," Canadian Journal of
Psychology, 10 19-22, March, 1956.

 

 

Passmore, R. "A Note on the Relation of Appetite
to Exercise," Lancet, 1:29, January, 1958.

Price-Jones, Cecil. "The Effect of Exercise on
the Growth of White Rats," Quarterly Journal
of Experimental Physiology, 16:61-67, 1927.

 

Sheer, B. T., S. Dorst, J. F. Codie, and D. F.
Soule. "Physical Capacity of Rats in Relation
to Energy and Fat Content of the diet," Ameri-
can Journal of Physiology, 149:194-203, April,
1947.

Shirley, M. "Studies of Activity. I. Consistency
of the Revolving Drum Method of Measuring the
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Psychology, 8:23-38, February, 1928.

 

Slanetz, Charles A. "The Adequacy of Improved
Stock Diets for Laboratory Animals," American
Journal of Veterinary Research, 4:182—189,
April, 1943.

 

 

Slonaker, James Rollin. "The Effect of a Strictly
Vegetable Diet on the Spontaneous Activity,
the Rate of Growth, and Longevity of the Albino
Rat," Leland Stanford Jr. University Publica-
tions, pp. 1-52, April 2, 1912.

Slonaker, James Rollin. "Long Fluctuations in Volun-
tary Activity of the Albino Rat," The American
Journal of Physiology, 77:503—508, August, 1926.

 

'Slonaker, James Rollin. ”The Normal Activity of the
Albino Rat from Birth to Natural Death, Its
Rate of Growth, and Duration of Life," Journal
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49

'42.w:510naker, James Rollin. "The Normal Activity of the

43.

44.

45.

46.

47.

48.

White Rat at Different Ages," Journal of Com~
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July, 1907.

 

 

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Activity in Relation to Diet in the Albino
Rat," The American Journal of Physiology,
142: 663-665, December, 1944}

 

Thomas, Barbara M., and A. T. Miller, Jr. "Adap-
tation to Forced Exercise in the Rat," The
American Journal of Physiology, 193:350—35 ,
May, 1958.

Tribe, D. E. "The Influence of Exercise on the
Selection of Food by Rats," British Journal
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Weininger, O. ”The Effects of Early Experience on
Behavior and Growth Characteristics," Journal
of Comparative and Ppysiological Psychology,
49:1-9, October, 1956.

Weininger,0 ., W. J. McClelland, and R. K. Arima.
Gentling and Weight Gain in the Albino Rat, "
Canadian Journal of Psychology, 14: 219- 224,
September, 1954.

Whalley, J., J. Howden, and F. R. Goetzl. "On the
Influence of Enforced Exercise Upon Voluntary
Food Intake of Rats," Permanente Foundation
Medical Bulletin, 9:114-118, October, 1951.

 

 

APPENDICES

51

 

 

 

 

 

 

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APPENDIX D

SEDENTARY GROUP FOOD CONSUMPTION FOR
POST—TREATMENT PHASE (IN GRAMS)

Aver.
Per Test
Period

Gr.

A
C

3.87
24.74

20.84
22.50
22.76
23.69
22.27
24.56
23.80
23.02
23.78
23.16
23.94
21.61
22.54

24.07
22.‘

25.04
23.82
24.01
24.16
25.14
24.06
2

20.12

 

10

Animals
5 6

4

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MNNNNNNNNNNNNCUCUNNCUNNC‘UCUCJCU

HCOUNHLnOM¢CUK3mw4rJN<DKDOKOGDOHflc>HrAO\

Punt-acnmhtc3HCUdnJCJmuneqoquunqu:rN)
NNNNNNNNNNNNNNNNNNNNNNNN

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NHNNNNHNNNNNNNNNNNNNNHHH

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.249

26

21.666

.429

I, —.
J

‘1
I-

9,

3
2e

21.828

20

24.024

.487,

L). E;

 

 

 

Testing Dates

 

4
2

11-14
15-18
19-23
24-27
28—Sep. 1
September 2-5
6-10
11-14
15-19
20—24
25-29
30-Oct.
5-9
lO-13
14-17
18-21
22=25
26-29
30—Nov.
3~6
7—10
11—14
5-18
19-22

1

November

Ochfler

August

 

Rat

lndiv.

