THE

RELATIONSHIP

B E T W E E N I N T E S T I N A L TR A NSI T ,

I N T E S T I N A L A B S O R P T I O N , AND S K E L E T A L
U P T A K E OF S T R O N T I U M 9 0 - Y T T R I U M 90
IN T H E

YOUNG CHICK

By
P a u l A. .T hg rn to n

AN A B S T RA CT
S u b m i t t e d t o t h e S choo l of A d v a n c e d G r a d u a t e S t u d i e s of
M i c h i g a n State U n i v e r s i t y of A g r i c u l t u r e a n d A p p l i e d
S c i e n c e in p a r t i a l f u l f i l l m e n t of the r e q u i r e m e n t s
f o r the d e g r e e of

D O C T O R O F P H I L O SO P H Y

D epartm ent

of P o u l t r y H u s b a n d r y
a nd

Institute

of N u t r i t i o n

Year

Approved
Date

1955

ABSTRACT

The t r a n s i t

rate

of S r

90

t h r o u g h th e d i g e s t i v e t r a c t of the

young c h i c k w as

s t u d i e d to d e t e r m i n e :

d ifferen t feeding

regim es,

other nutrients
affects

the

sidered

is

in

the

(2) if t h e t r a n s i t r a t e f o r

sim ilar,

retention
light

(1) if t r a n s i t is

a nd
90
<
,

(3)

w hether

of

Sr

The

of

the i n c o r p o r a t i o n

the

findings
of

Sr

rate

are
the

a f f e c t e d by
90

of

a nd f o r
transit

finally con­
tracer

into

t he

skeleton.
Two
per

cent

p.p.m .)
w ater
ported

different

of

t he

w ere

total

ration

studied.

movement
to

feeding

and

T he f i r s t

through

decrease

regim es

the

the
rate

procaine
is

bowel.
of

(lactose

penicillin

known
The

passage

incorporated

to

G

at

at
15

in crease rate

second

has

25

been

of
re­

of f e e d t h r o u g h the a l i ­

m entary tract.
S t r o n t i u m - 9 0, i n j e c t e d i n to t h e c r o p , i s
u p p e r gu t a t two d i f f e r e n t
approxim ately
initial
served.

30 p e r

"sp u rt."
This

rates.

removed from

D u r in g th e f i r s t 7 - 1 / 2

c e n t of the t o t a l d o s e i s

A fter this, a second

and

t i o n w h i c h is h i g h ly c o r r e l a t e d w it h t i m e .

m inutes,

r e m o v e d in an

slo w er

c a n be d e s c r i b e d by a s i m p l e

the

rate

is ob­

exponential
The p r e s e n c e

rate

equa­

of e i t h e r

lactose

o r penicillin has

no e f f e c t on t h e s e t r a n s i t

rates from

the

u p p e r gut s e g m e n t .
A. d e s c r i p t i o n
the
Sr

low er

90

intestine

into

idly.

the

In

However,
be

portions
as

absorbed

much
only

employed).

actually

absorbed

presented.

as

The

of
the

th e

to

Sr

per

s l o w ly

90

fraction

depends

upon

(of

after

the

initial

enters

at

reaches
spurt

last

the

(from

rap-

to

the

blood
dose)
the

portion

required

of

extrem ely

injected

a ll

tim e

it

m o v e s s lo w l y

the

this

the

90

absorbed

also

n ot

of

of

which

cent
or

Sr

be

in testine

50

of

All

appears

part
of

recovery

rapidly,
may

norm al

which

for

it

is

to

reach

vent.
The

rate

of S r

ing the f i r s t 7 - 1 / 2
to

the

gut

very

rations

the

is

lower

addition,

absorptive

of

120 m i n u t e s .

90

e x c h a n g e into th e b o n e is

m inutes,

a nd a g a i n f r o m

either

very

rapid d u r­

60 to 90, o r

90

At o t h e r t i m e s , bone u p ta k e a p p a r e n t l y d e p e n d s

up on the b a l a n c e b e t w e e n t h e a m o u n t of new bone f o r m a t i o n and
resorption.

Th e

4-hour

( 2 4 0 - m i n u t e ) n e t a c c u m u l a t i o n is l a r g e l y

i n d e p e n d e n t of d i f f e r e n c e s
absorption.

T h e n e t r e m o v a l of Sr

ton d o e s n o t s e e m
Sr

90

both in t r a n s i t r a te s

in the b l o o d .

to b e d i r e c t l y

90

from

a nd in i n t e s t i n a l

the b lo o d b y the

skele­

r e l a t e d to the c o n c e n t r a t i o n of

P a u l A. T h o r n t o n

In t h e l i g h t of t h e s e
quantity

of a n a l k a l i n e

l a i d down i n bone
m etabolism

of the

earth

findings,

it m a y be h y p o t h e s i z e d t h a t the

(strontiu m ,

is p r i m a r i l y

barium ,

radium ,

a f u n c t i o n of t h e i n t r i n s i c

o r calcium)
m ineral

s k e l e t o n i t s e l f r a t h e r t h a n of i n t e s t i n a l t r a n s i t

absorption.

iv

or

THE

RELATIONSHIP

B E T W E E N I N T E S T I N A L TR A NS IT ,

I N T E S T I N A L A B S O R P T I O N , AND S K E L E T A L
UPTAKE O F STRONTIUM90-Y T T R IU M 9°
IN T H E

YOUNG CHICK

By
P A U L A.

TH OR N TO N

A THESIS
S u b m i t t e d to the School of A d v a n c e d G r a d u a t e S t u d i e s of
M i c h i g a n State U n i v e r s i t y of A g r i c u l t u r e and A p p l ie d
S c i e n c e in p a r t i a l f u l f i l l m e n t of the r e q u i r e m e n t s
f o r the d e g r e e of

D O C T O R O F P H IL O SO P HY

D epartm ent

of P o u l t r y

Husbandry

and
Institute

of N u t r i t i o n

1955

P roQ uest Number: 1 0 0 0 8 6 7 6

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and there are m issing p a g e s, th e s e will b e noted. A lso, if material had to be rem oved,
a note will indicate the deletion.

uest,
ProQ uest 1 0 0 0 8 6 7 6
Published by P roQ uest LLC (2016). Copyright of the Dissertation is held by the Author.
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P a u l A.
candidate
D octor

f o r the

Outline

degree

of

of P h i l o s o p h y

F i n a l e x a m i n a t i o n , N o v e m b e r 23,
P o u l t r y Building.
D issertation:

T hornto n

1955,

9:00 A . M . ,

Conference

Room,

The R ela tio n sh ip B etw een In te s tin a l T r a n s i t , In te s tin a l
A b s o r p t i o n , and S k e l e t a l U ptake of S t r o n t i u m ^ ^ Y t t r i u m ^ i n the Young Chick.

of S t u d i e s
M a jo r subject:
M in o r subjects:

P o u ltry N utrition
Physiology, Biological C h e m istry

B iographical Item s
Born,

June

29,

U ndergraduate

G raduate

C a m p b e l l County,

Studies,

Studies,

Experience:

M ember

1925,

Kentucky

V e n t u r a J r . C o l l e g e , 1945-1946
U n i v e r s i t y of K e n tu c ky , 1 9 4 6 -1949

U n i v e r s i t y of Ke ntu c ky , 1952- 1953
M i c h i g a n State U n i v e r s i t y , 1953-1955

U.S. Navy, 1 94 3- 1946, 1950-1951.
R esearch
A s s i s t a n t , E x p e r i m e n t Sub-Station, P r i n c e t o n ,
K e n tu c ky , 19 49 -1 9 5 0 ; C a l i f o r n i a A g r i c u l t u r a l
E x t e n s i o n S e r v i c e , M o d e s to , C a l i f o r n i a , 19511952; G r a d u a t e A s s i s t a n t , U n i v e r s i t y of K e n tu c k y ,
19 52 -1 9 5 3 ; G r a d u a t e A s s i s t a n t , M i c h i g a n State
U n i v e r s i t y , 1953- 1955.

of P o u l t r y S c i e n c e ,
Sigm a.

v

S o c ie ty

of S ig m a Xi, S ig m a P i

ACKNOWLEDGMENTS

The

a u th o r w ishes to e x p r e s s

W o l t e r i n k of t h e
his

D epartm ent

guidance, a s s i s t a n c e ,

w ork was

conducted.

assistance

h i s a p p r e c i a t i o n to D r . L .

of P h y s i o l o g y and P h a r m a c o l o g y

F.

for

an d e n c o u r a g e m e n t d u r i n g the t i m e t h i s

He i s a l s o g r a t e f u l t o D r .

P.

J.

S ch a i b le f o r

and advice d u rin g th is tim e .

He i s i n d e b t e d to D r .
P o u l t r y H u s b a n d r y , and h i s

H. C. Z i n d e l , H e ad , D e p a r t m e n t of
s ta ff, a nd to D r .

B.

V. A l f r e d s o n , H e a d ,

D e p a r t m e n t of P h y s i o l o g y a n d P h a r m a c o l o g y , who f u r n i s h e d l a b o r ­
atory

space fo r th is
Gratitude

assisted

greatly

Sincere
m ission,

is

w o rk .
also

expressed

in the l a b o r a t o r y

to

M rs.

Cole,

who

and s ta tis tic a l an aly ses.

a c k n o w l e d g m e n t i s m a d e to the A t o m i c E n e r g y

D i v i s i o n of Biology and M e d i c i n e , f o r

a nd f o r f u n d s

Lois

in p a r t i a l

s u p p o r t of t h i s

stu dy .

Com­

s upplying t h e i s o t o p e

TABLE OF

CONTENTS

Page
I N T R O D U C T IO N AND R E V I E W O F L I T E R A T U R E

.

1

T r a n s i t R a t e s T h r o u g h the D i g e s t i v e T r a c t
i n th e B i r d
....................................................................................
Effect

of T r a n s i t R a t e s

Objective

of T h e s i s

C om parison

1

on A b s o r p t i o n ...................

8

.....................................................................

of R a d io s t r o n t i u m and R a d i o c a l c i u m

9
.

12

Dye M e t h o d s f o r Studying I n t e s t i n a l T r a n s i t

15

F e e d i n g R e g i m e s Which M i gh t be E x p e c t e d to
A lte r Intestinal T ra n s it or A bsorption
...................

17

L a c t o s e ........................................................................................

17

Penicillin

Z1

.............................................................

S u m m a t i o n and S t a t e m e n t of P r o b l e m
EXPERIM ENTAL
D e s c r i p t i o n of
Feeding
P reparation

METHO DS

G eneral P ro c e d u re s

R egim es

Studied

of S a m p l e s

D e s c r i p t i o n of

. . . .

. .

..................

Additional T r i a l s . .

D e r i v a t i o n of th e

Data

Z4
27

.

27
27
32
34
36

R E S U L T S AND DISCUSSION

46

vii

Page
R E S U L T S AND D I S C U S S I O N .........................................................................
L actose

E xperim ents

........................................................................................

46
46

B o d y W e i g h t ............................................................................................

46

Gat Weight

51

Bone Weight

......................................

52

M o v e m e n t of S t r o n t i u m - 9 0
Loss from

54

the u p p e r g u t .................................................................

M ovem ent through low er

gut

.....................................................

54
63

V a r i a t i o n i n e x t r a c e l l u l a r f l u i d a n d sof t t i s s u e
S r ^ w i th t i m e a f t e r i n j e c t i o n ..............................................

73

D e p o s i t i o n of S r ^ ^ i n the s k e l e t o n

..........................................

77

V a r i a t i o n due to h a n dl in g ...................................................................

88

Penicillin Experim ents
Body W e ig h t
G ut W e i g h t
B o n e W e ig h t

............................................................................

.......................................................................................................

92

...........................................................................................................

95

...................................................................................................

96

M o v e m e n t of S t r o n t i u m - 9 0 .....................................................................
Loss

from

92

the u p p e r g u t .....................................................

M o v e m e n t t h r o u g h t h e l o w e r gut

..........................................

90
U p t a k e of S r
b y the b l o o d a nd bo n e
d e p o s i t i o n ...............................................................................................

viii

100
100
103

108

Pag e
SUMMARY AND CONCLUSIONS
APPENDIX

......................................................

...................................................................................................................

LITERATURE

CITED

....................................................................................

ix

113
117
129

LIST

OF

TABLES

TABLE
I.
II.

Page
C alculated R a tes fo r Crop Emptying
C o m p o s i t i o n of B a s a l R a t i o n

III.

E f f e c t of L a c t o s e

IV.

F o u r - H o u r F e e d and

V.
VI.

VII.

VIII.

IX.

X.
XI.

XII.
XIII.
XIV.

XV.

Effect

of L a c t o s e

...

3

........................................

28

on Body a n d Gut Weight
W ater O b serv atio n s

on C a l c i f i c a t i o n

47
.

50

.....................

53

90
D i s t r i b u t i o n of S r
Recovered, Lactose
Experim ent
................................................................................

54

T h e E f f e c t of V i s i b l e D i f f e r e n c e s i n the
C r o p on I n i t i a l E m p t y i n g .....................................................

59

C o m b i n e d C o n t r o l a n d L a c t o s e D a ta ,
Bow el R e c o v e r i e s ........................................................................

61

90
90
M e a n P e r C e n t R e c o v e r i e s of S r
-Y
A f t e r C r o p I n j e c t i o n ....................................................................

72

Skeletal C learan ce

of Whole Blood S r

90

-Y

90

. .

90
E f f e c t of C h i c k H a n dlin g on Sr
Movement
a nd A b s o r p t i o n ............................................................................
Effect

of P e n i c i l l i n on Body and Gut Weight

E f f e c t of P e n i c i l l i n on F e m u r

W e i g h t ....................

90
D i s t r i b u t i o n of S r
Recovered, Penicillin
....................................................................................
E xperim ent
C o m b i n e d C o n t r o l a n d P e n i c i l l i n D ata,
B o w e l R e c o v e r i e s .............................................................

x

80

89
93
97

98

106

TABLE
XVI.

XVII.

XVIII.

XIX.

XX.

XXI.
XXII.

Page

90
90
M e a n P e r C en t R e c o v e r i e s of S r
-Y
A f t e r C r o p I n j e c t i o n ..................................................

107

90
90
S k e l e t a l C l e a r a n c e of S r
-Y
in
t h e P e n i c i l l i n S e r i e s .....................................................

Ill

E q u i l i b r a t i o n of Y
P a r e n t in Sam ples

D e t e r m i n a t i o n of the F a c t o r Used, to
C o n v e r t Whole Blood A c t i v i t y f r o m
C o u n t s p e r Second p e r M i l l i l i t e r to
P e r Cen t of I n j e c t e d Dose p e r M i l l i l i t e r
Tube

Conversion F a c to rs

...

XXIV.

C alculation

of Sr

. . . .

........................................................

V a l u e s ......................................................
90
-Y

90
A c t i v i t y in

118

119

L o s s Due t o H o m o g e n i z a t i o n of I n t e s t i n a l
Sam ples
...........................................................................................
Standard Activity

XXVI.

90
D a u g h t e r w i th Sr
of D i g e s t e d I n t e s t i n e

D e t e r m i n a t i o n of t h e F a c t o r U s e d to
C o n v e r t F e m u r A c t i v i t y to A c t i v i ty
of t h e T o t a l S k e l e t o n
..............................................

XXIII.

XXV.

90

t h e Blood

120
121

122
123
124

90
90
C a l c u l a t i o n of Sr -Y
P e r C en t
inT otal
S k e le to n
...............................................................................................

125

C alculation

12 6

of U p p e r Gut A c t iv i t y

xi

..................

LIST O F

FIGU RES

F ig u re
1.

2.

Page
90
T h e t r a n s i t r a t e of S r
f r o m the u p p e r to
....................................................................
t h e l o w e r gut s e g m e n t

5

90
R a t e of S r
p a s s a g e f r o m u p p e r to l o w e r
s e g m e n t of b o w e l .........................................................................................
90
present

3.

P e r c e n t of S r

4.

Sr

5.

90
T h e t r a n s i t r a t e of S r
f r o m t h e u p p e r to
.......................................................................
t h e l o w e r gut s e g m e n t

101

90
R a te of S r
p a s s a g e f r o m th e u p p e r to the
l o w e r b o w e l .......................................................................................................

102

6.

90

recoveries

in the l o w e r

90
present

P e r c e n t of Sr

8.

Sr

9.

W olte rink equation,

lactose

10.

W olte rink equation,

penicillin s e r i e s

in the

lower

. .
66

b ow e l . .

fro m lo w er b o w e l
series

xii

104
105

.

recoveries

64

f r o m the l o w e r b ow e l . . . . . . .

7.

90

b ow el . .

5

..............................
...

12 7
128

7

8

IN T R O D U C T I O N AND R E V I E W O F L I T E R A T U R E

T r a n s i t R a t e s T h r o u g h the D ig estiv e T r a c t in the

As early as

1904 B r o w n m e a s u r e d the t i m e

c o rn and oats to leave the
am ounts

of t h e s e

grains

c r o p of m a t u r e

w e r e fed.

of c o r n w a s f ed , a p p r o x i m a t e l y
c o r n to leave the

crop.

12 h o u r s w a s n e e d e d f o r a l l of t he

o v e r 24 h o u r s .

Browne

(1922) i n d i c a t e d t h a t t h e

starv ed for

35 g r a m s

O a t s gave e s s e n t i a l l y

of p a s s a g e to

sim ilar

results.

c r o p f i l l e d w i th o a t s e m p t i e d

C a r d e t a l.

(1926), u s i n g m a t u r e

hens

24 h o u r s , found t h a t a t t h e e nd of 11 h o u r s t h e

b i r d s which r e c e iv e d

50 g r a m s

of whole

c o r n fo ll o w e d by

w a t e r w a s e m p t y , w hile it t o o k a l o n g e r t i m e
t a n k a g e t o p a s s f r o m t he
F ro m these
m a y be e a s i l y

crop

f o r g r o u n d c o r n a nd

rate

of f e e d p a s s a g e

It if i s a s s u m e d t h a t t h e

e x p o n e n t i a l l y , a s d o e s the m a m m a l i a n

1

of

75 m l of

crop.

f i g u r e s , t he a p p r o x i m a t e

calculated.

in­

15 h o u r s , w hile i n c r e a s i n g the

lengthened the tim e

slightly

w i t h i n 18 t o 20 h o u r s .

hens when different

R a i s i n g t h e a m o u n t fed t o

50 g r a m s

req u ired for

He r e p o r t e d t h a t w h e n 30 g r a m s

c r e a s e d the tim e to a p p r o x im a te ly
a m o u n t of c o r n t o

Bird

crop em pties

s t o m a c h ( G o o d m a n e t a l .,

2
1952), and
periods
I are

that

are

for

"com plete”

r e q u i r e d , th e

obtained.

is based
strated

are

true

rate

' 1h a l f - e m p t y i n g 1'

co n stants listed

of c a l c u l a t i o n is i l l u s t r a t e d

That the a s s u m p tio n s

in T a b l e

in a f o o t ­

upon w h ic h t h e

calculation

t o a t l e a s t a f i r s t a p p r o x i m a t i o n will be d e m o n ­

in th is th e s is .
From

the

though the b ir d s

last

c o l u m n of T a b l e I, it w i ll be n o t e d t h a t a l ­

w ith f ul l c r o p s

lo n g er p e rio d th an those
leaving the
the

approxim ate

The method

n ote t o t h i s t a b l e .

e m p ty in g ,1 six

w i th l e s s

crop p e r m inute

crudeness

of th e

f e e d l e a v i n g th e

empty t h e ir

is

crops

over a much

f e ed , the a c t u a l a m o u n t of f e e d

rem arkably

constant.

o r i g i n a l d a t a , it i s l i k e l y t h a t the

crop p e r minute

is

considerably m o re

C onsidering
amount

of

uniform than

th is tabulation indicates.
Physiologically,

data

of t h i s t yp e

the

cro p is an i m p o r ta n t

the

l o w e r p o r t i o n of the b ow e l.

would s e e m to i n d i c a t e t h a t

r e g u l a t o r of the a m o u n t of f e e d p r e s e n t in

c o n tra ct e ith e r l e s s frequently

When t h e

c r o p i s v e r y full, it m u s t

or else m o re

weakly.