Averages

 

 

r4
(\=

1""!

Gr.

Total

\j‘.

 

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.___'"_-._

 

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—-——uu-.— -0-

 

 

 

 

APPENDIX E

POST-TREATMENT PHASE (IN GRAMS)

CONTROL GROUP FOOD CONSUMPTION FOR

Aver.
Per'Test
Period

Gr.

22.34
21.06
23.25
23.65
22.30
21.57
22.20
22.58
21.47
23.03
22.31
22.86
21.54
22.47
22.40
23.06
21.56
23.15
21.58
22.82
21.75
21.23
21.62
21.97

 

 

10

Animals
5 6

4

I‘d'OCULfiln

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FJOHAFJFJN

20.8
23.7
18.6
20.9
19-7
21.3
20.0
21.2
22.0
21.9
19-3
21.1
18.8
21.2
19.2
20.1
21.1
20.3

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”3.308 21.937 25.329

22.179

20.312

21.191

1.174

2

54

J

20.5

 

 

Testing Dates

 

4

11-14
15-18
19—23
24-27
28-Sep. 1
2-5
6-10
11-14
15-19
20-24
25-29
30-Oct.
5-9
lO-l3
14-17
18-21
22-25
26-29
30—Nov.
3-6
7~lO
11-14
15—18
19—22

August
SepUHMBT
October
November

 

Rat

Indiv.

Averages

 

22.245

0
0

Average

Total Gr.

55

 

 

 

APPENDIX F

FORCED EXERCISE GROUP FOOD CONSUMPTION FOR

POST-TREATMENT PHASE (IN GRAMS)

Aver.
Period

Gr.
.Per Test

22.39
22.40
25.64
23.06
22.50
21.85
22.03
22.92
22.55
23.12
23.45
23.26
22.77
23.15
23.28
24.49
21.50
23.03
22.18
23.48
22.79
22.62
23.08
22.90

 

10

Animals
5 6

4

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NNN CU NNNNNCUNNNNNNNNN

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O O O O O O C O O O O O O O O I 0 O O O O I
HMNSNd'MHNNMMNNNN r-ir-szNNMN
NNNNNNNNNNNNNNNN NNNNNNN

CNN-\ON LCN\.0\OONI-lc-lo LCNd'kDONNMCDOr-iko LCNNLCN

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HHNNH HNNNNN NNNNNNNNNN
QNOHOQOQMQQQNNQNHQQQMNQN
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HHNNNHHHNNNNNNNNNNNN NNN
mHQQmQQNQONQ-fi; MQONONUD Ono:- ONCUQ
N mm QMN :NN: NmaNzN N
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#HMQHHHQQQNNMHDQOOQDMOOO
8flNNQHQNNH£NQQ£ HN NNNNH

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NNNNN NNN NNNNNNNNNNNN

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HM-d'UNr-ir-i (Ode-1H r-lr-iu-ld'r-iNl-INN NM
NNNNNN NNNCU NNNNNNNNN NN

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HNNHNd’NNNMNNd‘MMMONNONr-{NNi—l
NNNNNNHNNNNNNNNNHNNNNNNN

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0000:? mm m
r—lu—I N

21.
23
24.
21.
23.
21
22.
25
24
2O
22.
23
23.
23
22.
24.

N H
N N

2
2
2

25.320 25.324 21.574 22.420

22.179

22.037 23.374 21.979 22.649

22.487

 

 

 

Testing Dates

 

2

11-14
15—18
19-23
24-27
28-Sep. l
2-5
6-10
11-14
15-19
20-24
25—29
30-Oct. 4
5-9
10-13
14-17
18-21
22—25
26-29
30—Nov.
3-6
7-10
11-14
15-18
19-22

August
Sepumba?
October
NOVGMIT

Rat

Indiv.
Averages

 

 

22.934

Average

Total Gr.