The n e t

result,

F o r e x a m p l e , a c r o p c o n t a i n i n g 50 g r a m s of f e e d would
c o n t a i n 25 g r a m s a t the end of one h a l f - e m p t y i n g p e r i o d , 12.5
gram s
at t he end of two h a l f - e m p t y i n g p e r i o d s , e t c .
T h u s , at the
end of s i x h a l f - e m p t y i n g p e r i o d s , the c r o p would c o n t a i n 0.8 g r a m s ,
w h i c h p h y s i o l o g i c a l l y would be " c o m p l e t e " e m p t y i n g .

3
TABLE I
CALCULATED

R A T E S FO R

CRO P EMPTYING

H alfEmptying
T im e
(m in.)

Rate
Constant
(m in . “ 1 )

Amount
E m ptied
in F i r s t
Minute
(gram )

Inv e s t i gator

Condition
of C r o p

B rown
(1904)

30 g r a m s
corn

12 h o u r s
(720 m i n . )

120

- 0 .0 0 5 7 7 5

0.173

35 g r a m s
corn

15 h o u r s
(900 m i n .)

150

-0.004620

0.162

50 g ram s
corn

24 h o u r s
(1440 m i n . )

240

-0.002888

0.114

f ul l of
oats ( a s ­
s u m e 50
g ram s )

1 8 -2 0 h r s .
( a s s u m e 19
h o u r s ; 1140
m inutes )

190

- 0.003647

0.181

Browne
(1922)

T i m e to
Em pty

E x a m p l e of c a l c u l a t i o n :
H a l f - e m p t y i n g t i m e = ( ti m e to e m p t y )/6 = 7 2 0 / 6 - 120 m i n u t e s .
K = r a t e c o n s t a n t = (InO. 5 ) / ( h a l f - e m p t y i n g t i m e ) = - 0 . 6 9 3 / 1 2 0
= -0.005775.
A m o u n t e m p t i e d in f i r s t m i n u t e :
- Kt
A s s u m e s i m p l e e x p o n e n t i a l = A /A
t
1o
Solve f o r t = 1 m i n u t e
A 0 = am ount in c ro p at s t a r t
At = 3 0 e - 0 . 005775
log At =
=
=
A.-t =
amount em ptied =

log 30 + ( - 0 . 0 0 5 7 7 5 ) log e
1.47712 - ( 0 . 0 0 5 7 7 5 )( 0 . 434294)
1.47461
29.827 = g r a m s i n c r o p a f t e r 1 m i n u t e
30.000 - 29.827 = 0.173 g r a m s

4
then,

would

s e e m to be t h a t a u n i f o r m a m o u n t of f e e d

the

Lower b o w e l p e r

the

dig estive t r a c t

p e r m inute).
obvious.

The

physiologic advantage

How ever, the data

One
rate

c it e d do not

objectives

of t h i s

l o a d of r a d i o a c t i v e

bowel fo r ab so rp tio n .
such m in e ra l

In o t h e r w o r d s ,

clear
exists.

study w ill be to i n v e s t i g a t e
c o n d i t i o n s and w it h s p e c i a l
lo w er

see w hether

r a t i o n p a s s t h r o u g h th e b ow e l m o r e
constituents

such as w ater.

b ow el a t d i f f e r e n t

rates.

and th e e f f e c t of t h e s e

H o w e v e r , the

differences,

if any,

on a c t u a l i n t e s t i n a l a b s o r p t i o n of the m a t e r i a l i s not known.
employing

radioactive t r a c e r s

answ ering

such questions.

im proved
ta tio n s

recent

on t h e

is

it i s i n i t i a l l y p o s t u l a t e d t h a t d i f f e r e n t f o o d s t u f f s

of d i f f e r e n c e

More

of

of f eed

s t r o n t i u m p a s s e d t o the

slowly t h a n o t h e r feed

p robably p a s s through the
m agnitude

of t he

a very

actually

It will a l s o be of i n t e r e s t t o

constitu ents

or m ore

(in g r a m s

constitute

such a physiologic m e c h a n is m

of t h e f i r s t

to the

load

segment

of s u c h a m e c h a n i s m

of t r a n s i t u n d e r d i f f e r e n t f e e d i n g

reference

rapidly

T h u s t he a b s o r p t i v e

would be g i v e n a u n i f o r m

d em o n stratio n that

the

unit of t i m e .

i s p a s s e d to

reports

early

would s e e m to be u n iq u e ly v a l u a b l e

found in t he l i t e r a t u r e

f in d i n g s

of B r o w n .

rate

of p a s s a g e

in

have not g r e a t l y

M o r e o v e r , the i n t e r p r e ­

u s u a l l y h a v e not b e e n p a r t i c u l a r l y p e r t i n e n t .

(1927) d e t e r m i n e d t h e

Methods

of whole

corn,

K e i t h et a l .
ground

corn,

5
and a m i x t u r e
the

results

alone
t wo

was

w ere

have

in i t s

rate

it w a s

in the

rate

of p a s s a g e

b e e n no d i f f e r e n c e

in the

rates

A l th o u g h
corn

c r o p t h a n the

other

w e r e n o t e d b e t w e e n th e

w e r e the

lower

segment

same, there

of

could
The

reflected m e r e ly different d e ­

of c r o p f il l i n g .
A g a i n Kaupp a nd Ivey

m easured
birds

crop.

load g i v e n the l o w e r b o w e l.

c r o p e m p t y i n g t i m e , t h e n , m u s t have
grees

f r o m the

of p a s s a g e t h r o u g h t he

If l o w e r t r a c t t r a n s i t

th e

c o n c lu d ed t h a t g r o u n d

S i g n i f i c a n t l y , no d i f f e r e n c e s

rations

the t r a c t .

c o r n a nd t a n k a g e f r o m

h ig hly v a r i a b l e ,

slow er

rations.

three

of g r o u n d

the

rate

(1923), u si n g l a m p b l a c k a s a m a r k e r ,

of t r a n s i t t h r o u g h t h e

gastrointestinal tra c t

w h i c h ha d b e e n s t a r v e d f o r 24 h o u r s .

feeds,

such as

of

T h e y found t h a t g r o u n d

w h e a t m i d d l i n g s and c o r n m e a l ,

required

3 hours

a nd

5 0 m i n u t e s to p a s s t h r o u g h g r o w i n g f o w l s a nd l a y i n g h e n s , while
m ature

h e n s not in p r o d u c t i o n r e q u i r e d a p p r o x i m a t e l y

b r o o d y h e n s a bo u t
lated

as

However,
is

before, a re
i n the

o b v io u s t h a t

m ate

12 h o u r s .

of the

-0.0181,

absence

respective

- 0 . 0 0 8 7 , and

of d a t a

on the

rate

constants,

and

calcu­

-0.0058 p e r m in u te .

s iz e

of the

crop

" p o o l , ” it

s u c h v a l u e s p r o v i d e no b a s i s f o r a r e a s o n a b l e

g r a m s p e r m inute

H euser

The

8 hours,

esti­

rate.

(1944) o b s e r v e d t h a t the

c r o p i s i n f l u e n c e d by t h e a m o u n t

rate

of p a s s a g e

f r o m the

of f ee d i n t a k e , w h i c h i s

in a g r e e m e n t

6
with B ro w n e 's
passes

observations made

f a s t e r th a n g r a in s ,

f a s te r than a dry m ash.
the t r a n s i t

th e f i r s t tw o

or three

hens.

It t h e r e f o r e

an X -ray

H o w e v e r , it t o o k
canal.

T he

a f t e r the

T his

segment

H euser
birds

as

5

suggesting a

of s t a r c h and a b s o r p t i o n of

sh o w t h a t a m o l e c u l e

of g l u c o s e

passes

c r o p pool due
w ith a m o l e c u l a r

a c r o s s th e i n t e s t i n a l m u ­

rapidly.
(1944) a l s o

may a lte r

p oi n t e d

Likew ise

out t h a t a c c e s s to f ee d in

crop emptying tim e,

f i r s t feed e a t e n m a y not e n t e r th e
proventricuius.

completely

sharply as e a rly

of c o u r s e , t h e e f f e c t of a s m a l l

g re a t a s that

150 m i n ­

r a p i d p a s s a g e t h r o u g h the n o n a b s o r p t i v e

The data a ls o

cosa extrem ely

starved

rapid h y d ro ly sis

reflected,

weight as

rose

in

au th o rs also noted that

i n g e s t i o n of a s t a r c h m e a l , t h u s

extrem ely

starvation.

c l o a c a a bo u t

16 t o 24 h o u r s to

same

blood s u g a r l e v e l of s t a r v e d h e n s

glucose.

study and o b s e r v e d t h a t

T h e f i r s t food a r r i v e d i n th e

correspondingly

to

s e e m s that w a te r m ay in c r e a s e

b o l u s e s go d i r e c t l y to the p r o v e n t r i c u l i s

e v a c u a t e th e a l i m e n t a r y

m inutes

He found a l s o t h a t m a s h

and i n t u r n , t h a t a wet m a s h p a s s e s

(1932) m a d e

utes a f te r ingestion.

t he

1922.

of f e e d t h r o u g h t h e i n t e s t i n a l t r a c t .

H e n r y e t a l.

starved

in

since

c r o p , but p a s s

Steinm etzer

(1924) and

show ed t h a t in s t a r v e d b i r d s th e f i r s t food

some

of the

d i r e c t l y to the
Fedorovskii

(1951)

consumed p assed directly

7
across

t h e d o r s a l p o r t i o n of t he

arrived

at the

p e a rs that

gizzard

starved

c ro p th ro u g h the p r o v e n t r i c u l u s ,

within 2 m inutes.

birds

sh o ul d u s u a l l y

and t h a t in n o r m a l b i r d s th e t r a n s i t

F ro m these

results,

and

it a p ­

give m i n i m a l t r a n s i t t i m e s ,

rates

would be

considerably

slow er.
H i l l e r m a n et a l .
perim ents

(1952) c o n d u c t e d a n e x t e n s i v e

d e a l i n g w i th food p a s s a g e

bio tics,

are

and age.

and p o w d e r e d
beforehand.
turkey

d i f f e r e n t a nd a r e

hens

c h a rco a l o r lam pblack.
results

(age 4 to

old t u r k e y h e n s

(age

in rate

Chicken hens,

were

b i r d s w e r e not

oxide
starved

years).

A c o m p a ris o n betw een

showe d a s i g n i f i c a n t l y f a s t e r t r a n s i t in the

was

ra tio n caused

on t h e b o r d e r l i n e

c o n t r a r y to t h e

show ed no i n c r e a s e

The

used; f e r r i c

anti­

significantly f a s t e r than through

1 - 1 /2 to 2 - 1 / 2

P e n i c i l l i n i n the

difference

c a n a l of c h i c k e n s

by e gg p r o d u c t i o n ,
were

re­

showed t h a t food p a s s e s t h r o u g h young

5 months)

l a y e r s and n o n l a y e r s
layers.

affected

Two d i f f e r e n t m a r k e r s

T he

of e x ­

a nd found t h a t the t i m e s

q u i r e d f o r food to p a s s t h r o u g h the a l i m e n t a r y
and t u r k e y s

series

in the

rate

report

a slo w er p a s s a g e ,
of s t a t i s t i c a l

significance.

of Kaupp a nd Ive y

of the p a s s a g e

but the

(192 3),

of food when t h e y

in p r o d u c t i o n .
A l l of t h e s e

studies,

while i n t e r e s t i n g ,

w h i c h i s b a s i c to a c l e a r u n d e r s t a n d i n g

of the

r e a l l y m i s s the p oint
re la tio n between

transit

rates

significance
is

on

tim e,

sorptive
I),

th e

other

th e

considered

tim e

so

equally

em ptied

in

not

usually

been

empty

im portant

the

of

w ith

the

are

tim e.

The

h ib i t e d

gastric

T he

rates

glucose
rate

m otility

w e r e found t o be

number

This

last

a
ab­
(T able

of

gram s

inform ation

on A b s o r p t i o n

rats

which w e r e f a s t e d f o r 48

solution was given orally,

le ft the

s t o m a c h in t h e

first

d e c re a s e d precipitously

due to c o n t a c t

closely

functional hyperm otility
rate,

w h ic h

of the

of g l u c o s e

m o re than

15 m i n u t e s .

f o r a 30 m i n u t e
result from

w ith the

e m p t y i n g a n d of i n t e s t i n a l a b s o r p t i o n

U y e y a m a e t al.

at a n o r m a l

half

lag in g l u c o s e t r a n s i t w a s a s s u m e d to

of g a s t r i c

th e

a bove

only

the

in

of

crop

is

food

o bt a i n e d .

A f t e r t h i s , the t r a n s i t
period.

area

th e
data

physio logical

b ow e l

dem onstrated

(1945), u s i n g m a t u r e

c e n t of the

t he

As

m easured

h o u r s , found t h a t w h en g l u c o s e
50 p e r

h av in g

of a b s o r b a b l e

E f f e c t of T r a n s i t R a t e s

Fenton

factor

surface

together

to

T he

gram s

covered.

required

The

of

absorptive

feed

has

variables.

number

the

surface

picture.
of

is

placed

given

and

in ­

duodenum.
of g l u c o s e

correlated.

(1953) o b s e r v e d t h a t h u m a n p a t i e n t s
of t h e d i g e s t i v e t r a c t a b s o r b e d

but t h a t a b s o r p t i o n of g a l a c t o s e

with

Vitam in A

was th re e

tim es

9
f a s t e r than in n o r m a l in d iv id u a ls.
b e tw e e n the i n c r e a s e

A c lo s e

c o r r e la t io n w a s found

in a b s o r p tio n of g a la c t o s e and the d e g r e e

in t e s t in a l h y p e r m o t ilit y .

The

rate

of

of a b s o r p tio n of m e th io n in e w as

found to be n o r m a l in t h e s e p a tie n ts .

C um m ins and A lm y (1953)

found that the a b s o r p tio n rate fo r g lu c o s e and m e th io n in e w a s i n ­
c r e a s e d by d r u g s w hich i n c r e a s e d in t e s t in a l m o t ilit y .

P a t i e n t s with

sp ru e who e x h ib ite d h y p o m o tility a c te d s im i l a r l y when t r e a t e d with
th ese drugs.

The m e c h a n i s m w h ereb y in c r e a s e d a b s o r p tio n i s

a s s o c i a t e d w ith i n c r e a s e d in t e s t in a l m o t ilit y i s unknown; h o w e v e r ,
C u m m in s and A lm y

s u g g e s t that i n c r e a s e d in t e s t in a l blood flo w m a y

be a f a c t o r .
M acDonald et al.

(1951) found that so m e la x a tiv e a g e n ts d e ­

c r e a s e d the uptake of str o n tiu m , w hile
of t h is e le m e n t in r a t s .

o th e r s i n c r e a s e d the uptake

T h is i s p r e s u m a b ly

r e la te d to the a b ility

of d iffe r e n t a g e n ts to bind d iffe r in g am oun ts of stro n tiu m w ithin the
tract.

The p r e c i s e

m e c h a n i s m s w e r e not e x p e r im e n t a lly w orked

out, h o w e v e r .

O b jectiv e of T h e s i s

It i s v e r y apparent fr o m the data just d i s c u s s e d that re l a %
t i v e l y lit t le i s

known about the rate of in t e s t in a l t r a n s i t (in t e r m s

10
of g r a m s

p e r m i n u t e ) an d t h a t e v e n l e s s

of d i f f e r e n c e s

in in te s tin a l t r a n s i t

T his th e s is
testinal tra n sit
to a l t e r th e
rates

rates

rate

rates

whe n f e e d i n g

a c r o s s th e

at which the
be r e p o r t e d .

t a k e n up by t he

of a t t e m p t s to m e a s u r e

regim es

l u m e n of the

of i n t e s t i n a l f u nc tio n,

rate

of t h i s type

blood, and t he
s k e l e t o n will

s hould c l a r i f y the

re­

skeleton.

we know v e r y l i t t l e

a bout t h e s e

aspects

it i s f i r s t n e c e s s a r y to e s t a b l i s h t h e n a t u r e

if any, b e t w e e n a b s o r p t i o n and t r a n s i t
d e p o s i t i o n (or

t h e o t h e r . It i s w e l l to b e g i n with what
sim ple

and

of t r a n s i t and the a m o u n t of m a t e r i a l

h a n d , and b e t w e e n a b s o r p t i o n an d t i s s u e

fairly

intestine

blood d e p o s i t s a m i n e r a l in t h e

S in c e , p h y s i o l o g i c a l l y ,

on

In a d d i t io n , the

i n t e s t i n a l m u s c o a t o e n t e r the

M easurem ents

in­

w h ic h m i g h t be e x p e c t e d

used.

blood a nd d e p o s i t e d in t he

relationships,

known a b o u t th e e f f e c t

on th e a b s o r p t i o n of n u t r i e n t s .

of i n t e s t i n a l t r a n s i t a r e

la tio n sh ip betw een the

the

results

at which a n u trie n t le a v e s the

travels

also

r e p o r t s th e

rates

is

situation.

on t h e

of

one

' ' u t i l i z a t i o n 1')

one m i g h t e x p e c t t o be

a

T h i s t h e s i s w i l l be c o n c e r n e d w ith the i n ­

t e s t i n a l b e h a v i o r of r a d i o a c t i v e

s t r o n t i u m - 90 f o r the fo llow ing

reasons:
1.

It i s e a s i l y m e a s u r e d .

2.

With c e r t a i n r e s t r i c t i o n s ,

it i s a good t r a c e r

for

calcium .

11
The in t e s t in a l a b s o r p tio n and ''apparent d i g e s t i b i l i t y ' 1 of c a l ­
cium a r e a s yet p o o r ly u n d ersto od d e s p ite a la r g e lit e r a t u r e b ea rin g
in d ir e c t ly
3.

on t h i s

sub ject.

The m e t a b o li s m

of im m e d ia t e

concern,

of S t r o n t iu m - 90 in plants and a n im a ls i s

s in c e it i s a sig n ific a n t con trib u to r to the

h a z a r d s a s s o c i a t e d w ith f a ll- o u t fo r n u c le a r f i s s i o n s .
4.

Strontium ,

Since the

s i m i l a r to c a lc iu m , i s a b o n e - s e e k in g e le m e n t .

s k e le to n i s the

sin g le quantitatively im portant

' ' s i n k 1' f o r S tr o n tiu m -9 0 in a n im a ls , we need not be c o n c e r n e d
w ith n u m e r o u s m e t a b o lic p o o ls and the
fo r m a t io n s w hich one m u st

c o m p le x m e t a b o lic t r a n s ­

c o n s id e r in dealin g w ith o r g a n ic f e e d ­

stuff s .
5.
is

F r o m the e c o n o m ic

s t i l l su ffe r in g e x t e n s iv e

Crooked t o e s ,

crooked

standpoint, the baby chick in d u stry

l o s s e s fr o m

Legs,

s k e le t a l a b n o r m a lit ie s .

crooked k e e ls , p e r o s i s ,

stargazers,

and e v e n r i c k e t s a r e am ong the m o s t co m m o n r e a s o n s for d e ­
str o y in g o th e r w is e

s a le a b le

c h ic k s.

A study su ch a s t h is m ay be

valuable in that it could le a d to a lo g ic a l b ase fo r applied
c o n c e r n in g t h e s e a b n o r m a lit ie s .

stu d ie s

12
C o m p a r i s o n of R a d i o s t r o n t i u m

Radioactive
uring the

absorption rate

sk ele to n , the
half-life
decay

because

suitable energy

chosen as a m ean s

of i t s

of i t s b e t a

selective

of m e a s ­

f i x a t i o n in t h e

r a y s and the

convenient

of 25 y e a r s w h i c h e l i m i n a t e s the n e c e s s i t y f o r p h y s i c a l

corrections

vantage

S tro n tiu m -9 0 was

and R a d i o c a l c i u m

in s h o r t - t i m e

experim ents.

i s t h a t it h a s a r a d i o a c t i v e

has a half-life
m aterials

m

of only

61 h o u r s .

secu lar equilibrium

T he

daughter,^

Y t t r i u m - 90, w h i c h

The p r e s e n c e
2

a t i o n of r a d i o a c t i v i t y i n u p p e r gut

chief d i s a d ­

of two r a d i o a c t i v e

i n t r o d u c e s no e r r o r
sam ples,

in d e t e r m i n ­

si nc e n e i t h e r i s a b ­

s o r b e d f r o m t h i s p o r t i o n of t h e d i g e s t i v e t r a c t .

However,

s in c e

Y t t r i u m - 9 0 i s not a b s o r b e d a p p r e c i a b l y f r o m t h e l o w e r bowel,
blood and bone

sam ples

90 i s f i r s t a b s o r b e d .

w i ll c o n t a i n no Y t t r i u m - 9 0 w h e n S t r o n t i u m -

T h e r e f o r e , the p r o d u c t i o n of Y t t r i u m - 9 0 f r o m

decay

of S t r o n t i u m - 9 0 w i l l c a u s e a n i n c r e a s e

th e se

s a m p l e s w it h t i m e

a v o id l a r g e
t i o n of th e

form

errors,

i n the a c t i v i t y

until a new e q u ilib riu m is atta ined.

of
To

it i s t h e n n e c e s s a r y e i t h e r , t o d e la y d e t e r m i n a ­

a m o u n t of r a d i o a c t i v i t y u n t i l the p a r e n t d e c a y and

^ Radioactive dau g h te r Y ttriu m -9 0
w he n S t r o n t i u m - 9 0 d e c a y s .
2

refers

to t h e

elem ent

S e c u l a r e q u i l i b r i u m e x i s t s w hen t h e p a r e n t h a s a h a l f - l i f e
v e r y m u c h l o n g e r t h a n the d a u g h t e r . C o n s e q u e n t l y , whe n e q u i l i b r i u m is
a t t a i n e d , the n u m b e r of p a r e n t a t o m s a nd d a u g h t e r a t o m s d i s i n t e g r a t i n g ,
p e r s e c o n d , is t h e s a m e .

13
d a u g h t e r g ro w th have

come

m a k e the

corrections.

appropriate

R adio active
l a r l y to
groups
Sr

90

into th e n e w s e c u l a r e q u i l i b r i u m ,

strontium

radioactive

has generally

calciu m .

of a d u l t w hite m i c e

However,

strontium

other tis s u e s

of the

w a s a bo ut t h e

by i n d i v i d u a l s a t two d a y s .

was f r o m
cent.

13 to

30 p e r

G r e e n b e r g and T r o e s c h e r

strontium ,
had

and

excreted

feces.

An a v e ra g e

G reenberg
rats

D istri­

r e s u lte d in a v a ria b le
of up ta ke f o r
6 to

(1942) i n j e c t e d m a t u r e

calcium

14 p e r

rats

intra-

a nd a s i m i l a r g r o u p w ith r a d i o ­
65 h o u r s the

25 p e r

rats

of b ot h g r o u p s

c ent of the i n j e c t e d d o s e i n the
cent of the t o t a l d o se

b ile ,

(1945)

c o m p a r e d the a c t i v i t y of Ca

45

and Sr

89

in

w h i c h had p r e v i o u s l y b e e n g iv en a r a c h i t o g e n i c d i e t and

found t h a t a l t h o u g h u pta ke
paralleled

a nd

s a m e f o r the two e l e m e n t s .

range

e l i m i n a t i o n of 4,5 p e r

o c c u r r e d t h r o u g h the

young

The

observed that a fte r

approxim ately

45

c a l c i u m t h a n of

cent; t h a t of s t r o n t i u m f r o m

p e r i t o n e a l l y w i th r a d i o c a l c i u m

Ca

s k e l e t o n in a 24 h o u r p e r i o d .

O r a l a d m i n i s t r a t i o n of t h e s e tw o e l e m e n t s
uptake

sim i­

(1941) i n j e c t e d

i n t r a v e n o u s l y w ith r a d i o a c t i v e

w a s t a k e n up by t h e

bution am ong

b e e n found to a c t

Pecher

and found t h a t a l a r g e r p e r c e n t a g e

o r to

of S r

89

e a c h o t h e r w h en t h e s e

Young r a c h i t i c

rats

was

Less t h a n Ca

isotopes w ere

45

, differences

orally a d m in is te re d .

a f t e r tw o d a y s e x c r e t e d abou t

67 p e r

c e n t of

0 90 .
Sr
m the
u p ta k e
the

feces,

of a b o u t

carcass.

89

to Sr

approxim ately

12 p e r

21 p e r

c e n t in the

Treatm ent

adm inistration

c en t in the u r i n e

skeleton, teeth,

of r a c h i t i c

rats

r e t a i n e d i n th e

cass,

and a b o u t

cent was e x c r e t e d

feces
The

33 p e r

45

orally,

when r a c h itic ;
45

Ca

2 0.5 p e r

c o n t e n t of t h e

w i t h V i t a m i n D.

Copp e t a l.
tor

s a l l y w ith S r

jected

w i t h Ca

45

had

(1945) found

and

when

and

car­

Sim ilar
in

the

g i v e n V i t a m i n D.

carcass

38 per
of

urine.

c en t of the d o s e

c ent

la rg e st percentage

a m o u n t e d to

cent in t h o s e t r e a t e d
b ot h Ca

45

and S r

89

taken

skeleton.

89 90
45
J , and Ca
.

approxim ately

Tweedy

rats

in the

60 p e r

skeleton, teeth ,

(1951) i n j e c t e d

tain ed

skeleton , te e th , and the

32.5 p e r

rachitic

The

up w a s found i n t h e

excreted

but only

c ent in the

of

r e s u l t e d in a f e c e s e x c r e t i o n of only 49 p e r

18.5 p e r c e n t w a s

g i v e n Ca

rem ainder

with V i t a m i n D p r e v i o u s

cent,

groups,

and

and had an

72 p e r

a bout

of r a t s d u r i n g the f i r s t

73 p e r

t io n .

Repeating the

24 h o u r s

intraperiskeleton con­

s t r o n t i u m given:

cent of a d o s e

urine

sam e

of young r a t s

A f t e r e i g h t d a y s the

cent of the

retained
11 p e r

groups

those

c ent of c a l c i u m giv en.
of S r

89

excreted

in the

after intraperitoneal injec-

e x p e r i m e n t w i th Ca

45

.
, the to tal u rin a ry

excretion

o v e r a 6 6 - h o u r p e r i o d a m o u n t e d t o only a bo ut 2 p e r

R ecently,

C o m a r et al.

(1955) have

in-

show n f i v e - f o l d d i f f e r e n c e s

cen t.

15
b e t w e e n Ca

45

a nd Sr

90

o v e r tw o g e n e r a t i o n s
From

these

r e s u l t when Sr
results

are

activity.

90

behavior

data,

or Sr

it a p p e a r s t h a t

89

are

t o be i n t e r p r e t e d

such d ifferences a re

certain discrepancies

as quantitative

considered

to d if fe r with d iffe ren t

and t i s s u e

components

90

a nd Ca

of c a l c i u m
45

a p p e a r to

of t h e p r e s e n t e x p e r i m e n t s ,

critical.

h o w e v e r , t h a t the e x a c t q u a n t i t a t i v e
absorption,

indications

between Sr

F o r the p u r p o s e s

not

may

u s e d i n bioLogical e x p e r i m e n t s , if the

H o w e v e r , the d i f f e r e n c e s

intestinal tra n sit,

running

of r a t s .

be m a i n l y q u a n t i t a t i v e .

m ind,

in l o n g - t e r m e x p e r i m e n t s

It m u s t be b o r n e in

re la tio n sh ip s betw een

d e p o s i t i o n wouLd be e x p e c t e d

of the

ration.

Dye M e t h o d s f o r Studying I n t e s t i n a l T r a n s i t

It w a s th o u g h t a d v i s a b l e to m e a s u r e the
t h r o u g h the d i g e s t i v e t r a c t by t h e u se
isotope m e a s u r e m e n t .
of m e a s u r e m e n t
To use

sim ultaneously

of t r a n s i t

of a dye i n a d d i t i o n to the
of u s in g two
same

separate

individual a r e

intestinal tra n s it,

m ethods

o b v io u s.

it m u s t f i r s t

It shouLd be u n c h a n g e d in i t s p a s s a g e t h r o u g h

gastrointestinal tract.

l u m e n a l wa ll.

on t h e

a dye in m e a s u r i n g

of a l l be n o n t o x i c .
th e

The a d v a n t a g e s

rate

Fin ally,

It m u s t not a d h e r e to o r

it m u s t be n o n a b s o r b a b l e .

s t a i n the

16
G oodm an et a l.
w ith an aq u eou s

(1952)

s to m a c h -tu b e d n o n sta r v e d adult r a ts

so lu tio n of E v a n s Blue dye and found that g a s t r i c

em p ty in g p r o g r e s s e d at an e x p o n e n tia l rate.
Blue w a s v e r y high in a i l c a s e s .
blood p ro d u ced no to x ic e f f e c t s .
a n a ly se s

A f t e r o r a l a d m in is tr a tio n , blood

shown l a t e r ,

f o r th e u p p e r s e g m e n t

a single

in t h e

of the

exponential ra te p r o c e s s .

with E v a n s

Blue w i t h t h e

t i o n , th e E v a n s
the

E vans Blue in t h is

research.
A s w i l l be

90

T h e r e f o r e , t h is dye

r e q u ir e m e n t s p r e v io u s ly outlined for study in the rat.

C o n se q u e n tly , an attem pt w as made to u se

Sr

of E v a n s

In jection of E v a n s Blue into the

r e v e a le d no E v a n s Blue at any t im e .

m e t a ll of the

th e sis

The r e c o v e r y

duodenum.

intestinal tra n s it
procedure.

However,

of th e d ye, b e g a n a s

i n th e

sim ilar m easurem ents

c h i c k gave v e r y e r r a t i c

T his breakdown,

Therefore,

Evans

Blue

r a t , th e l o s s

of

.
.
.
.
.
chick digestive t r a c t is e sse n tia lly

Blue w a s b r o k e n dow n v e r y

low er intestine.

afte r extraction

c h i c k a s in t h e

as

results.

rapidly

In a d d i ­

upon r e a c h i n g

shown by a c o l o r ch ange

s oon a s the dye

r e a c h e d the

couLd not be u t i l i z e d to m e a s u r e

c h ic k, w ith o u t f u r t h e r e f f o r t s to r e f i n e th e

17
F e e d i n g R e g i m e s Which M i g ht be E x p e c t e d to A l t e r
In testinal T ra n s it or A bsorption

Lactose

A 25 p e r
ing

regim es

cent la c to s e

on t h e

chosen as

one of t h e f e e d ­

s u p p o s i t i o n t h a t it would i n c r e a s e

gastrointestinal tran sit,
been

ra tio n was

s h ow n to p r o d u c e
R u tte r et al.

since high le v e ls

of t h i s

the

substance

rate

of

have

diarrhea.

(1952) f ound t h a t l a c t o s e up to 20 p e r c e n t of th e

r a t i o n did not i m p a i r g r o w t h o r o b vi ou s ly a f f e c t the p h y s i o l o g i c a l w e l l ­
be ing of young c h i c k s .

A bove t h i s l e v e l , l a c t o s e c a u s e d g r o w t h i m p a i r m e n t ,

d i a r r h e a , and a c r o o k e d t o e d e f o r m i t y i n p r o p o r t i o n to t h e l e v e l of l a c t o s e
f ed .

Blood a n a l y s e s showed s m a l l a m o u n t s of l a c t o s e , but no g a l a c t o s e ,

indicating that the

young c h i c k i s u n a b l e to h y d r o l y z e

lactose.

F i s c h e r and Sutto n (1949) s u g g e s t t h a t l a c t o s e m i g h t p r o d u c e
diarrhea
include

in one o r a n u m b e r
(1) d i r e c t

of d i f f e r e n t w a y s .

T he p o s s i b i l i t i e s

s t i m u l a t i o n of i n t e s t i n a l m o t i l i t y ;

actio n--interference

(2) h y d r a g o g u e

w i th a b s o r p t i o n of w a t e r and o r g a n i c n u t r i e n t s ;

(3) a l t e r a t i o n of i n t e s t i n a l f l o r a to a n a c i d u r i c t y p e , the a c i d u r i c
f l o r a th e n acting in s e v e r a l dif feren t po ssib le
of pH e n o u g h t o

s tim u la te the i n te s tin a l m u s c u l a t u r e ,

l a t i o n of i n t e s t i n a l m u s c l e
am ounts

w a y s;

th ro u g h its n erv e

of u n h y d r o l y z e d l a c t o s e

in t h e blood.

(4) a l t e r a t i o n
and

supply by

(5)

stim u­

sm all

They fu r th e r

point

18
out t h a t ,

of t h e s e

p o s s i b i l i t i e s , the hydragogue

hypothesis

seem s

m o s t pla usible.
Although lacto se

is

u n d o u b te d ly a n o s m o t i c

laxative, tis s u e

d e h y d r a t i o n d o e s not i n v a r i a b l y a c c o m p a n y a p e r s i s t e n t
diarrhea.

D e h y d r a t i o n would be e x p e c t e d

action were
urine

involved, u n le ss

w ater

if a c o n t in u o u s h y d r a g o g u e

consum ption is in c r e a s e d

or

volume d e c r e a s e d .
Roccuzzo

(1945) found t h a t a s o l u t i o n of l a c t o s e

m outh, w h eth er h y p e r-,
c r e t i o n of l e s s
that lactose

of c a l c i u m .

iso-,

o r h yp o to n ic ,

interfered

Kline et a l.

show t h a t l a c t o s e e n h a n c e s the

intestinal tra c t.

ce nt in a r a c h i t i c

utilization

r a t i o n , a n d found t h a t

skeleton were

improved.

r e d u c e d th e pH t h r o u g h o u t the e n t i r e
In t he p r e s e n c e

calcification, th e r e f o r e

indicating

(1932) fed l a c t o s e to White L e g h o r n

a b s o r p t i o n and m i n e r a l i z a t i o n of th e
l e v e l of l a c t o s e

of w a t e r ,

w ith w a t e r a b s o r p t i o n .

reports

a t a l e v e l of 40 p e r

g i v e n by

a l w a y s led to t h e e x ­

u r i n e t h a n did a n e q u a l v olu m e

N um erous

that

lacto se

chicks
calciu m
This

l e n g t h of the

of V i t a m i n D, l a c t o s e a l s o i n c r e a s e d

i n d i c a t i n g t h a t it h a s a r o l e

o v e r an d above

of V i t a m i n D in c a l c i u m a b s o r p t i o n .
Bergeim

increased

(1926) found t h a t 25 p e r

c a lc iu m a b s o r p tio n in r a t s ,

cent l a c t o s e

i n the

ration

both on r a c h i t i c a n d in V i t a m i n

D s u p p l e m e n t e d d i e t s , t h u s c o n f i r m i n g the f in d i n g s

of Kline, e t al.

19
F r e n c h and

C o w g i ll (193 7) found t h a t 2 0 p e r

ration increased
pups,

a l t h o u g h no

However,
or its

such r e s u l t s

(1942) found t h a t

At th e

20 p e r

2 m illiliters

tion in c r e a s e d nonradioactive

in young

5 per

cent la c to s e

lactose

was

M a c D o n a l d et al.

of a p o w d e r e d m i l k

strontium absorption from

c ent e v e n t h o u g h t h e m i l k a n d the

dog.

f a i l e d to i n c r e a s e

cent l e v e l ,

to a c t u a l l y i n h i b i t c a l c i u m a b s o r p t i o n .
as

in the

o b ta in e d w it h a m a t u r e

g l u c o s e and g a l a c t o s e ,

a b s o r p t i o n in r a t s .

(1952) found t h a t a s l i t t l e

per

were

R o b e r t s a nd C h r i s t m a n

reported

lactose

u t i l i z a t i o n of caL c i u m a nd p h o s p h o r o u s

hydrolysis products,

calcium

cent

strontium were

16 to

solu­
30

given con­

currently .
It h a s long b e e n known t h a t V i t a m i n D i n c r e a s e s the
of t h e

i n t e s t i n a l t r a c t and it h a s

cause

of t h e

position,

increased

D i n c r e a s e d th e a m o u n t
chicks,
was

w h en t h e

injected

these
D was

results

Ca

45

In s u p p o r t of t h i s

J a m i e s o n (1955) have
of Ca

45

d e p o s i t e d in the

c o n c l u d e d t h a t the p r i m a r y

and large

of m a t u r e

Vitamin

h o w e v e r , w h en Ca
From

e f f e c t of V i t a m i n

intestine.

(1953) m a r k e d l y i n c r e a s e d the a c i d i t y

intestine

sup­

s k e l e t o n of young

V i t a m i n D h ad no e f f e c t .

on th e a b s o r p t i o n of c a l c i u m f r o m the
A shcraft

shown t h a t

was a d m in is te r e d orally;

intram uscularly,
it w a s

o fte n b e e n t h o ug h t t h a t t h i s i s the

calcium absorptio n.

M i g i c o v s k y and

acidity

of the

ceca

h e n s by f e e din g a r a t i o n c o n ta in in g

45

20
20 p e r

cent lacto se .

affected.

of r a t s

b e l o w t h e d u o d e n u m whe n t h e

i l e u m and

cecum,

rats

scraps.

s i d e r a b l y i n f l u e n c e s th e
The evidence
at a

occurred

results

are

cite d) who u s e d a d ie t w h i c h c o n t a i n e d
T h u s the n a t u r e
results

o b ta in ed .

t i o n of c a l c i u m .
in e x p e r i m e n t s
The use
cedure.

be

It a l s o

r a t i o n i n c r e a s e s the

T h e s e two o b s e r v a t i o n s m a k e the u se

in­

It m i g h t be

is also in c re a s e d .

in the

whe n fed

and c e r t a i n l y

absorp­

of l a c t o s e

on t r a n s i t a nd a b s o r p t i o n a v e r y f e a s i b l e p r o j e c t .
of l a c t o s e ,

however,

i s not an u n a m b i g u o u s p r o ­

Since a lactose , r a t i o n d o e s l o w e r the pH of the

any e f f e c t
ally

of f ee d p a s s a g e

c e rta in that lacto se

10

of the b a s a l r a t i o n c o n ­

c r e a s e s t h e w a t e r f low t h r o u g h t h e a l i m e n t a r y t r a c t .

se e m s fairly

was le s s .

s i m i l a r to those

just d is c u s s e d in dicates that lactose

rate

d ie t.

only in the

of c hange

sufficient le vel c a u se s an osm otic d i a r r h e a

s u p p o s e d t h a t the

c e nt l a c t o s e

on a v e g e t a b l e

a nd e v e n h e r e th e m a g n i t u d e

(previously

cent m e a t

were

change

It i s i n t e r e s t i n g to note t h a t the l a t t e r
of A s h c r a f t

not

s h a r p l y i n c r e a s e d the a c i d i t y in the d i g e s t i v e

When m e a t w a s a dded t o the d i e t t h e

per

were

R o b i n s o n and D u n c a n (1931) found t h a t 2 5 p e r

in t h e d i e t
tract

H o w e v e r , t h e d u o d e n u m and i l e u m

on i n t e s t i n a l a b s o r p t i o n of s t r o n t i u m

re g a r d e d a s an effect a s s o c ia te d

r a t e th r o u g h the t r a c t .