56

 

 

APPENDIX G

AVERAGE COUNTER REVOLUTIONS PER GROUP
PER TEST PERIOD

 

 

 

 

 

 

Forced
Sedentary Spontaneous Exercise
Treatment Phase

Test Periods
1 1124 1705
2 863 1182
3 713 726
4 614 520
5 702 391
6 849 462
7 826 468

PostTreatment Phase

8 360 ' 580 785
9 483 619 801
10 524 ‘498 1005
11 648 542 1371
12 699 476 1202
13 747 543 1423
14 787 551 1430
15 830 664 1775
16 675 517 1558
17 922 584 1685
18 851 551 1266
19 964 631 1467
20 655 466 1153
21 640 430 1181
22 718 447 1155
23 603 409 1060
24 489 397 1031
25 568 452 1131
26 614 495 906
27 524 419 842

28 484 420 780

 

 

ANIMAL BODY WEIGHTS

 

 

 

 

 

 

Treatment Phase I Post—Treatment
Jul Jul Jul Jul Jul Jul Aug. Aug. Aug. Aug. Aug. Aug. Sept. Sept. Sept.
GPOUp 6 y 11y 16y 21y 26y 31y 5 10 15 20 25 30 4 9 14
5* 1 110.3 130.5 175.1 214.6 248.2 285.3 321.3 35 .5 338.8 342.6 363.5 372.0 382.5 395.0 400.4
g3 2 109.0 135.4 175.0 213.2 240.3 274.6 308.5 335.0 335.0 344.2 364.0 377.0 391.6 395.7 408.2
.61 3 108.2 135.5 177.6 203.2 237.1 272 0 303.2 331.5 325.5 337.3 346.7 358.8 372.8 381.0 388.1
g 4 110.1 143.4 186.2 220.3 245.5 276.5 297.5 326 0 320.3 337.1 343.6 353.0 360.7 388.5 401.6
3* 5 103.4 127.6 171.0 202.5 224.0 247.5 282.5 310.3 308.6 322.2 328.3 340.7 355.2 350.0 361.8
5 6 103.7 126.2 174.7 215.0 237.6 261.5 297.1 323.2 325.7 337.1 353.6 364.0 379.4 366.3 375.5
9 7 108.5 134.9 180.8 216.4 242.5 266 0 276.0 297.4 296.7 307.6 319.9 330.5 324.4 333.3 351.5
g> 8 106.4 133.4 174.2 216.0 224.6 252.7 273.4 296.0 307.0 321.0 335.8 348.0 361.5 35 .4 370.4
0 9 109.3 143.7 192.2 226.0 253.2 274.0 303.1 325.8 334.0 348.2 363.5 371.9 385.4 398.1 399.8
g 10 109.7 140.4 190.0 229.5 256.0 276.3 290.2 305.1 317.8 332.1 354.0 368.5 386.2 398.0 408.1
Avg. -107.9 135.1 179.7 215.7 240.9 268.6 295.3 320.4 320.9 332.9 347.3 358.4 370.0 376.5 .386 5.
E 1 109.2 124.0 168.5 197.8 235.3 258.5 287.0 306.6 323.8 346.4 363.3 372.4 385.4 392.5 398 0