In a d d i t i o n , the

bowel,

could not a u t o m a t i c ­

only w it h a n a l t e r e d t r a n s i t
altered

w a te r balance

in

21
the

b o w e l m i g h t i n i t s e l f m o d i f y t h e a b s o r p t i o n of a n a l k a l i n e

The e n tir e
ever,

earth.

situation obviously needs e x p e rim e n ta l c larification.

How­

it i s d e s i r a b l e t h a t o t h e r a g e n t s w h i c h m o dif y b o th food p a s s a g e

and c a l c i u m m e t a b o l i s m
d i r e c t e d to t h e e f f e c t s

be

studied.

C o n s e q u e n t l y , a t t e n t i o n w as a l s o

of p e n i c i l l i n .

Penicillin

P e n i c i l l i n and
s h o w n to

s e v e r a l o t h e r a n t i b i o t i c s have b e e n r e p e a t e d l y

s t i m u l a t e g r o w t h a nd i m p r o v e f e e d e f f i c i e n c y i n c h i c k e n s .

H ow ever, the m e c h a n is m
entirely

by w h i c h g r o w t h s t i m u l a t i o n o c c u r s

i s not

clear.
R o s s and Y a c o w i t z

(1954) found t h a t p r o c a i n e p e n i c i l l i n G at

a l e v e l of 2.4 g r a m s p e r t o n s i g n i f i c a n t l y i n c r e a s e d th e w ei gh t of bone
a s h in c h i c k s .
sorption.
adequate

R esu lts indicated that penicillin in c r e a s e d

H o w e v e r , th e
supplies

L indblad

et al.

aureom ycin

tibiae

s i n c e no i n c r e a s e

of, o r a t v e r y lo w l e v e l s

of,

in bone a s h w a s
V i t a m i n D.

(1952) h a v e p r e s e n t e d e v i d e n c e w h i c h s h ow s t h a t

supplem entation may also in c re a s e

M i g i c o v s k y et a l.
kilogram

i n c r e a s e d a b s o r p t i o n w a s d e p e n d e n t up o n

of V i t a m i n D,

n o t e d in t h e a b s e n c e

calciu m a b ­

of r a t i o n

(1951) r e p o r t e d t h a t

calcium utilization.

30 m i l l i g r a m s

s i g n i f i c a n t l y i n c r e a s e d t h e u p ta ke

o v e r t h a t of the

control,

w h en Ca

45

of p e n i c i l l i n p e r
of Ca

45

w as given orally .

by t h e

22
V'/aibel et a l.
of t h e
a

(1952) i n c r e a s e d th e b i o ti n and f o li c a c i d

egg yolk by i n c o r p o r a t i n g two d i f f e r e n t l e v e l s

standard

procaine

breeding

ration.

penicillin G per

by 29.6 and

38.5 p e r

Folic

acid

cent,

respectively,

penicillin

increases

of p e n i c i l l i n into

P e n i c i l l i n a t 5 and 200 m i l l i g r a m s

kilogram

ce nt,

of r a t i o n i n c r e a s e d

respectively,

f o r the

same

after

16 d a y s

groups were

w i th t h e i n c r e a s e s

occurring

yolk b i o t i n
of t r e a t m e n t .

31.0 and 2 6 ,7 p e r
30 d a y s a f t e r

(1953) o b s e r v e d t h a t p e n i c i l l i n o r a u r e o m y c i n in the

c h i c k s ’ r a t i o n d e c r e a s e d t h e gut w e ig ht.

This

c h ange in gut w eigh t

i s no t r e f l e c t e d in the l e n g t h of the gut, w h ic h w a s m u c h t h e
in both t r e a t e d
w ater

no d i f f e r e n c e
gut.

to

and u n t r e a t e d
o r fat c o n t e n t.

b e t w e e n the

chicks.

H istological exam inations

heavier

raised

stim ulate

in a g e r m - f r e e

growth.

revealed

c o n t r o l and th e l i g h t e r p e n i c i l l i n

T reatm ent

environm ent.

sterile

reported

antibio tics fail

gut to be m u c h l i g h t e r

conventionally-kept, n o n ste rile

of c o n v e n t i o n a l l y - k e p t

gut w e i g h t i n a m a n n e r
germ -free

environm ent where

He found the

i n w eig ht t h a n t h a t f r o m

same

N e i t h e r i s it due t o a n

G o r d o n (1952), in c o n f i r m a t i o n of C o a t e s ’ o b s e r v a t i o n ,

on c h i c k s

of

s u p p l e m e n t a t i o n had begun.

C oates

altered

c o n te n t

chicks.

c h i c k s w ith a n t i b i o t i c s d e c r e a s e d

s i m i l a r to t h e d e c r e a s e

o b s e r v e d in the

23
The

effect

of d i e t a r y

(Migicovsky et al.,
folic a c id

1951) a nd on th e

( W a i b el et a l . ,

(1) t h e a m o u n t
absorptive

of t h e s e

capacity

s e r v e d by C o a t e s
to indicate

p e n i c i l l i n on c a l c i u m a b s o r p t i o n

1952)

s u g g e s t s t h a t the a n t i b i o t i c i n c r e a s e d

n u t r i e n t s a v a i l a b l e to the

of t h e

gut w all.

(1953) and G o r d o n

a higher absorptive

of t h e gut.

c o n c e n t r a t i o n of y olk b i o t i n and

E i t h e r of t h e s e

The

host,

added to a ration.

(2) the

l i g h t e r gut w e i g h t s

ob­

(1952) have a l s o b e e n t h o u g h t

capacity

or an in c r e a s e d

' ’e f f i c i e n c y ' '

h y p o t h e s e s m i g h t w e l l e x p l a i n th e i n ­

c r e a s e d feed efficiency and stim ula te d growth rate
are

or

H o w e v e r , n e i t h e r c a n be

when antibio tics

regarded as

proved.
H i l l e r m a n e t a l.

(1953) s h o w e d t h a t p e n i c i l l i n a d d e d to the

r a t i o n of a d u l t c h i c k e n s and t u r k e y s
sage.

T his m ight well cause

d o w n th e

rate

m o re tim e
segm ents
the

of p a s s a g e

d e c r e a s e d the

growth stim ulation,

s l ig h t ly , the d i g e s t i v e

rate

of food p a s ­

s i n c e by s low ing

j u i c e s would have

to c o m p l e t e d i g e s t i o n a nd , i n a d d i t i on , t h e a b s o r p t i v e
of t h e i n t e s t i n a l t r a c t

available

would have m o r e t i m e to a b s o r b

nutrients.

Gabutin and Shaffner

(1954) have

shown t h a t r a t i o n s

c on ta i n in g

p e n i c i l l i n m a i n t a i n h i g h e r blood c a l c i u m l e v e l s in laying h e n s .
Sturkie

and P o l i n

com plish this

(1954)

result.

r e p o r t that

sin g le

i n j e c t i o n s w i l l not a c ­

24
To
transit
in the

sum m arize,

i n c l u s i o n of a n t i b i o t i c s i n the

r a t e s th ro u g h the
ordinary

accom plished
of t h e s e

sense.

b o w e l and i n c r e a s e s

a g e n t s i n th e p r e s e n t

particularly

since t h e i r effects

s u p p o s e d t o be

N evertheless,

included in this

11u t i l i z a t i o n , ''

whereby these

are

f u r th e r investigation

c o n te x t a p p e a r s to be highly d e s i r a b l e ,
on c a l c i u m

s i m i l a r to th e e f f e c t s

a c o m p a r i s o n of p e n i c i l l i n - f r e e
is

calcium

A g a in , the m e c h a n i s m s

rem ain obscure.

ration reduces

' ' a b s o r p t i o n 1' h a v e b e e n

of l a c t o s e .

F o r this

reason,

and p e n i c i l l i n - s u p p l e m e n t e d

ratio ns

st u d y .

S u m m a t i o n a n d S t a t e m e n t of P r o b l e m

It i s

o b vi ou s f r o m t h i s

a b s o r p t i o n ' 1 of s t r o n t i u m
am in ed
from

r e v i e w of the l i t e r a t u r e t h a t

( o r of c a l c i u m ) h a s not b e e n c r i t i c a l l y e x ­

by d i r e c t e x p e r i m e n t a l p r o c e d u r e s .

various

been altered
very te r m

in d ire c t r e s u l t s that
in t h e

''in te s tin a l

course

It i s a l w a y s i n f e r r e d

' ' i n t e s t i n a l a b s o r p t i o n ' ' m u s t have

of a g i v e n e x p e r i m e n t a l t r i a l .

T h u s the

' ' a b s o r p t i o n ' 1 h as a c q u ir e d a m o s t indefinite m ean in g b e ­

cause a num ber

of d i f f e r e n t end p o i n t s have b e e n u s e d .

Specifically,
of a b s o r p t i o n a n d

it i s n e c e s s a r y to d i s t i n g u i s h b e t w e e n t r u e

s t u d i e s i n w h ic h m e r e l y the

orally ad m in istered

m aterials

d istin g u ish betw een p a ssa g e

have

been m ade.

of s t r o n t i u m f r o m

studies

s k e l e t a l r e t e n t i o n of
Further,

one m u s t

gut to blood

(which

25
is the

only p r o c e s s

process

whereby

of a b s o r p t i o n in t h e

strontium p a s s e s

To th e n u t r i t i o n i s t ,

loose

not

reverse

f r o m th e blood b a c k to t h e gut.

The d i f f e r e n c e s

''a b s o rp tio n .''

' ' a b s o r p t i o n 1' c o n c e p t

factors

s e n s e ) a nd the

' ' a b s o r p t i o n ' 1 is the a r i t h m e t i c d if fe re n c e

t w e e n t h e s e tw o p r o c e s s e s .
absorption,

strict

be­

sh ou ld be c a l l e d ne t

Only by d i s s e c t i n g and

r e f i n i n g the

c a n we e x p e c t t o u n r a v e l t h e t a n g l e

w h i c h c u r r e n t l y a r e th o u g h t t o have a b e a r i n g

i m p o r t a n t but n e g l e c t e d a r e a

on t h i s

of

highly

of m u t u a l i n t e r e s t to t h e n u t r i t i o n i s t

and the physiologist.
As

c a n be d e d u c e d f r o m the f o r e g o i n g d e v e l o p m e n t

background
m ore

for this th esis,

u n i f i e d a nd c o h e r e n t

its m a in objective is to a r r i v e

posited
To

f r o m th e t i m e

i n the

stu dy

skeleton,

it i s p l a c e d in th e

how f a s t t h e

radioactive

c r o p u n t i l it i s d e ­

and u n d e r tw o d i f f e r e n t n u t r i t i o n a l r e g i m e s .

' ' a b s o r p t i o n 1' one m u s t know how m u c h

f e r e d to t h e a b s o r p t i v e

at a

c o n c e p t of ' ' a b s o r p t i o n ' 1 by c o n s i d e r i n g

e x p e r i m e n t a l l y w hat h a p p e n s to a s in g l e g i v e n m a t e r i a l ,
strontium ,

of the

intestinal m ucosa.

crop e m p ties,

one m u s t

strontium

is

of­

Since t h i s d e p e n d s upon

study t r a n s i t

rates.

Then,

the m a n n e r

in w h i c h s t r o n t i u m d i s a p p e a r s f r o m t h e a b s o r p t i v e

ment

b o w e l w i l l be d e s c r i b e d .

the
to

of the

difference

Since t h e blood l e v e l s

b e t w e e n input and output,

c o n s i d e r a t l e a s t the m a j o r

site

seg­

reflect

it w i l l t h e n be n e c e s s a r y

of s t r o n t i u m d e p o s i t i o n , n a m e l y

26
the

skeleton,

sorbed

in t e r m s

strontium .

When t h e

s hould be p o s s i b l e t o
quite

a number

of i t s

ability to
analysis

critically

of f a c t o r s

c l e a r the
of t h e s e

e v a l u a t e the

broader

ab­

d a t a a r e c o m p l e t e d , it

relative

im portance

specific for

but t o b r i n g f o r t h c e r t a i n o t h e r c o n c e p t s

significance.

of th e

of

w h i c h c o n t r i b u t e to i n t e s t i n a l a b s o r p t i o n .

It i s hoped not only t o o b t a i n i n f o r m a t i o n
strontium ,

plasm a

radioactive

of c o n s i d e r a b l y

EXPERIM ENTAL

METHODS

D e s c r i p t i o n of G e n e r a l P r o c e d u r e s

White L e g h o r n c o c k e r e l s ,
purchased from

of unknown g e n e t i c b a c k g r o u n d

local c o m m e rc ia l h a tc h e rie s,

were

u s e d in a l l of

the t e s t s

c o n d u c t e d e x c e p t i n the f i n a l

experim ent.

c a s e , the

chicks w e re p u rc h a s e d fro m

a localpoultry

chicks when p u r c h a s e d
placed

were

one day

of age and w e r e

on t h e f e e d i n g r e g i m e to be t e s t e d .

The

w a s u s e d t h r o u g h o u t th e e x p e r i m e n t a l p e r i o d ,
cillin

regim e,

c o n tro l ratio n.
regim es

at which tim e
T he

studied a r e

ra tio n was

same

latter

breeder.

The

im m ediately
b a s a l ration

e x c e p t f o r the p e n i ­

p e n i c i l l i n w a s not in c lu d e d in the

c o m p o s i t i o n of th e b a s a l r a t i o n a n d the f e e di n g
given in T able

c a l c u l a t e d to be

1.52 p e r

Feedin g

II.

The l e v e l of c a l c i u m in t h i s

cent.

R egim es

Studied

Basal
B a sa l plus

25 p e r

c ent l a c t o s e

B a s a l pLus

15 p p m p r o c a i n e

1 Substituted

In t h e

1

penicillin

at e x p e n s e

G

of c o m p l e t e

27

ration.

28
T A B L E II
C O M P O SI T IO N OF

BASAL RATION

Item

Ground

y e ll o w c o r n .......................................................................................................

Ground heavy oats
Wheat b r a n
Wheat

...............................................................................................

......................................................................................................................

sta n d a rd m iddlings

Alfalfa m e a l,

dehydrated,

M e a t and bone

scraps,

S o y b e a n oil m e a l
Red fish m eal
D ried

Pounds

............................................................................
17 p e r

50 p e r

centpro tein

..........................

dried

5
5
5
5

c e n t p r o t e i n ..................................

5

.......................................................................................................

20

...........................................................................................................

W h e y ......................................................................................................................

B rew ers

45

2.5
2.5

y e a s t ...............................................................................................

2.5

G r o u n d l i m e s t o n e .......................................................................................................

1.5

S t e a m e d b o n e m e a l ...................................................................................................

1.0

S a l t ( i o d i z e d ) ..................................................................................................................

0.3

V i t a m i n A a nd D f e e d i n g

0.25

M anganese
Choline

2250 A,

300 D

....................

s u l f a t e , f e e d i n g g r a d e .........................................................

chloride,

V itam in B jg

B

oil,

25 p e r

c e n t d r y m i x t u r e ..................................
1, 2
ancl a n t i b i o t i c f e e d s u p p l e m e n t ( M e r c k )
.

^ E a c h pound c o n t a i n s n ot l e s s t h a n 3 m i l l i g r a m s
a n d 1.2 g r a m s of c r y s t a l l i n e p e n i c i l l i n G.
12
5
2
R e m o v e d f r o m t h e d i e t f o r p e n i c i l l i n study.

0.02
0.15
0.05

of V i t a m i n

29
The
raised

chicks w ere

s c re e n floors.

reared

in an e le c tric a lly heated b attery

The t e m p e r a t u r e

u n d e r the h o v er

w a s t h e r m o s t a t i c a l l y m a i n t a i n e d a t 95° F. a t a l l t i m e s .
was

located in the a n im a l room

a constant

room te m p e ra tu re

were

of 75° F.

followed d u rin g the

of th e b a t t e r y
The b a t t e r y

of t h e P h y s i o l o g y D e p a r t m e n t w h e r e

l i g h t d a y of 14 h o u r s wa s p r o v i d e d .
tices

with

w a s m a i n t a i n e d and a c o n s t a n t

Recommended m anagem ent p r a c ­

g ro w i n g p e r i o d w ith f ee d and w a t e r

p ro v id ed ad libitum .
In the f i r s t
m e n te d at 2 5 p e r
ject

of t h e s e

would h av e

of e x p e r i m e n t s the e f f e c t

c e n t of t he

ration was

e x p e r i m e n t s w a s to

on t h e t r a n s i t

and in t u r n ,
would h av e

series

studied.

r a t e of Sr

of S r

90

90

t h r o u g h the

by t h e

of Sr

90

intestinal tra c t,

were

system .

With t h i s

s e t up to d e t e r m i n e

t r a n s i t t h r o u g h the t r a c t p e r unit t i m e .
7-1/2
15
30
45
60
90
120
180
240
1440

ob­

r a t e , if one e x i s t e d ,

skeletal

c o n c e p t in m i n d , the f ollowing t i m e p e r i o d s
th e p r o g r e s s

The p r i m a r y

supple­

se e what e f f e c t t h i s l e v e l of l a c t o s e

what e f f e c t a n i n c r e a s e d t r a n s i t
on th e u p ta ke

of l a c t o s e

minute
minute
minute
minute
minute
minute
minute
minute
minute
minute

g r oup
g r ou p
group
group
group
g r oup
group
group
g roup
group

30
The
the

l e n g t h of t i m e

r e f e r s to the i n t e r v a l b e t w e e n the t i m e

c h i c k w a s i n j e c t e d w i th S r
Sim ilar

groups were

90

and t h e t i m e

u s e d i n th e

a 50 m i n u t e

group

1440 m i n u t e

group was elim inated.

of e x p e r i m e n t s ,
provide

r e p l a c e d the 45 and

adequate

series

w ere t r e a t e d the

and the

in b oth s e r i e s
u s e d to

chicks w ere

old, while t h o s e u s e d i n t h e p e n i c i l l i n

s a m e way at

14 t o

18 d a y s

of a ge .

of t e n to t w e l v e b i r d s

It w a s found t h a t

sixty

chicks

so t h a t

g r o u p s e a c h of c o n t r o l a nd e x p e r i m e n t a l b i r d s w e r e

selected.
chicks

r e m o v e d f r o m th e b a t t e r y a s a g r o u p , a s m a l l i n c i s i o n w a s

m ad e th r o u g h the
aqueous

skin just

s o l u t i o n of S r

30 m i l l i g r a m s
was

The

could be p r o c e s s e d i n one

T he d ay follow ing the w e ig hin g and w i n g - b a n d i n g the
w ere

w e ig h e d

w e i g h t would be a p p r o x i m a t e l y e q u a l w i t h i n a f e e d i n g

d ay ; t h u s t h r e e
usually

groups,

F o r all groups

of e x p e r i m e n t s the

c h i c k s w e r e t h e n d i v i d e d into g r o u p s

regim e.

60 m i n u t e

however,

com parisons.

a n d w i n g - b a n d e d w h e n 14 d a y s

the a v e ra g e

study of p e n i c i l l i n ;

a n e q u a l n u m b e r of c o n t r o l c h i c k s w e r e

In t h e l a c t o s e

series

of c h i c k s a c r i f i c e .

90

of E v a n s

Cl

o v e r the

2

c r o p , and a

, c on ta in in g

B lue,

accurate

injection,

m illiliter

10.23 pc r a d i o a c t i v i t y

w a s i n j e c t e d into th e

c o n s i d e r e d n e c e s s a r y to e x p o s e the

to i n s u r e

\/Z

and

crop lumen.

c r o p by i n c i s i o n i n o r d e r

and in a d d i t i o n , the

c o n di t i o n of the

It

31
crop

could be

observed.

c lu d e d t h e a m o u n t
crop

contraction,

cedure.

O bservations

of f ee d p r e s e n t ,
which was

The effects

c e p t in th e

case

of the

c h i c k s a g a i n had f r e e

presence

chicks were

7-1/2

and

a c c e s s to f e e d and w a t e r .

a large

pair

e a c h c h i c k.

carried

so t h a t a l l c h i c k s
same

A blood

Im m ediately
in the

alim entary tr a c t

prevent fu rth er movement

one m i n u t e .

later.

battery

(ex­

w h e r e the

This p ro c e d u re

routine

The

of the

chicks

only

after

c h i c k s w e r e t h e n killed

of any one g r o u p w e r e

a m o u n t of t i m e .

out by r e m o v i n g the

of s h e a r s .

k il l i n g t o t y i n g

recorded.

s o l u t i o n the

o p e n in g w a s m a d e
a n d t he

carefully

order

t o t he i n j e c t e d
chicks was

and

c h i c k w a s i n j e c t e d , th e wingband n u m b e r and t i m e

of i n j e c t i o n w e r e
same

of g a s ,

injection p r o ­

r e t u r n e d to the

15 m i n u t e g r o u p s ) ,

in ­

a m o u n t of t i m e .

As each

in the

or absence

c o n d i t i o n w ill be d i s c u s s e d

r e s t r i c t e d t h e n o r m a l e a t i n g and d r i n k i n g
a m inim um

c r o p at t h i s t i m e

o ften o b s e r v e d d u r i n g the

of c r o p

A f t e r i n j e c t i o n th e

of the

sample

head in one swift

c o l l e c t i o n of the blood

snip w ith

gizzard

sam ple,

an

was exposed,

l ig ate d j u s t p o s t e r i o r to the g i z z a r d to

of the

off the l o w e r

of the

w as t h e n c o l l e c t e d f r o m

a b d o m i n a l r e g i o n , the
was

Killing

exposed

gut

Sr

90

.

The

complete

seg m en t usually

operation fro m

required

le s s than

32
Preparation

A sam ple
m ents

was

From

the

placed

of whole blood,

blood

collected,

oven at

oughly a s h e d a t
activity

90
600

o

nuclear

same

The

(see the
bone

sample

A f t e r d r y i n g th e

C.

i n a m uff le f u r n a c e .

l o w e r gut

and

the a c t i v i t y

of whole blood.

dried,
th e

The

fem ur

and a s h e d

sam ple

fem ur

was

T h e d e g r e e of

T his was l a t e r

dose p e r m illilite r
the i n j e c t i o n

collected

set-up
per

described

second

so lu ti on and

was the

a s h e d , the

a m o u n t of S r
previously.

per fem ur.

Sr

90

c o u nt s p e r
c o n v e r t e d to

of blood by c o m p a r i s o n
c ounted in the

T his

90

bone

36).

r i g h t f e m u r w h ic h w a s
w e ighed b e f o r e

18 m i l l i l i t e r

s a m e m a n n e r a s the blood

com pletely

p o w d e r a nd the

was then th o r -

r e c o r d e d in

w a s t h e n p l a c e d in a n

in the

and

d r i e d t h o r o u g h l y in

c l e a n e d t h o r o u g h l y upon r e m o v a l a nd i m m e d i a t e l y
drying.

present.

w as e x t r a c t e d

s e c t i o n on d e r i v a t i o n of d a t a , p age

sample

90

seg-

w ith a t h i n end window G M tube and a

standard made from
way

1 m illiliter

0 porcelain crucible,

c e n t of a d m i n i s t e r e d

with a

u p p e r and

C.

scaler,

second p e r m i l l i l i t e r
per

a

was determ in ed

standard

bone,

c o l l e c t e d and p r e p a r e d f o r t he a m o u n t of Sr

in a (18 m l . ) s i z e

a drying

of S a m p l e s

crucible,

sam ple.

When

was g r o u n d into a fine

activity d e term in ed

with the

The

a c t i v i t y w as o b ta in ed a s

w as

c o n v e r t e d into p e r

counting
co u nt s

c e n t of i n j e c t e d

33
dose

in the

com plete

skeLeton by m e a n s

This

fa c to r is explained

l a t e r in the

of a Linear

section,

c o r r e c tio n facto r.

' ' D e r i v a t i o n of the

D a ta .''
The

com plete

a n t e r i o r end
and

of t h e

separated

d i g e s t i v e t r a c t and i t s

e s o p h a g u s and e nding a t th e

j u s t p o s t e r i o r to t h e g i z z a r d .

were tre a te d

sim ilarly

removed,

wet w eight w a s d e t e r m i n e d .

its

placed, in a W a r i n g
t o t a l w e i g h t to
liter

sam ple

a n d the

in e v e r y

respect.

A fter

of t h e

sam ple

t h e n a dd ed and the

sam ple

90

Waring
ment

sample

made

The

Evans

c o n t e nt ,

cent n i t r i c a c i d w as

sample

A fter

w a s d i lu t e d with

The d i l u t e d

sample

w as

m i l l i l i t e r aliquote taken for d e te r m in a tio n

m easured

by i n j e c t i n g the

activity
same

was t h e n c o m p a r e d

amount

of Sr

B l e n d o r w h i c h c o n t a i n e d e i t h e r the u p p e r

of a n o n i n j e c t e d

Blue

1 m illi­

w a s a ll o w e d to c o m p l e t e l y d i g e s t .

of 2 50 m i l l i l i t e r s .

a nd a 4 - 1 / 2

activity.

was then

homogenization, a

of 50 p e r

d i g e s t i o n had t a k e n p l a c e the

standard,

The

s e g m e n t w as

w a s p l a c e d in a 2 50 m i l l i l i t e r

Fifty m i l l i l i t e r s

of S r

rem oved

T h e s e two gut s a m p l e s

A s e a c h gut

complete

E r l e n m e y e r flask.

th e n m ixed

cloaca, was

w a s t a k e n f o r d e t e r m i n a t i o n of the

w a t e r to a v o l u m e

s t a r t i n g a t the

B l e n d o r w i th e n o u g h w a t e r a d d e d to b r i n g the

50 g r a m s .

rem ainder

com plete

contents,

c h ic k and

90

or

CL

w ith a

into a

l o w e r gut

s u f f i c i e n t w a t e r to t o t a l

seg­

50 g r a m s .

The

same

procedure

activ ity in the

w a s t h e n u s e d to d e t e r m i n e t h e a m o u n t

standard

A f t e r th e

a s in the e x p e rim e n ta l

sixth e x p e rim e n t,

m e t h o d w a s n ot r e l i a b l e ,
determ ination.
t h e r e f o r e , the
in to t h e
the

gut

w hen it b e c a m e

The

o bvious t h a t t h i s
Evans

Blue

s a m p l e s n e e d not be h o m o g e n i z e d

s a m p l e s a f t e r be in g w e ig h e d w e r e

E r l e n m e y e r f l a s k an d d i g e s t e d .

same.

sam ples.

it w a s d e c i d e d to e l i m i n a t e the

T h i s m e a n t t h a t the

of S r 9°

placed d irectly

O ther p ro c e d u re s

rem ained

c o n v e r s i o n of v a l u e s f r o m th e f i r s t p r o c e d u r e to the

l a t t e r i s fully d e s c r i b e d i n t h e Appe ndix.

D e s c r i p t i o n of A d d i t i o n a l T r i a l s

Two additio nal e x p e rim e n ts
t i o n of t h e

series

m anagem ent,

just d escrib ed .

environm ental

were

conducted a f t e r the t e r m i n a ­

In th e f i r s t e x p e r i m e n t , the

conditions,

etc., were

chicks

s i m i l a r to t h o s e

previously described.
The b ird s

were

w eigh ed a nd w i n g - b a n d e d at

T h e y w e r e t h e n d i v i d e d into g r o u p s
m etabolism

brooding la m p s.

The

g iv e n ad l i b i t u m .

w a s m a i n t a i n e d at 95° F .

chicks

of a ge .

of f o u r and p l a c e d i n s m a l l r a t

c a g e s w h e r e feed a n d w a t e r w e r e

environm ental te m p e ratu re

13 d a y s

r e m a i n e d in the

The

by i n f r a - r e d

cages for 2 days,

at

w h i c h t i m e t h e y a p p e a r e d t o be w ell a c c u s t o m e d to t h e i r n ew e n ­
vironm ent .

35
The
back in the
placed

chicks w ere
cage.

injected as d e s c rib e d p rev io u sly ,

E xcreta

was

c o l l e c t e d a s vo id ed , w e i g h e d and

in a c o n ta in e r fo r futu re

4 h o u rs the

chicks w ere
described.

The

organ tis su e s

removed,

an d the

w ater for a

Sr

g a s tro in te s tin a l t r a c t was tr e a te d

At th e

When the m u s c l e

was easily

r e m o v e d f r o m t he

and s k e l e t a l t i s s u e

was

dried

in a d r y i n g

were

skel­

completely

oven o v e r n i g h t ,
at

600° C.

The

s k e l e t o n w a s t h e n g r o u n d into a p o w d e r and d i s s o l v e d w ith

c o n c e n t r a t e d HC1.

The

t o 250 m i l l i l i t e r s .

A Sr

i l a r t o the
per

soft

c o n c l u s i o n of t h i s p e r i o d it

a n d t h e n c o m p l e t e l y a s h e d i n the m u f fl e f u r n a c e
ashed

A fter

c h i c k w a s t h e n p l a c e d i n boiling

30 m i n u t e p e r i o d .

s e p a r a t e d , t h e bone t i s s u e

activity d eterm in atio n .

c h i c k s w e r e t h e n s k in n e d , a l l t h e

w a s f ound t h a t t h e m u s c l e t i s s u e
etal tissu e.

90

k il l e d , the

as previously

and p l a c e d

bone
90

s o l u t i o n w as t h e n d i l u t e d w i th t a p w a t e r

d e te rm in a tio n was then c a r r ie d

out

gut d e t e r m i n a t i o n p r o c e d u r e p r e v i o u s l y d e s c r i b e d .

sim ­
The

c e n t u p t a k e by t h e t o t a l s k e l e t o n could t h e n be c a l c u l a t e d by a

com parison

of t h e s e

v a l u e s w it h the

standard

value.

D e t e r m i n a t i o n s i n t h i s e x p e r i m e n t i n c l u d e d f e e d a nd w a t e r
intake, e x c r e ta

output,

p r e s e n t in e x c r e t a ,
The
chicks w ere

m oisture

c o n t e n t of e x c r e t a ,

per

^

ce nt Sr

90

u p p e r gut, l o w e r gut, a nd s k e l e t o n .

second t r i a l was very

s i m i l a r to the

placed, i n i n d i v i d u a l c a g e s

so t h a t

f i r s t e x c e p t the

separate

values for

36
t h e e x c r e t a , could be

computed.

w ith the m u s c l e t i s s u e
tissue

intact,

In a d d i t i o n t h e
since Sr

90

skeleto n was ash e d

activity

of t h e m u s c l e

w a s f ound t o be v e r y l o w f o r t he 4 h o u r g r o u p in t h e p r e -

vious e x p e rim e n t.
a n d l o w e r gut,

O bservations

skeletal

system ,

included Sr

90

a c t i v i t y of t he u p p e r

a nd t h e e x c r e t a f o r 2 and 4 h o u r

groups.
The

data

d e t e r m i n e the
and the uptake

collected

i n t h e s e two t r i a l s p r o v i d e d m e a n s to

e f f e c t of d i f f e r e n t h an d l i n g m e t h o d s
of S r

90

by th e

skeleton.

on t r a n s i t

rates

T his is d is c u s s e d la te r.

In a d d i t i o n , w a t e r and f e e d i n t a k e a nd e x c r e t i o n w e r e m e a s u r e d
and a fa c to r was d eterm in ed
of t h e f e m u r t o t h a t of t h e
could be

co n v erted to p e r

show in g th e

entire

90

i n the

skeleton.

of t h e

separate

90

activity

T h e d a t a and

found in t h e Appe ndix.

A d e s c r i p t i o n of a l l c a l c u l a t i o n s , th e f o r m u l a e
lustrations

of S r

s k e l e t o n so t h a t a l l f e m u r a c t i v i t i e s

c e n t of Sr

c a l c u l a t i o n of t h i s f a c t o r a r e

relationship

calculations a r e

used, and i l ­

found i n t h e A ppendix.

D e r i v a t i o n of th e Data

The

o b se rv e d e x p e rim e n ta l values usually

of c o r r e c t i o n s

of v a r i o u s

the b a s ic data fo r th e se

required a num ber

k in d s b e f o r e t h e y could be
experim ents.

reg ard ed as

T his was p a r tic u la r ly tru e

37
of t h e

radioactivity m e a s u re m e n ts ,

w h i c h c o n s t i t u t e t h e b u l k of t he

observations.
The

rad io activ ity data a r e

injected dose
H ansard,
of Ca

45

recovered

C om ar,
in the

from

v a r i o u s p o r t i o n s of the

and P l u m l e e

rat,

best expressed as p er

(1951) h ave

if e x p r e s s e d a s p e r

concurrently

t h e n only if t h e

u n less m o r e than

rats

are

c en t of i n j e c t e d d o s e , i s
carrier

calcium ad m in ­

10 m i l l i g r a m s

on a low c a l c i u m d ie t.

t a i n i n g n o r m a l a m o u n t s of c a l c i u m , a s m u c h a s
nonradioactive

chick.

shown t h a t f e c a l e x c r e t i o n

n ot a f f e c t e d by t h e a m o u n t of n o n r a d i o a c t i v e
istered

ce nt of the

c a l c i u m g i v e n o r a l l y w i t h the

i s give n, a nd

On a r a t i o n c o n ­
50 m i l l i g r a m s

of

radio active t r a c e r failed

to a l t e r t h e f e c a l e x c r e t i o n , if th e e x c r e t i o n w a s e x p r e s s e d a s p e r
cent

of d o s e .

Of c o u r s e , t h e r e

the n o n ra d io a c tiv e
There
curie

is

calcium

l e v e l in th e f e ed .

only 0.0063

m icrogram s

of t h i s t r a c e r and it i s

i s 4.9 X 10

-10

was usually

pg of Y
of t h e

ent, f i r s t , th at tr u e

90

per

i s a m a r k e d e f f e c t a s s o c i a t e d w ith

of a c t u a l Sr

c a r r i e r free.

pc of Sr

90

.

s e c o n d , t h a t t h e a c t u a l Sr

are

expressed

as per

cent

90

dose

Sr

90

Since the i n j e c t e d d o s e
chick,

e x p e rim e n ts w ere actually

and

in a m i c r o -

A c c o m p a n y i n g the

o r d e r of 10 m i c r o c u r i e s p e r
tracer

90

it i s a p p a r ­
c o n d u c te d,

w a s not c r i t i c a l , if the d a t a

of i n j e c t e d d o s e .

38
The
roughly

a c tu a l e x p e r im e n t a l work pro c eed ed

18 m o n t h s ,

different

during which tim e

c ou n t i n g e f f i c i e n c i e s w e r e

s i b l e to p r o c e s s th e
most

of t h e

large

c o n v e r s i o n of th e

different

employed.

m asses

counting t u b e s w ith
Sinc e it w a s i m p o s ­

c r u d e m e a s u r e m e n t s w a s done a f t e r
It i s th e p u r p o s e

s e c t i o n t o o u tlin e th e a r i t h m e t i c w h i c h w a s n e c e s s a r y .

actu a l data which

s u p p o r t th e

of

of d a t a a s t h e y a c c u m u l a t e d ,

t h e e x p e r i m e n t a l w o r k h ad b e e n c o m p l e t e d .
this

over a period

conversion fac to rs

of

The

e m p l o y e d have

b e e n p l a c e d i n t h e A p pe nd ix .
1.

Background

c o r r e c tio n s w ere always m ade

a t the t i m e

of c o un ti n g .
2.

D e c a y c o r r e c t i o n s f o r Sr

necessary,

s i n c e th e h a l f - l i f e

p e rio d , th is am ounts to a lo s s
Since

90

c o n s i d e r e d u n-

i s 25 y e a r s .

90

in Sr

decay w ere

activity

In a n

of 4.0 7 p e r

18 m o n t h
cent.

s t a n d a r d s w e r e p r e p a r e d i n t e r m i t t e n t l y d u r i n g t h i s t i m e , the

actual e r r o r
3.

could not have e x c e e d e d h a lf t h i s v a l u e .

C o r r e c t i o n s f o r the

In five h a l f - l i v e s ,
half-life
days

of S r

90

of Y

before

correction.

90

is

g r o w t h of Y

growth is

97 per

90

from

H ow ever,

sam ples,

s i n c e the

Y

or else
90

90

cent c o m p l e t e .

2.54 d a y s , t h i s would r e q u i r e

c ou n ti ng t h e

Sr

Since the

e i t h e r waiting

12.7

m a k i n g the a p p r o p r i a t e

count is about t w o - t h i r d s

t h e t o t a l c o u n t, t o t a l a c t i v i t i e s m e a s u r e d a t f o u r h a l f - l i v e s

of

(10.2

39
days)

are

only 4 . 5 p e r

At t h r e e

half-lives

negative

8.3 p e r

It i s
of S r

90

to be

cent

Low.

T his was

(7.6 d a y s ) th e

regarded

as acceptable.

corresponding e r r o r

would be a

c en t.

r e c o g n i z e d t h a t t h e e f f e c t of f a i l u r e to

.
p lu s the f a ilu r e to c o r r e c t fo r

Y

90

c o r r e c t for decay

g r o w t h m i g h t be e x p e c t e d

c u m u l a t i v e , w h i c h would r e s u l t at the w o r s t in t o t a l

10 p e r

c e n t low.

This negative

s i n c e b o t h e x p e r i m e n t a l and
concurrently

control groups

for

Sr

90

decay,

realized,

control groups were

(w ith in a day).

and

bias was n e v e r

T h u s no d i s p a r i t y

c ould p o s s i b l y a r i s e

a l t h o u g h b oth m i g h t be

c o n s id e r a tio n holds for the effect

of Y

always

recoveries
however,

co un te d

between e x p e rim e n ta l

f r o m the f a i l u r e to c o r r e c t
s li g h t l y low.

90

The

same

g r o w t h , a l t h o u g h w i th l e s s e r

force.
That equilibrium
achieved

in th e t i m e

of Y

w ith i t s p a r e n t ,

b e t w e e n killing and the

p r e p a r a t i o n is illu s tra te d
first

90

in Table

a c t i v i t y d e t e r m i n a t i o n gave

A n o th er aliquot

rem oved

p e r aliquot, an in c r e a s e
activity

of 166 c o u n t s p e r

tw e lf th day,
count.

90

, was esse n tia lly

c o m p l e t i o n of s a m p l e

XVIII (in the Appen dix).

159 c o u n ts p e r

Z0 d a y s l a t e r gave
of only

Sr

1.9 p e r

s eco nd p e r a l iq u ot .