- 2 106.2 125.1 162.0 187.2 193.5 217.0 247.0 271.3 297.3 319.4 333.5 342.3 360.6 374.8 383.2
6 3 108 5 127.3 166.0 201.0 219.0 240.5 269.2 286.5 308.8 322.7 333.4 349.0 359.4 361.0 368.8
g 4 109.7 134.1 174.6 209 0 252.0 274.5 298.3 316.2 328 3 343.2 360.9 363.5 379.0 384.2 396.0
g' 5 104.6 134 1 166.0 204.9 225.4 246.1 272.0 298.2 315.2 331.6 343.8 361.6 386.0 391.7 398.8
g 6 110.2 133.5 147.8 181.0 210.0 236.0L 270.0 306.4 317 .0 327.4 345.2 360.4 366.6 373.6 367.5
7 105.2 121.2 174.0 203.4 232.6 258.4 285.4 293.0 316.5 330.0 340.8 355.2 361.4 360.0 373.4
3 8 106 2 129.4 146.1 195.0 214.3 239.5 257 0 293.5 314.8 337.8 355 4 378.5 398.5 402.2 421.7
O 9 —— _- __ -.. __ __ __ __ __ '__ __ ~—. __ __ ..
g 10 105.2 120.8 156.1 180.6 211.6 245.7" 279.0 292.1 311 2 320.5 328.5 350.6 367.1 366 6 378.4
Avg. 107.2 127.7 162.3 195.5 221.5 246.2 273.9 296 0 314 8 331.0 345.0 359 3 373.8 378.5 387.3
l—l
E 1 108.3 138.6 173.5 202.3 218.2 244.5 258 5 279.8 298.1 308.5 321 0 330.2 342.6 348.6 349.2
1? 2 99.3 105.7 149.1 185.0 212.5 231.5 257 4 271.5 289.0 311.7 318.2 332.4 343.6 353.0 361 8
Q0 3 96.7 101.1 137.6 181.7 211.0 227.2 252.0 261 3 278.3 303.5 309.9 321.6 338.0 335 0 341.4
3g 4- 109.4 141.5 183.5 215.8 243.5 265.2 290.0 309 2 330.0 357.6 367 1 382.4 395 0 404.9 417.1
.gg 5 103 7 114.1 148.6 177.3 198.2 227.9 248.0 277 4 298.4 325.5 334.3 354.0 368 0 376.0 385.2 ,
m 6 106.7 124.8 169.9 216.3 238.9 269.2 298.2 330 8 351.0 367.1 376.6 389.0 403.8 412.7 422.6 i
N 7 107.3 132.2 148.0 191.5 222.5 240.4 267.5 276 0 280.5 306.5 311.7 316.8 328.6 330.0 336.4 E
g 8 106.1 118.8 150.1 179.9 184 1 203.9 228.5 243 5 255.2 279.1 289 2 301 0 320 8 320.2 323.3 1
3 9 104.9 122.5 159.9 187.5 212 1 234.9 259.4 217.7 268.8 302.2 320.0 338.5 353 5 366.0 373.3 1
m 10 110.4 143.0 184.0 217.5 230 6 240.0 260 0 278.2 268.8 319.8 333.8 348 .4 362 7 369 2 373 6 ?_
mAvg. 105.3 124.2 160.4 195 5 217 2 238.5 262.0 279.5 293.8 318.2 328.2 341.4 355 “ 361.6 368.9 E2 ;

 

 

 

 

 

 

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NOV‘ 23
Nov‘ Lé-‘—_:;9:;7
_NOV' 17 12.0 :EO'g
. 8 4 A7 .2 487~
NOV 468-9 518 1 0.8
001‘ 3 71_6 111.2 479'5 775.4
oct- ‘29 . 2 1196 471'6 169736 448°O
24 470'O 4-3 0 462- 41 .5 06.3
Oct. AR .6 £03.? M72.5 @18.3 4&3 8 %33.u
19 . .6 11 167-1 46 .o 474 4014 4.7.6
t 14 6 413 3 £20.? 455-3 A§$ 2 304.2 433'0 i04.0
QC 1 “ R" A8 ' .9 78' A .O 6-/ A8 ' 9 :.3
432 6 6'0 449 A 6 411' 43 R 7.6 462
Oct“ 9 . 7.3 473- A5 5 19.7 433 O - 6.4 473-8 49 6
t. 4 A2” O L:30 437'6 A _2 313- A3 O 7.11 461‘ 3.3
. Sep 6.21 469' 111/ 3 .16. 1133 3 7;.7 7778- 119 l 48 6
' 29 8 926 8 443-3 A27'0 1 0.8 409- A3“ 6 87-0 459' 7.2 445'
. / 7 I ‘ 43 .5 A76. 4 6 47 u 1-0
.. 24 414 6 A 3.4 416-0 410 2 7.7 M30 .8 “£8. 6.0 an. M1
. 2 43- 43 5 3.A 422- 39 2 1.0 470 4 9 47 A 0 786.7
19 415°; 121.5 118.2 40 7 97.1 425-0 470 O 714. 2 O “38.3 412-1 3.5
8-4 437'fi A 4 96-7 919.0 3 0.0 462° 46 8 J 47 '2 9.6 4727/ 46 3
GI’Oup 4 912.A 409- A17 o 3 1.7 383' A2 8 7771-11 4A7' O-2 438‘ 90 3 -66 O 431'4
A10'6 136 5 417-8 387‘4 116.3 418°C A51'3 42.1 6 176.1 412 O A73'4 7 3.7 429'
1 415' A1 '0 6.3 4OA°' 37 0 46.11 470'1 A 470- A3 0 54.0 458'6 136 2 120.2
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CD 3 399'5 380'3 366.3 400' 43 2 28-9r 0 0.0 405' A5; 3 ”3.5 419' 52 - 0.0
m . .1 O- 3 .7 450- A 50. 43 6.2 52- 42 -,0 _— 45 0
cl 4 372 8 39 6 5.0 419 0 A . 2 2.3 46 A“ 0 -6-7 521 5 1 8-
m . 2' 39 0 . 0.8 422‘ . 8,0 427° A0 a ,3 ugl- 41 a 2 1 440. 45
5 5 375 5 37 _3 425- A9 7 AA 6 ' 5 50-9 450 4 2 R09,3 - _ 4,A 2
Cr 6 69') 390. .6 412" 8 18- 388.4 4/ O 16' 410.6 2 _ 32.7 45 «29L
8 3 8 26-3 439 437' A 7 43- -4A9° A 0 4. ~, A 1 ~ 1.4 5
’1 7 380'6 A O 08-1 6 2.5 381' A 0 '8.3 413° 50 1.2 450° 25.7 42 '9
7: 8 412' A278 9 A29 4 413.11 435-9 A38 6 913.3 499-0 0 937.9 , 7 9 2.? 383'“
17 8 932'; 405.0 42g’i $3326 338 g 432'9 718 8 191.2 g 2 iii 5 993 4 71717 333.2 497';
1 3 1' . 9 ' ,6 4. 4O ' , °8 __ 43— ' 41 '. ---I0 . 3.; 40 Q
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1 390 37 5 22.5 389 A 4 _ 426 — A3 406 A 5 774°. 4: g 7 c--
11 2 65-7 418'5 A .6 5-2 460' ‘ .9 30-b 2 0.5-5 371‘4 1. .7 461-“ 379" 391.6 H
11 3 7 3. 371 39 1 _ 420 A _ 399' 1 O '82-. 453 291 0 1.3 4A1 5 8
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8 .0 9-. 38 _ 10. 42 8. 4O 3 ,6 47. 493 3 402 716. 441 8
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6+ 5 381 3 3 5 __ 40A A 397 . 1 2 '68'0 451 4 3 3-0 43* 4 4 7 4
*1 6 780 437. .l 09.1 .4 05.7 370'_ 4.4.8) 0.8 36f- 39 .2 hi . 434-
o 3 4 __ 395 A- I 397 ) A O 6.1 44; A9 3 L.7 431 z A . 1
,1 7 42A' 2 04.0 . 4 99.1 363°1 A5 0 6.8 362' 393 0 -33.8 431'
Q) 8 _- 393'5 A 387'0 3'?.© 450-‘ 415'9 9770A 78-7 AEB'G 9A4.8
a 4.0 404. .2 88'- 36— 5 76GB 471' 36C . 3- .7 1 6.b :-
E3 3 383 7 .,3 371.3 955'“ A38:6 121.? 63-5 359'? 113.5 :33-0
T51 39 377 0 3E9 5 ”7-g 426 g 9:0 U 361.7 418'3 45g 3
g. 0 " . C E7.J °7 7 7' . 43.°
AV 365:3 373.6 32;.0 407‘1 477.? 323.4 u17-8 413.2 /L
3
14 l 377 0 358I4 “00.6 453-4 35 / 3I5I3 478.I 41
L1 2 353 121.0 18.6.5 351»? 3 6 4:21
. 3 430 9 40 2 9 l b 78.3 40'°3 9 4 7
61 4 3961 446:1 3:44 332,. 3943 48'
*1 R 1 9 316‘7 397'8 K71.0 399°
643 J 43 O 45.3 410' 39 l
*44 8 339'5 389.8 389 3 399°
gpj.( 391.8 385.1L 388° 3
Tx4 8 38u'4 3 E 1
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8 9 38 0
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134