162 c ou n ts p e r

cent.

s e co n d

An a b e r r a n t high

s e c o n d p e r a liq uo t w a s m e a s u r e d

but e v e n t h i s w a s

The

in the

only 4.4 p e r cent above the i n i t i a l

40
Since t h e

blood

d r y i n g a nd d r y a s h i n g
bias

sam ples

and at l e a s t

s h o u l d be fo und i n t h e m .

trem ely

sm all percentage

0.1 p e r

cent p e r m i l l i l i t e r

values w ere
These

rarely

points,

further

Y

90

.

half-lives,

(never m o re

If t h e

light of th e f a c t t h a t t h e y w e r e

period make

it e v i d e n t t h a t d r a m a t i c

i s not lik e l y to r e s u l t f r o m

Y
Y

90

90

d e c a y in t i s s u e s
p l a c e d in t h e

w h ic h have a b s o r b e d

c r o p w e r e a b s o r b e d and

w h i c h did not a c c u m u l a t e

c o u n te d.

This might

if the t o t a l q u a n t i t y

of Y

90

Sr

90

require

, the e a s i e s t

to d e c a y b e f o r e

f r o m f o u r to t e n Y

accum ulated

were

(1955), m u c h l e s s t h a n

m inistered

i s a b s o r b e d f r o m the d i g e s t i v e t r a c t .

quently,

in t h e s e

e x p e r i m e n t s , the t i m e

c o u n t i ng i s m o r e t h a n
bility .

1 per

90

c o r d i n g to P e a r s o n
Y ttrium

than

In a d d i t i o n , th e blood

c o n f u s i o n would be to a l l o w the Y

sam ples were

g reatest percentage

of t e c h n i q u e ,

in a t i s s u e

way t o a void
the

90

(after a i r

c o n t r o l and e x p e r i m e n t a l g r o u p s .

of t h e blood t i m e - c u r v e s

C orrections for

accum ulated

7 d a y s ) , th e

of whole blood).

c o n s i d e r e d in th e

and not S r

counte d f i r s t

H o w e v e r , the blood c o n t a in e d a n e x ­

d i f f e r e n t in t h e

refinem ents
4.

always

of the i n j e c t e d a c t i v i t y

counted within a sh o rt tim e
im provem ent

were

large.

90

Ac­

cent of o r a l l y a d ­
Conse­

e l a p s i n g b e t w e e n k illing and

s u f f i c i e n t to a vo id c o n f u s io n f r o m t h i s p o s s i ­

41
5.

SeIf-absorption co rrectio n s.

m in e d at infinite t h i c k n e s s
or were

(intestinal

segm ents

c o r r e c t e d t o in fi n it e t h i c k n e s s by the

tion f a c to r s
activities,

experim entally determ ined

e x p re s s e d as p e r

of c o u n t s p e r

A ctivities w ere

cou nted in s olu tio n )
u se

of l i n e a r c o r r e c ­

( f e m u r and blood).

ce nt of i n j e c t e d d o s e ,

T h u s , the

r e f e r to the

s e c o n d p e r i n f i n i t e l y t h i c k l a y e r of t i s s u e

to

s e c o n d p e r i n f i n i t e l y t h i c k l a y e r of s t a n d a r d , m u l t i p l i e d by
A l l of t h e
of S r

90

to p e r

-Y

90

sk eletal data a r e

found i n th e

c e n t of t o t a l i n j e c t e d d o s e

T he f a c t o r f o r t h i s
of t h i r t y - t w o
activity

ash from

taken.
the

-Y

90

solution.

the activ ity

of the

to ta l activity
a c t i v i t y in t h e

The

s k e le t o n .

a nd the
The

a s h f r o m th e t o t a l

cent n i t r i c a c i d ,

and a 4.5 m i l l i l i t e r a l iq u o t

ratio

of t h e gut

b e t w e e n th e

aliquot gives a facto r

whole

s e co n d

in t h i s a l i q u o t wa s t h e n d e t e r m i n e d in

radioactivity

in the whole

100.

determ ined as described e a rlie r.

of 2 50 m i l l i l i t e r s

m a n n e r a s the

co u n t s p e r

A series

for 4 or Z hours,

T he t o t a l a s h w as d i s s o l v e d i n 50 p e r

The t r a c e r activ ity

injection

90

ratio

converted

r e c o v e r e d f ro m the to ta l

g i v e n Sr

right f e m u r s

up t o a v o l u m e

same

data were

e a c h f e m u r w a s t h e n c o m p o s i t e d with the

skeleton.
made

These

c o u nt s p e r

c o n v e r s i o n w a s d e t e r m i n e d a s f o l l ow s .

chicks w ere

of t h e i r

b a s e d on the

rig ht fe m u r.

either d e te r ­

a nd of the

single f e m u r a c t i v i t y a nd
(8.84 ± 0.13) by which the

s k e l e t o n c a n be

skeleton is

sam ples

calculated.

c o n v e r t e d to p e r

F i n a l l y , the

c en t of i n j e c t e d

42
dose

by d i v i d i n g by th e

jection

s o l u t i o n in t h e
Table

activity
same

of a s t a n d a r d

factor

8.84 ± 0.13.

per

c e n t.

It c o r r e c t s

g i v e s t h e b a s i c d a t a l e a d i n g to

The e r r o r

in t h i s f a c t o r i s

s k e l e t o n d i s s o l v e d i n 250 m i l l i l i t e r s
tion, at infinite t h i c k n e s s .

The

c o r r e c tin g p r o c e d u r e s , then,
infinitely th ick aliquot.

dose

counte d

moved

in s o l u -

f o r a l l the
on s u c h an

The

f in a l d a t a a g r e e

very

closely

mixing,

in c r u c i b l e s

The

in a n ov en at

(in c o u n ts p e r

c o n v e r t e d to p e r

i n j e c t i o n of 10.23 pc of Sr

of 50 p e r

com plete

and p l a c e d

of whole blood

second

cent of i n ­

of whole blood by a f a c t o r d e r i v e d i n the

A standard

A fter

c o u n ti ng blood.
dried

point of r e f e r e n c e

on the a s h ) w a s

p l a c e d i n 2 50 m i l l i l i t e r s
before.

and c ounted,

in je c te d activ ity d i s p e r s e d u n ifo rm ly in s o lu ­

per m illiliter

fo ll ow in g way.

as

base

3

of the whole

of J o n e s a nd Copp (1951) in p r e c i s i o n .

m illiliter,

jected

1.5

T he a l iq u o t r e p r e s e n t s a d e f i ni te f r a c t i o n

The m e a s u r e d activity
per

of HNO

is an activity d eterm in ed

t i o n by d i g e s t i o n w i th HNO^.
with those

le s s than

count on a 4.5 m i l l i l i t e r a l iq u o t

of the e n t i r e

in­

a count on th e p o w d e r e d a s h f r o m a single

bone t o a n e q u i v a l e n t

(4.5/250)

f o r th e

way.

XIX (in the A ppend ix )

the

prepared

cent HNO^ in the
1 m illiliter

of the

1 m illiliter

sam e

standard

90

-Y

same

sam ples

size
sample

was

manner

were

as those
w as

90

re­

u s e d in

completely

95° C. and t h e n a s h e d at 600° C. f o r

several

43
hours.

S in c e t h i s

activity

of t h e

be d i r e c t l y
liter

exact procedure

b lo o d

s a m p l e s , the a c t i v i t y m e a s u r e d in t h i s

c o m p a r e d w i t h whole blood v a l u e s .

sam ple

is

0.4 p e r

the t o t a l in je c te d d o se .

cent p e r m i llilite r .

T he

c an

1 m illi­

s o l u t i o n c on ta i n in g

Thus a d ire c t proportionality
c o u n ts p e r

case

However, the

c e n t of t h e 250 m i l l i l i t e r

w h i c h c o n v e r t s blood v a l u e s f r o m
to p e r

w a s followed, in d e t e r m i n i n g the

c a n be

s et up

second p e r m i llilite r

c a l c u l a t i o n of t h i s f a c t o r i s i l l u s t r a t e d

in T able

XX (in t h e A p p e n d ix ) .

It sh ou ld be n o t e d t h a t 4.5 m i l l i l i t e r

aliquots,

c o u n t e d wet, gave e x a c t l y the

sam e activity a s

1 m illiliter

c o u n t e d a f t e r d r y i n g an d a s h i n g .
6.
different
q u i r e d to

C o r r e c tio n s for different G M tubes.
c o u nt i n g t u b e s w e r e

are

listed
7.

S he lf a n d g e o m e t r y

n a m e l y t ube n u m b e r 2 - D I - 2 9 .

These

corrections.

Since a l l s a m p l e s w e r e

0 C o r r s p o r c e l a i n c r u c i b l e s p l a c e d on the b o t t o m
of t h i s type a r e

C o rre c tio n s for loss

sam ples.

w h ich would have b e e n o b ­

XXI (in t h e Appen dix).

no c o r r e c t i o n s
8.

gut

sta n d a rd tube,

in T able

counted in size
shelf,

u s e d , a s e r i e s of c o r r e c t i o n s w a s r e ­

c o r r e c t a l l a c t i v i t i e s to t h o s e

s e r v e d w i t h one

Since a n u m b e r of

It w a s

of a c t i v i t y d u r i n g h o m o g e n i z a t i o n of

soon evident that

j e c t i n g a known v o l u m e

required.

s t a n d a r d s p r e p a r e d by i n ­

of t h e i n j e c t i o n s o lu t i o n into f r e s h l y

i n t e s t i n a l s e g m e n t s f o ll o w e d by the h o m o g e n i z a t i o n a nd

removed

sampling

44
p r o c e d u r e s uniform ly
anticipated.

gave v a l u e s t h a t w e r e

Data in Table

XXII (in th e Appe ndix)

d i g e s t i o n w i th o ut p r e l i m i n a r y
B le n d o r g iv es the
130 c o u n t s p e r

20 per

a n t i c i p a t e d v alue

c en t l o w e r t h a n
sh o w t h a t wet

h o m o g e n i z a t i o n in the

(in the n e i g h b o r h o o d of 125 to

s e c o n d p e r a liq u o t) a nd t h a t h o m o g e n i z a t i o n i s a c ­

c o m p a n i e d by a f a i l u r e

to

obtain complete

recovery.

h o m o g e n i z a t i o n w a s due t o m a t e r i a l a d h e r i n g to the
and t h e

sam ple taken for Evans
The

data

indicated

of t h e f i r s t

correction

ization p ro c e d u re.

was

Blue

W a rin g B l e n d o r

of 0.8 w a s a p p l i e d , t h e n , to the gut
It w a s in t h e s e

sim ultaneously

When the E v a n s

to be i n a d e q u a t e ,

T h i s l o s s in

Blue a n a l y s i s .

six e x p e rim e n ts .

t h e a t t e m p t to u s e E v a n s

strated

W a r i ng

e x p e rim e n ts that

r e q u i r e d the h o m o g e n ­

Blue m e t h o d w a s

homogenization,

clearly d em o n ­

b e in g no l o n g e r n e c e s s a r y ,

also dropped,
9.

ash

c ounted

use

of t h i s
10.

rately

E xternal a b so rb er

In a f ew c a s e s , t h e bone

so r a p i d l y t h a t a n e x t e r n a l a b s o r b e r w a s r e q u i r e d .

T he

c o r r e c t i o n i s i l l u s t r a t e d in T a b l e XXV (in the Append ix ).
S ta n d ard s.

The

for each ex p e rim en t,

Sr ^ ° - Y ^ ^ .

corrections.

There

were

p r e p a r a t i o n a nd the l a s t
s t r o n g e r than those

injection

solutions w ere prepared,

but f r o m th e

same

s l ig h t d i f f e r e n c e s in the

stock

s h i p m e n t of

activity

s o l u t i o n u s e d w a s about 20 p e r

used e a r lie r .

These

standard

sepa­

of e a c h ne w
cent

values a re

listed

45
i*1 T a b l e

XXIII (in t h e A p pe nd ix ) .

counts p e r

sam ple

of i n f i n i t e t h i c k n e s s .

n o te d t h a t i t c a n not be
4.5 m i l l i l i t e r s , t h e

ard

concluded that

are

It should

e x p r e s s e d as

counts p e r

s a y,

second

s p e c i f i c a l l y be

s i n c e t h e a l iq u o t

a c t i v i t y w a s , l e t us

c on ta in e d

1 2 5 / 4 . 5 o r 27.8 c o u n ts

second p e r m illilite r.
11.

or

activities

second p e r aliquot; in o t h e r w o rd s, a s

f o r a l iq ui d

per

T he

som e

Statistics.

In m o s t

l i n e a r r e g r e s s i o n e q u a t i o n w ith i t s

errors

are

G enerally, these

the fin al s t a t i s t i c s
were

of t h e

being

corrected

r e c tio n fa c to r itself.

The

c a l c u l a t i o n of s t a n d a r d

standard e r r o r

c o n s t a n t s a nd t h e i r

upon w h i c h c o n c l u s i o n s a r e

The

stand­
based.

c o rre c tio n s w ere then applied

m e a n v a l u e s , w i t h due a l l o w a n c e

v a lu e

a m e a n a nd i t s

c o m p u t e d on v a l u e s a s c l o s e to th e o r i g i n a l

ob serv atio n s as was p e rm is s ib le .
to t h e s e

cases

and f o r t h e

a nd f o r the

products,

standard e r r o r

standard e r r o r

standard form ulae

errors

e r r o r s in s u m s , d i f f e r e n c e s ,

b oth f o r the

employed,

of t h e

cor­

b oth f o r the

c o m b i n a t i o n of s t a n d a r d

a nd q u o t i e n t s a r e

l i s t e d in the

Appendix.
To i l l u s t r a t e
Tables

XIV,

the nature

of t h e s e

a rith m e tic m anipulations,

XV, and XVI (in the Appe ndix) a r e

presented.

R E S U L T S AND DISCUSSION

L actose

O bservations made

E xperim ents

in the la c to s e

s t u d i e s i n c l u d e d live body

w eight , u p p e r and l o w e r gut wet w e ig h t s, f e m u r wet weight, f em u r
a s h e d weight, upper
(which w as

and l o w e r gut Sr

act ivity, f em ur Sr

90

ac tivity

c on v e r te d to to tal s k e le t o n S r 9 ° a ct ivity), and S r 9°

a c t iv it y p e r m i l l i l i t e r of whoLe blood.
discussed

90

in the

The o b s e r v a t i o n s w i l l be

o r d e r m ent ione d above.

Body Weight

The m e a n body weight af te r 2 w e e k s on a ration containing
25 p e r cent l a c t o s e w as
a difference

w hich i s highly significa nt (Table III).

parison s between la ctose
highly

10 p e r cent l e s s than i t s n o r m a l control,

and control groups, l a c t o s e

In eight c o m ­
c ause d a

s ig nif ic an t r e d u ct ion in s i x i n s t a n c e s , a r eduction of doubtful

s i g n i f i c a n c e in one c o m p a r i s o n ,
ference.

and in one c a s e , no significant dif­

Since the o v e r - a l l c o m p a r i s o n was based on m o r e than

s e v e n t y c h i c k s in e a c h group, it can be stated without r e s e r v a t i o n
that fe ed in g l a c t o s e

at 25 p e r cent of the tot al ration i m p a i r e d

n o r m a l growth.

46

47
TABLE

II I

E F F E C T O F L A C T O S E ON BODY AND G UT W E I G H T
(all v a l u e s g i v e n a s m e a n ± s t a n d a r d e r r o r )

N o . of
Chicks

T i m e a nd T r e a t m e n t

7.5 m i n . c o n t r o l g r o u p
7 .5 m i n . l a c t o se g r o u p

...........................
...........................

Body
We ight
(gm s.)

-

p 1
(pet.)

_2

-

10

93 ±

1.6

118 ±
114 ±

4.0
4.0

-

15 m i n . c o n t r o l g r o u p
15 m i n . l a c t o s e g r o u p

...............................
...............................

12

30 m i n . c o n t r o l g r o u p
30 m i n . l a c t o s e g r o u p

...............................
...............................

11
11

128
3.1
104 =fc 1.3

1

45 m i n . c o n t r o l g r o u p
45 m i n . l a c t o s e g r o u p

...............................
...............................

9
11

116
103

3.3
2.7

1

60 m i n . c o n t r o l g r o u p
60 m i n . l a c t o s e g r o u p

...............................
...............................

12
12

124 ± 5.6
103 ± 2.3

1

90 m i n . c o n t r o l g r o u p
90 m i n . l a c t o s e g r o u p

...............................
...............................

10
11

116 =fc 2.3
107 ± 4.0

6.2

9

NSD

120 m i n
120 m i n

control group
la c to se group

...........................
...........................

11

97 ± 3.0

180 m i n
180 m i n

control group
lactose group

...........................
...........................

10
8

103 ± 4.0
87 ± 2.0

1

240 m i n
240 m i n

control group
lactose group

...........................
...........................

9
10

102 ± 4.0
84 ± 4.0

1

73
93

119 ± 1.4
106
1.1

1

Total co ntrols
Total la c to s e
1
2

Statistical probability.

W e ig h t d a t a f o r t h e
w e re lost.

3

-

7.5 a n d

120 m i n u t e

NSD = no s i g n i f i c a n t d i f f e r e n c e .

control groups

48
TABLE

Uppe r Gut
p e r 100
g m s . Bo d y
W ei gh t

10.6

P
(pet.)

III ( C o n t i n u e d )

L o w e r Gut
p e r 100
g m s . Body
W e ig h t

P
(pet.)

T o t a l Gut
p e r 100
g m s . Body
W e ight

P
(pet.)

Z .6

1Z.Z

0.4

ZZ.8

1 .3

8.9 ± 0.8
10.4 ± 0.9

9-9
11.3

0.8
0.6

18.8 ±
Z1.0 ±

0.6
1.6

1

1

Z1 .0 db 0.6
Z3.8 ± 0,5

1

Z1 .8 db 0.4
Z4.5 ± 0.4

1

Z3.1 ±
Z1.3 ±

0.6
0.5

Z

Z3.5 ± 0.4
Z5.5 d: 0.4

1

NSD

NSD

1

9.3 ± 0.4
13.4 ± 0.7

0.4
0.4

1

11.6 ± 0.6
14.3 db 0.6

1

1Z.4 ±
13.8 ±

1.1
1 .0

NSD

10.7 db 0.6
11.8 ± 0.6

NSD

10.8 ±
1Z.8 ±

0.6
0.1

1

10.4 ±

0.5

11.7 ±
15.9 ±

0.9
0.6

10.3 ±
13.3 ±

1 O.Z ± 0.5
0.3
9-9
10.1 ±
10.6 ±

0.5
0.8

10.Z db 0.6
1Z.Z ± 0.6

1Z.8 ±
14.9 ±

0.6
0.8

13.3 ±

0.7

Z3.7 rfc 0.5

NSD

11.3 ± 0.4
13.0 ± 0.6

1.8

0.3
Z1.5
±
0.3
ZZ.9

1

NSD

1Z.3 ± 0.8
11.9 ± 0.7

NSD

ZZ.4 ± 0.5
ZZ.5 ± 0.5

NSD

1.9

10.0 ± 0.7
1Z.5 ± 0.6

1

ZO.Z ± O . Z
Z4.7 ± O.Z

1

3.6

49
T h is weight lo ss
the

energy

(1924)

in la c to se .

w a s p r o b a b l y due t o th e n o n a v a i l a b i l i t y
P lim m er

r e p o r te d that la c ta s e

w h ile

(1906) and H a m i l t o n and M i t c h e l l

i s found only i n t h e

Shaw (1913) found no l a c t a s e

a t a ll .

crop

S in ce t h e
ration,

lactose-fed

chicks

it would be n e c e s s a r y

cent m o r e

lactose

groups

lactose.

25 per

for th em to consume

It i s
chick.

c e n t of the
at le a s t 2 5 p e r

of f e e d c o n s u m p t i o n (T ab le IV) s h o w ed t h a t the
c o n s u m e d a bo u t th e

control group.

lactose-fed

s a m e t o t a l a m o u n t of f e e d a s

T h e r e f o r e , it w a s not s u r p r i s i n g t h a t the

chicks weighed l e s s .

In v i e w of th e h y d r a g o g u e t h e o r y
S utto n (1949), the
sidered.

(1953) d e m ­

of e n e r g y f o r t h e

c ould not u t i l i z e

chicken,

f e e d to e q u a l i z e t h e e n e r g y i n ta k e b e t w e e n the two g r o u p s .

A m easurem ent

their

i s not a s o u r c e

of the

R u t t e r e t al.

o n s t r a t e d t h a t t h e young c h i c k i s u n a b le to h y d r o l y z e
evident, then, th at la cto se

of

w ater balance

Roccuzzo

interefered

w ith the

of the b i r d s m u s t a l s o be

(1945) h a s p r e s e n t e d d a t a to
"absorption"

principle.

lactose

45 p e r

"retained"

c o n tro ls during a 4 hour p erio d

con­

show t h a t l a c t o s e

of w a t e r , w h ic h i s in g e n e r a l a g r e e ­

m e n t w i t h the h y d r a g o g u e
chicks

s u g g e s t e d by F i s c h e r and

In the p r e s e n t e x p e r i m e n t s , the

cent m o r e

w a t e r t h a n did t h e i r

(Ta ble IV), a l th o u g h m o s t

of the

r

excess
is

a p p e a r e d to h ave

s u b t r a c t e d f r o m th e

r e m a i n e d in t h e bowel.

body we ight, the

If the gut w eight

c a rc a ss averaged

99 g r a m s

50
TABLE
FOUR-HOUR

FEED

IV

AND WATER

OBSERVATIONS

T r eatm ent
Item
Control

Number

of c h i c k s ..............................................

W ater

intake p e r b i r d

Wa t e r

excreted

W ater

" r e t a i n e d 11 p e r b i r d ( m l.)

F eed

intake

2

per

( m l . ) ...............

per bird

(ml.)
. .

b ir d (gm s.) . . . . .

10

7

+ 64

7.70

+ 86

2.13

3.96

+ 46

2.57

3.74

- 2

4.1

4. 0

4.70

^ B i r d s w e r e g r o u p e d b y l o t s of f o u r ;
given a r e weighted a v e ra g e s .
2

U r i n a r y and f e c a l w a t e r l o s s

4.0 g r a m s of f e e d = 1.0 g r a m
1.0 g r a m of l a c t o s e i n c r e a s e s
1.0 g r a m of l a c t o s e i n c r e a s e s
in a f o u r h o u r t r i a l .
P ercentage

of w a t e r

1

therefore,

all v a l u e s

only.

of l a c t o s e .
w a t e r c o n s u m p t i o n by 3.0 g r a m s (ml.);
w a t e r " r e t e n t i o n ' ' b y 1.17 g r a m s (ml.);

c o n su m e d th at is

C o n t r o l = 2 . 5 7 / 4 . 7 0 X 100 = 54.7.
Lactose

Lactos e

P et.
Diff.

= 3 . 7 4 / 7 . 7 0 X 100 = 48.6.

retained:

51
in t h e

control groups,

U nfortu nately , the

but only

w ater

81 g r a m s

c o n t e n t of the

in the

lacto se-fed groups.

carcass

w a s not d e t e r m i n e d .

Gut Weight

The

u p p e r and lo w e r weight data

digestive t r a c t

of th e

lactose-fed

i n b o th u p p e r a nd l o w e r
every

case

were

w a s n o t e d in t h e

case

out of s e v e n , d i f f e r e n c e s
significant.

w a s highly

were

highly

The

control,

It i s i m p o r t a n t t o

the

result

A sim ilar
i n two

s i g n i f i c a n t , a nd i n two

contents,

(Table III)

show

cent h e a v i e r in the l a c t o s e -

r e m e m b e r t h a t the

of d i f f e r e n c e s

significant.

" g u t ” weights were

T h u s the d i f f e r e n c e

noted m i g h t

in the w e ig h t of the bowe l i t s e l f ,
o r in both.

sig­

in one of w h ic h it

combined data

w hic h w a s highly

contents included.

w e i g h t of th e

group

of s e v e n

s i g n i f ic a n t .

i n a l l c o m p a r i s o n s e x c e p t two,

chick th a n in its

be t h e

In

The t o t a l gut weight w a s h e a v i e r to a highly

significantly different.

t a k e n w ith t h e

lactose

of t h e l o w e r gut, w h e r e

t h a t t h e d i g e s t i v e t r a c t w a s a bo u t 20 p e r
fed

controls.

c o n t r o l g r o u p , and in t h r e e

m a d e , the d if fe re n c e

nificant d e g re e
was

significantly h ea v ie r,

e x c e p t one, t h e u p p e r gut w eight of the

com parisons

cases

chicks was

s e g m e n t s , t h a n t h a t of th e

w a s h e a v i e r t h a n t h a t of th e

difference

(Ta ble III) show t h a t the

F urther,

o r in

it w as not d e t e r ­

m i n e d w h e t h e r t he d i f f e r e n c e s w e r e a s s o c i a t e d p u r e l y

with w a t e r

52
content o r w h e th e r actu a l dry

w e i g h t d i f f e r e n c e s m i g h t have

tr ib u te d to the findings.

In v ie w of the f i n a l o u t c o m e

sorptive

data,

these

and t r a n s i t

rate

it did not

seem

con­

of the

ab­

p e r t i n e n t to p u r s u e

studies fu rth e r.

Bone

T he

Weight

w et w e i g h t of t h e f e m u r s

w a s found t o be g r e a t e r t h a n t h a t of
la c to se -fe d groups.

t a k e n f r o m the
fem urs

controL g r o u p s

r e m o v e d f r o m the

T h i s d i f f e r e n c e w a s highly

s i g n i f i c a n t ( T a b le V).

Since t h e f e m u r w e i g h t s w e r e a d j u s t e d t o m i l l i g r a m s p e r
body w e i g h t , t h e d i f f e r e n c e

100 g r a m s

n o t e d w a s o v e r and above t h a t a c c o u n t e d

f o r by

t he d i f f e r e n c e

i n body w e ig ht.

However,

a s h in

thef e m u r was

com pared, there

w a s no s i g n i f i c a n t d i f f e r e n c e

b e t w e e n th e l a c t o s e - f e d a nd t h e
r e d u c t i o n i n wet f e m u r
regim e
plu s
the

control groups

organic m a t t e r .

Lactose

size

of t h e

b i l it y

regim e

c a n n o t be

lactose

T h u s the

feeding

c o nte nt of w a t e r

i t s g e n e r a l e f f e c t on the

organism .

Although th e se
to se feeding

V).

fe ed in g d o e s not a p p e a r to in f l u e n c e

l a y - d o w n of bone m i n e r a l a p a r t f r o m

over-all

( T able

s i z e a s s o c i a t e d w ith t he

w a s due e n t i r e l y t o a r e d u c t i o n in the

w hen t h e w eight of

data w ere

were

what one m i g h t e x p e c t if the l a c ­

indeed a tis s u e

d eh y d ra to r, this p r o b a ­

s u b s t a n t i a t e d i n the a b s e n c e

of d r y

weight d a t a .

TABLE
EFFECT

V

O F L A C T O S E ON C A L C I F IC A T I O N

F eeding
Item

R egim e

Control

Control +
25% L a c t o s e

85

83

Mean fe m u r wet
w e i g h t p e r 100 g r a m s
bod y w e i g h t ( m g.) . . .

889 ± 12

827 ± 9

N u m b e r of s a m p l e s . .

20

20

Mean ashed fem u r
w e i g h t p e r 100 g r a m s
bod y w e i g h t ( m g.) . . .

115.3 ± 2.1

111.1 ± 2.2

Number

of s a m p l e s .

Differ enc e

62

4.2

P

1%

NSD

54
M o v e m e n t of S t r o n t i u m - 90

The fate
scribed,

of t h e

prim arily

loss from

injected

from

Table

radioactive

VI, in c h r o n o l o g i c a l o r d e r .

Thus,

the u p p e r gut w i l l be d e s c r i b e d f i r s t , t h e n l o s s f r o m the

l o w e r g ut, a n d

so on.

L o s s f r o m the u p per g u t.
(Table

s t r o n t i u m w ill be d e -

VI a n d F i g u r e s

injection th e re

is

It i s a p p a r e n t f r o m t h e d a t a

1 a n d 2) t h a t i n t he f i r s t f e w m i n u t e s a f t e r

an e a rly lo s s

of a p p r o x i m a t e l y

30 p e r

c e n t of th e

i n j e c t e d i s o t o p e w h i c h would o r d i n a r i l y not be a s s o c i a t e d w ith n o r m a l
ga stro in te stin a l m ovem ent.
m o v e m e n t in r a t s
the

chicks used

w h i c h h ad b e e n

s t a r v e d f o r 48 h o u r s .

in t h e s e e x p e r i m e n t s

until a few m in u te s before
It w a s

F e n t o n (1945) a l s o found a n e a r l y

rapid

However,

had feed and w a t e r a v a i l a b l e

injection.

o b s e r v e d t h a t th e f u l l n e s s

of the

crop varied greatly

b e t w e e n c h i c k s e v e n t h o u g h a l l had a c c e s s to f e e d a n d w a t e r with
am ple

space to p r e v e n t

conditions m ight have
made

at t h e t i m e

of g a s ,
VII)

crowding.

To d e t e r m i n e

on th e e a r l y S r

90

passage,

of i n j e c t i o n on t h e d e g r e e

a nd o c c u r r e n c e

of c r o p

contractions.

sh o w c l e a r l y t h a t t h e q u a n t i ty

what e f f e c t c r o p
observations were

of f u l l n e s s , p r e s e n c e
These data

of f e e d , the p r e s e n c e

(T able
of g a s

or

55
TABLE

VI

90
D IST RI BU TIO N O F Sr
RECOVERED, LACTOSE EXPERIM ENT
(all v a l u e s g i v e n a s m e a n p e r c e n t of i n j e c t e d d o s e ± s t a n d a r d e r r o r )
Per
No . of
Chicks

T i m e a nd T r e a t m e n t

Cent S r 9 °

U p pe r
G ut

P1

........................... . .
...........................

7
10

67 ± 2
6 4 db 3

NSD

15 m i n . c o n t r o l g r o u p ............................... . .
15 m i n . l a c t o s e g r o u p ............................... . .

12
9

2
51
60 ± 2

1%

30 m i n . c o n t r o l g r o u p ...............................
30 m i n . l a c t o s e g r o u p ...............................

11
11

2
51
d
b
2
51

NSD

45 m i n . c o n t r o l g r o u p ............................... . .
45 m i n . l a c t o s e g r o u p ...............................

9
11

35 ± 4
45 ± 2

1.4%

60 m i n . c o n t r o l g r o u p ...............................
60 m i n . l a c t o s e g r o u p ...............................

12
12

40
3
45 ± 3

NSD

90 m i n . c o n t r o l g r o u p ...............................
90 m i n . l a c t o s e g r o u p ...............................

10
11

26 ± 4
4
29

NSD

120 m i n . c o n t r o l g r o u p ...........................
120 m i n . l a c t o s e g r o u p ...........................

11
11

23 ± 4
21 ± 3

NSD

180 m in ,. c o n t r o l g r o u p ...........................
180 m i n . l a c t o s e g r o u p ...........................

10
8

9 rfc 5
9 d= 3

NSD

240 m i n , . c o n t r o l g r o u p ........................... . •
240 m i n , . l a c t o s e g r o u p ...........................

9
10

6 ± 4
10 rfc 3

NSD

24 h o u r c o n t r o l g r o u p ............................... . .
2 4 h o u r l a c t o s e g r o u p ............................... • •

8
9

7 .5 m i n .
7 .5 m i n .

1

control group
la c to se group

S tatistical probability.

C alculated value p e r
( c s p / f e m u r ) / (8.84 ) / s t a n d a r d .

cent Sr

in s k e l e t o n =

_

-

4

TABLE

YI ( C o n t i n u e d )

Per
Lower
Gut

P

24 ± 2
20 ± 2

33 ± 1
27 ± 2
54 ± 4
44 ± 3
4
3

62 ±
51 ±

5
3

55 ±
47 ±

4
5

2

NSD

0.1
1.9 ± 0.4

NSD

1.6 ± 0.2
1.1 ± 0.2

5.5%

0.3
3.5
2.4 ± 0.4

5.7%

9.9 ± 0.8
4.8 ± 0.4

7.2%

6.3
5.1 ±

0.8
0.6

5.3%

14.8 ±
12.8 ±

0.7
1 .4

NSD

18.0 ± 1 .4
2.8
26.0

NSD

15.5 ± 1 .3
12.3 ± 0.9

NSD

16.5 ± 1 .7
15.2 ± 1 .8

-

21.2 ± 4.2
5 .4
27.1

P

12 ifc 3
14 ± 1
7 ± 1
10 i 2
__

-

p e r ml .
Blood^

P

2.7%

0.043 ± 0.007
0.045 rb 0.005

NSD

1%

0.043
0.007
0.029 ± 0.005

NSD

9.3%

0.025 ±
0.022 ±

0.004
0.005

NSD

1%

0.042 ±
0.026

0.005
0.002

1%

NSD

0.020 ±
0.02 7

0.003
0.003

NSD

NSD

0.024 ± 0.004
0.025 ± 0.004

NSD

1.1%

0.041
0.005
0.057 ± 0.007

NSD

3.0%

0.015 ± 0.002
0.002
0.018

NSD

NSD

0.018 ± 0.001
0.013 ± 0.004

NSD

NSD

-

1.0

2
27
26 ± 2

50 ±
42 ±

Skeleton

C e n t Sr^

3

90
C a l c u l a t e d va lue p e r c e n t Sr
p e r m i l l i l i t e r of b lo od =
( c p s / m l . ) X 0.00315.
F a c t o r 0.00315 i s e x p l a i n e d in th e ap pe n d ix .
4

NSD = no s i g n i f i c a n t d i f f e r e n c e .

57
a:
LJ

£

o

_l

Ld

Ld

Ld

H f—

“ Z

— 8

Q

O

LJ

XN3WD39 dSddH Nl

%
a
u.

58

100

% S R 9 0 REMAINING IN
UPPER SEGMENT =
6 7 . | e — 0. 009 6 6 *
T 1/ 2 71.7 MIN.
rx* = “ 0 . 9 8 6

80 -

6 0 P-

z

Ld

2
O

40

* —

Ld
if )

QC
Ld
CL

CL

O

z
*
20
a:
if)

vp

0s

10 -

-

30

60

90

120

150

I

210

MINUTES
S I N C E D I F F E R E N C E S W E R E I N F R E Q U E N T L Y N O T E D B E T W E E N C ONT R OL
AND L A C T O S E T R E A T E D GR O U P S * T H I S F I G U R E IS C O N S T R U C T E D FROM
C O M B I N E D M E A N S . T H E S T A N D A R D E R R O R S OF E A C H P O I N T I NCREASE
FROM 2 . 3 %
A T 7 . 5 M I N U T E S T O 3 1 . 1 % AT 1 8 0 M I N U T E S . T H E COR­
RELATION
C O E F F I C I E N T W A S C A L C U L A T E D F R O M T H E MEAN P O I N T S
ON T H E F I G U R E AND N O T FROM I N D I V I D U A L B I R D S .

F I G U R E 2 . R A T E ON
TO LOWER SEGMENT

S R 9 0 P AS S AGE
OF B O W E L .

FROM

UPPER

59
TABLE
THE E F F E C T

V II

O F VISIBLE D I F F E R E N C E S IN THE C R O P
ON I NI T IA L E M P T Y I N G

C ro p Condition

90
P e t . Sr
Leaving
C r o p i n 7.5 m i n .

F a l l f e e d , no c o n t r a c t i o n .................................. ...................

35 ± 5

F a ll feed,

c o n t r a c t i o n ...............

32 ± 4

F all,

no c o n t r a c t i o n ...............................

36 ± 5

gas,

c o n t r a c t i o n ..................................................

33 ± 3

E m p t y , no c o n t r a c t i o n .......................................... ...................

40 ± 4

................................................................................................ ...................

35 ± 2

E m pty ,

X

60
c o n t r a c t i o n of t h e

c r o p at the t i m e

of i n j e c t i o n had no e f f e c t on t h i s

initial lo ss

isotope f r o m the

crop.

of t h e

A c o m p a r i s o n of t h e
( T a b le

VI) s h o w e d t h a t

f r o m the

co n tro l and la c to se -su p p le m en te d groups

Lactose p o s s i b l y

up p er bowel segm ent,

h a d no e f f e c t .

a f t e r injection, the

of u p p e r gut

ably u n if o r m
Therefore,

(Figure

c o n s t r u c t i o n of F i g u r e

noted before

segm ent emptying was r e m a r k ­

c o m b i n e the

2 and T a b l e

used

VIII.

w i t h the a c t i v i t y

s id e r e d a r a t h e r excitable

lowed

significant differences

a nd c o n t r o l g r o u p u p p e r gut r e c o v e r i e s w e r e

w a s t h e Single C o m b

injection, the

of the

observed afte r
chicks.

White L e g h o r n w h ic h i s
breed.

chicks frequently

battery.

struggled.

Th ey u s u a l l y

F o r t y - f i v e to

to

com pletely; they ate, drank,

activity.

The b r e e d
commonly

con­
during

This re a c tio n was f o l­
chicks w ere

r e f u s e d to e a t

this tim e.
recover

1 hour,

In the han dlin g n e c e s s a r y

by a q u i e t o r d e p r e s s e d b e h a v i o r a f t e r the

t u r n e d to t h e

w e r e note d.

d a ta , a s w a s done in the

60 m i n u t e s , and t h a t none w e r e

m i g h t be a s s o c i a t e d

out of n in e,

B e t w e e n 60 and 240 m i n u t e s

It w a s t h o u g h t t h a t t he f a c t t h a t a l l
lactose

of p a s s a g e

1) and no d i f f e r e n c e s due t o l a c t o s e

it w a s p o s s i b l e t o

betw een the

rate

but a t only two t i m e s

and th a t u s u a ll y l a c t o s e
rate

s low ed the

re­

or drink during

60 m i n u t e s a f t e r i n j e c t i o n t h e y a p p e a r e d

T h e f a i l u r e to find d i f f e r e n c e s

and a g a i n

showed n o r m a l

b e t w e e n the

c o n t r o l and

61
TABLE

VIII

CO M B IN E D C O N T R O L AND L A C T O S E DATA,
BOWEL RECOVERIES
90
P e t . Sr
in U p p er
G ut

Tim e
(m in.)

0

100

7.5

Ob s e r v e d
P e t . S r 90
in L o w e r
Gut

Calculated
P e t . S r 90
in L ow er
Gut

Loss from
L o w e r Gut
(pet.)

0

66

2

22 ± 2

34 ± 2

12 ± 2

15

56

2

27 ± 2

44 ± 2

17 ± 2

30

51 ± 1

30 ± 2

49 ± 1

18 ± 2

45

40 ± 2

49 ± 3

60

2

12 ± 3

60

42 ±

2

46 ± 2

58

2

11 ± 3

90

28 ±

2

56 ± 3

72 ± 3

16 ± 4

120

22 ±

2

51 ± 3

78

2

27 ± 4

180

9

3

13 ± 2

91 ± 3

78 ± 3

240

8 ± 2

8 ± 1

2

84 ± 3

1

92

90
90
One h u n d r e d p e r c e n t Sr
in u p p e r gut = c a l c u l a t e d Sr
in l o w e r , on t he a s s u m p t i o n t h a t no S r 9 ° h a d b e e n e i t h e r a b s o r b e d
or defecated.
T h i s i s the l i m i t i n g c a s e with w h ic h the o b s e r v e d
data m u s t be c o m p ared .

lactose

groups after

60 m i n u t e s m i g h t have b e e n r e l a t e d to t h i s

t u r n to a n o r m a l b e h a v i o r f ollo w in g i n j e c t i o n .
noticeable
very

p a t t e r n , h o w e v e r , t h e e m p t y i n g of t h e u p p e r

h ighly

of F i g u r e

correlated

2.

interfere

w ith t i m e ,

T h u s t he

with the

entirely

a s i n d i c a t e d by the

s t e a d y e m p t y i n g of t h e

crop.

between lactose

very-

s e g m e n t w as
s e m i l o g plot

o b s e r v e d b e h a v i o r p a t t e r n did not

co nnectio n w ith the d if f e r e n c e s
is

Despite this

re­

seriously

W h e t h e r it had any
and c o n t r o l g r o u p s

conjectural.

Th e d a t a i n d i c a t e , th en , t h a t i m m e d i a t e l y following i n j e c t i o n
in to t h e
dose)

crop,

a 11 s l u g ’’ of S r

90 ,
(about 30 p e r

p a s s e d i m m e d i a t e l y t o the

a process
presence

segment

injected

of the b ow el by

w h i c h w a s i n d e p e n d e n t of d i f f e r e n c e s in c r o p f u l l n e s s ,
of g a s ,

proxim ately

or noticeable

by t h e e q u a t i o n :

Following t h i s ,
t o the

absorptive

c e n t of the

contractions.
p e r cent S r

loss

is given ap-

u p p e r = 3 2 . 9e

—0 . Ot

n o r m a l u p p er segm ent m u s c u la r activity p a s s e s

absorptive

p o r t i o n at a u n i f o r m

e x a c t l y d e s c r i b e d by t h e e q u a t i on :
6 7 . 1 e " ° '° 0^ ^

90

T his

(t i n m i n u t e s ) .

per

exponential rate

rate

=
equation

w a s not a f f e c t e d by d i f f e r e n c e s in t he g e n e r a l b e h a v i o r of th e
The

com plete

m a t e l y by t h e

s i t u a t i o n in the u p p e r
combined

e q u a t io n :

90

which is

r. 90
cent S r
upper

On the a v e r a g e , t h i s

Sr

c h ic k s .

segm ent, then, is given a p p ro x i-

per

ce nt S r

90

n -0. 6t
u p p e r = 3 2 . 9e
+

l e "0-00966t^ in w h i c h t h e a p p r o x i m a t i o n i s only in the e x p o n e n t in

63
the f i r s t t e r m
u te s, t h e

(in t h e a b s e n c e

first process

a v e r a g e s 1’ t h e
actually
is

required

passage

of a l l e a r l y

There

25 p e r

f r o m the

e v e n t s , if m o r e t h a n one w e r e

chicks than f o r th e i r n o rm a l controls.

c en t l a c t o s e

in the

c r o p into t he a b s o r p t i v e

M o v e m e n t t h r o u g h l o w e r gut.
b ow e l, t h e p e r
determ ined
this

c e n t of S r

by t h r e e

90

factors;

r a t i o n did not a f f e c t S r

T u r n i n g now to the

r e c o v e r e d f r o m the
(1) the

am ount which has been ab so rb ed ,

90

s e g m e n t of th e bowel.

lower

l o w e r gut m u s t be

amount which has e n te red

s e g m e n t f r o m t h e u p p e r gut, a s d e s c r i b e d a bo v e ,

been defecated.

7.5 m i n ­

i s no i n d i c a t i o n t h a t a d i f f e r e n t e q u a t i o n

for la c to se -fe d

In o t h e r w o r d s ,

e a r l i e r th an

c a n only be d e s c r i b e d , by a n e q u a t i o n w h ic h

effects

in v o l v e d ) .

of d a t a a t t i m e s

(2) t h e net

and (3) the a m o u n t w h ic h h a s

No d e l a y i s i n v o l v e d in p r o c e s s e s

(1) a n d (2).

In

th e m a t h e m a t i c a l d e s c r i p t i o n of d e f e c a t i o n , h o w e v e r , a d e la y t e r m
m ust

be i n t r o d u c e d t o a l l o w f o r t r a n s i t t h r o u g h the l e n g t h of t h e

b ow e l to t h e

vent.

No s i g n i f i c a n t d i f f e r e n c e s

(due to l a c t o s e ) in S r

90

recovery

f r o m th e l o w e r gut a p p e a r e d u n t i l 30 m i n u t e s a f t e r i n j e c t i o n (see
Table

VI and F i g u r e

a p p e a r e d to be
of t h e

lactose

3).

From

s m a l l e r am ounts
groups,

30 m i n u t e s u n t i l 90 m i n u t e s , t h e r e
of Sr

90

p r e s e n t m the

but the d i f f e r e n c e s w e r e

l o w e r bow el

of b o r d e r l i n e

O
CM

BOWEL.

04

O
O

LOWER

O

CM
CM

O

IN

O

o

TVJ
O
O
O

00

o

CO

- :§

PER

CD

CENT

O

MINUTES
OF S R 90 PRESENT

00

THE

CM

co

if )

o

"M*

o

cO

J.N3WD3S W3MO"l NI^HS %

o

CM

FIGURE

_

3.

o

CM

65
significance.

They

u p ta k e , since

in e v e ry

percentage

of S r

th e d i f f e r e n c e s
differences

90

c a n n ot be a t t r i b u t e d to d i f f e r e n c e s
c a s e , the

i n blood a c t i v i t y w e r e

in t h e

skeletons

t h a n th o se f r o m the la c to se

i n t h e l o w e r bowel.

w i th l a c t o s e

control

in s k e l e t a l

contained a higher

groups.

L ikew ise,

i n a d e q u a t e to a c c o u n t f o r

T h e r e f o r e , the d i f f e r e n c e s a s s o c i a t e d

l o w e r b ow e l c a n only be a c c o u n t e d f o r by s m a l l

d iffe re n c e s in d efecation r a t e s

between lactose

It i s d o u b t f u l t h a t t h e p r e s e n t d a t a a r e

and c o n t r o l g r o u p s .

good e n o u g h to d e m o n s t r a t e

such effects.
In o r d e r to

obtain the best g e n e ral picture

from

the lo w e r bowel s e g m en t,

bined

as before

(Table

c o n t r o l and l a c t o s e

VIII) and the d a t a p l o t t e d

90
recovery

of S r

data w ere

( F i g u r e 4).

It w i l l be n o t e d t h a t it w a s n e v e r p o s s i b l e t o r e c o v e r
low er

s e g m e n t a c t i v i t y w h ic h sh ould have

quently,
initial

as

soon as S r

90

p o rtion m u st im m ediately

b o w e l and p a s s t o t h e p l a s m a ,

Conse-

This

l a r g e r fractio n , then,

form

of S r

l e a v e the

e x t r a c e l l u l a r fluid, and body t i s s u e s .

A s e c o n d an d l a r g e r f r a c t i o n left the

l o w e r gut m u c h m o r e

is a " b o u n d " ,

, a l t h o u g h the t y pe

not be d e t e r m i n e d

been p re se n t.

a l l the

a p p e a r e d in the l o w e r bowel (in the

s lu g) , a c o n s i d e r a b l e

90

com­

slowly.

o r at l e a s t a l e s s m o b i l e

of r e s t r i c t i o n on i t s m o b i l i t y

f r o m t h e s e d a t a a l on e .

c an

6 *>

80—

CALCULATED

60-

OBSERVED

£ •

20—

DIFFERENCE

l0 ~ ABSORPTION
OF INITIAL
g _ _ SLUG

/

COMBINED
ABSORPTION
AND
DEFECATION
180

240

MINUTES
F I G U R E 4 . S R 9 0 R E C OV E R I E S FROM THE LOWER
B O W E L . [ B A S E D O N C O N B I N E D M E A N S , FOR COMPARI SON
WITH FIGURE 2.]

67
In k i n e t i c t e r m s ,
pooLs (m t h e

then, th e r e

absorptive

cent

of t h e

defecated
m ate ly)

in jected dose

The

which p a sse d

extrem ely

rapidly,

slow t e r m

30 p e r

crop im m ediately

and w a s p r o b a b l y n e v e r
The

w h ic h l e a v e s the

70 p e r c e n t ( a p p r o x i ­
c r o p at the

rate

g iv en

in t h e e q u a t i o n m e n t i o n e d a bove m a y be a l m o s t

c o m p l e t e l y d e f e c a t e d w i th ou t a b s o r p t i o n a t a ll.
hand, th is

slowly

(approximately)

out of the

except as endogenous fecal Sr.

of t h e i n j e c t e d d o s e

by t h e

r a p i d l y m o v i n g and f r o m the

of t he i n j e c t e d d o s e .

s e e m e d to be a b s o r b e d

strontium

p o r t i o n of t he bowel) w h i c h a p p e a r to be d e ­

rived. r e s p e c t i v e l y f r o m t h e
moving f r a c tio n s

m u s t be a t l e a s t two

Or,

on the

other

s l o w m o v i n g S r m a y be

slowly c o n v e r t e d to a

' ' f r e e ' 1 Sr

which was its e lf rapidly ab so rb ed ,

but whose q u a n t i t a t i v e

im portance

of b r e a k d o w n of the

slow m o v in g

would d e p e n d p r i m a r i l y
Sr

"com plex."

entirely

on t h e

T h e s e p o i n t s w e r e not d i r e c t l y i n v e s t i g a t e d .

possible,

of c o u r s e , t h a t m o r e t h a n one f o r m

p r e s e n t i n the d i g e s t i v e t r a c t .
could not have
Sr atom s

rate

a simple

s y s te m in which all

sim ilarly.

T h e m a t h e m a t i c a l d e s c r i p t i o n of the S r
lower

s e g m e n t a s a f u n c t i o n of t i m e

lated fro m th e se
9 term

of Sr w as

In f a c t , the d a t a p r e s e n t e d h e r e

b e e n o b ta i n e d f r o m

behave

It is

90

r
r e c o v e r y f r o m the

a f te r injection has been c a lc u ­

d a t a by W o l t e r i n k ( p e r s o n a l c o m m u n i c a t i o n ) .

equation is n e c e s s a r y .

Two t e r m s

describe

A

input f r o m the

68
upper

segm ent.

about

60 m i n u t e s ) ,

required

D efecation r e q u i r e s
another.

to d e s c r ib e

1 term

This leaves

''a b s o rp tio n ”

a nd i t s d e l a y

5 term s,

of w h i c h 4 a r e

(gut t o blood) a n d t h e l a s t , to

d e s c r i b e t r a n s f e r b a c k into t h e b o w e l (blood to gut).
t h i s e q u a t i o n t h e fo llo win g
Of t h e
low er
cent

67.1 p e r

results

w ere

o b t a in e d.

of i n j e c t e d ) w a s a b s o r b e d

spurt,

82 p e r c e n t (thus,

was

of t h e i n j e c t e d d o s e

On t h e o t h e r ha nd, a l l of t h e i n i t i a l s p u r t of S r

absorptive
18 p e r

55.2 p e r

so slowly t h a t d e f e c a t i o n b e g a n

( a f t e r 5 7.3 m i n u t e s ) , w i t h 4 1.0 p e r ce nt

the

B a s e d on

c e n t of th e i n j e c t e d d o s e w h ic h e n t e r e d th e

seg m e n t a f t e r the initial

unabsorbed.

(until

segm ent was

c e n t of t h e

It w a s p o s s i b l e , t h e n , t o

into

90 .
fractio n.

su b d iv id e th e S r

90

i n j e c t e d into the

c r o p a s follows:
100% i n j e c t e d into
67.1% m o v e d ' ’n o r ­
m a l l y ” t o a b s o r p t i v e s e g.

crn n

absorptive

%)

seg.

all v ery rapidly
absorbed
1.9% r a p i d l y a b ­
s o r b e d (two r a t e s )

55.2% slo w ly
absorbed
(74%)
\k
4 1.0 % ( m a x i m u m )
d e f e c a t e d (without
absorption)

90

c a l c u l a t e d t o be r a p i d l y e x c h a n g e d , a s

slo w e r moving S r

(82%)

still

( 2 6 %)

' >>vvx^ 1 4 . 2 %

(minimum)

69
T w e n t y - f o u r h o u r s a f t e r t he
of t h e i n j e c t e d d o s e
cent was

rem oved

rem ained

crop injection,

in the

chick, w h e r e a s a t l e a s t

f r o m t h e b o w el by t h o s e p r o c e s s e s

d e f e c a t i o n w h i c h h av e b e e n l o o s e l y t e r m e d
q u e n t l y , i n one d a y, n e a r l y
lost th ro u g h the
fe c a l Sr

90

combined

60 p e r

c en t

59 p e r

other than

' ' absorption.' 1

c en t of 1' a b s o r b e d ' ' Sr

Conse90

wa s

r o u t e s of r e n a l e x c r e t i o n and e n d o g e n o u s

l o s s , i n a d d i t i o n to t h e d i r e c t l o s s

the fe c e s .

only 24 p e r

of u n a b s o r b e d

Sr

9q

in

T h i s m a y be s u m m a r i z e d a s follows:

Injected
A b so rb e d (m inim um)
R e t a i n e d , 24 h o u r s
L o s t , 24 h o u r s
U n a b s o r b e d S r9 0 i n f e c e s ( m a x i m u m )
L o s s via u r i n e + e n d o g e n o u s f e c a l Sr^O

100%
59%
24%
76%
41%
35%(59%

of 59%)

In t h e p r e s e n t e x p e r i m e n t s , m o s t of the l o s s e s d e s c r i b e d
above

occurred

in the f i r s t fo u r h o u rs .

+ s k e l e t o n + blood) w a s

32 p e r

24 p e r

a l l i n the

cent

tention,

(essentially

16 p e r

c en t w a s

cent, w h e r e a s

76 p e r

From

the

dose

lef t t h e

8 per

c e n t of t h e

retentio n

was

Of the f o u r h o u r r e ­

skeleton w here

h a l f (8 p e r

it w as

recovered

s t a n d p o i n t of l o s s , t h e n , the t o t a l l o s s wa s

c e n t of t h e i n j e c t e d d o s e .

jected

s k e le t o n ) .

24 h o u r s

s t i l l i n the bowel and of t h i s ,

c e n t of i n j e c t e d ) f i n a l ly went to the
a t 24 h o u r s .

T h u s , 4 h o u r r e t e n t i o n (gut

However,

66 p e r

c h i c k s b e t w e e n 0 an d 4 h o u r s .

i n j e c t e d d o s e le ft th e

cent of the i n ­
Only an a d d i t i o n a l

c h i c k s b e t w e e n 4 and 24 h o u r s .

70
This
t i o n of t h e

indicates

adm in istered, dose

pended p r i m a r i l y
was,

as

t h a t in t he m e t a b o l i s m

u p o n tw o

e x p e c te d , the

tinal absorption.

If t h e

wou ld be d e f e c a t e d .
t h r o u g h the

rate

which was

of s t r o n t i u m , the f r a c ­

r e t a i n e d in the

r a t h e r slow r a te s .
(or a s

skeleton d e ­

The f ir s t

shown h e r e , t h e

of t h e s e

rates)

strontium were absorbed m o re

of i n t e s ­

rapidly,

less

To put it a n o t h e r way, if t he t r a n s i t t i m e

b o w e l could be i n c r e a s e d

(by d e c r e a s i n g i n t e s t i n a l

Mm o t i l i t y ’1) w i th o u t a l t e r i n g the a b s o r p t i o n r a t e s , m o r e

strontium

w ould be a b s o r b e d .
How ever, th is is
slow ra te
f ro m the
m ost

was the

b lo o d by t h e

of s u c h s t r o n t i u m

ta tiv ely ,
uptake

of t he p r o b l e m .

(or ra te s) at which strontium was
skeleton.

state

chief r e a s o n that

it a n o t h e r way, the

s t r o n t i u m to the

feces

the

s k e le t o n .

growing

Q uanti­

s lo w s k e l e t a l

stro n tiu m was lost.

combined lo s s

of a b s o r b e d

of s t r o n t i u m by th e

t h e p r o p o r t i o n of 35:24 ( a p p r o x i m a t e l y

only i n c r e a s e

cleared

(as e n d o g e n o u s f e c a l s t r o n t i u m ) a nd to the

w a s f a s t e r t h a n the u pta ke

constant,

second

E v e n if a b s o r b e d into t h e blood,

w a s not r e t a i n e d by the

of s t r o n t i u m w a s t h e

m ained

The

j u d g i n g f r o m t h e p r e s e n t d a t a , th e r e l a t i v e l y

A g a in , to

urine

rate

only a p a r t

3:2).

s k e l e t o n in

If t h i s p r o p o r t i o n r e ­

i n c r e a s i n g t o t a l a b s o r p t i o n by 2 0 p e r

s k e l e t a l r e t e n t i o n of s t r o n t i u m by 8 p e r

ce nt would
cent.

s k e l e t o n w a s a l r e a d y e x p o s e d to m u c h m o r e

Since

strontium

71
t h a n it w a s t a k i n g up,

it m a y

be d ou b te d w h e t h e r s u c h a n i n c r e a s e

i n i n t e s t i n a l a b s o r p t i o n would give e v e n an 8 p e r
skeletal

in

strontium .

One
olism

cent i n c r e a s e

i s l e f t, t h e n , w ith t h e d i s t i n c t i m p r e s s i o n t h a t the m e t a b ­

of t h e

sk eleto n is

considerably m o re

i m p o r t a n t t h a n the p h y s i ­

ology of t h e

b o w e l , i n s o f a r a s the

concerned.

T h e f a c t t h a t d a t a of t h e l a c t o s e g r o u p s a nd c o n t r o l

groups a re
To
Table

very

s k e l e t a l r e t e n t i o n of S r 9 ° - Y 9 ° i s

s i m i l a r i s i n h a r m o n y w i th t h i s

s u m m a r i z e , the

g en eral picture

VI, a s f a r a s t h e b e h a v i o r of Sr

concerned,
words,

is that all

d e r i v e d f r o m t h e d a t a of
i n the l o w e r b o w el w a s

s t r o n t i u m d o e s not b ehave t h e

d i f f e r e n t gut p o o l s w e r e

gut f r a c t i o n t u r n s

90

c o n c ep t.

l a b e l e d by the t r a c e r .

o v e r at d i f f e r e n t r a t e s ,
90

Since e a c h

and, i n a d d it io n , v a r i e s

skeleton,

o r e x c r e t a at d i f f e r e n t t i m e s a f t e r i n j e c t i o n c hange in s u c h

way t h e s e
sheet

were

the type

recovered from

in

observed

require

of S r

In o t h e r

size, the

a way a s to

quantities

same.

of s y s t e m

gut, blood.,

outlined a b o v e .

o b s e r v e d to cha nge i s g iv e n in th e

The p r e c i s e

o v e r - a l l balance

of T a b l e IX.
With r e s p e c t t o t h e e f f e c t s

The data p r e s e n te d e a r l i e r
lactose:

of l a c t o s e ,

v e r y li t t l e

c an be

sh ow the u s u a l l y d e s c r i b e d e f f e c t s

re d u c ed grow th r a te ,

increased

said.

of

w ater consumption, in ­

c r e a s e d w e i g h t of i n t e s t i n a l t r a c t a nd c o n t e n t s , and p e r h a p s

some

72
T A B L E IX
M EA N P E R C E N T R E C O V E R I E S OF S r 9 ° - Y 9 °
A F T E R CROP INJECTION
( c o m b i n e d d a t a f r o m c o n t r o l a nd l a c t o s e - f e d g r o u p s )

Tim e
( m i n .)

Uppe r
to
Lower

R ecove red
from
Lower

Differenc e

Recovered
from
E xcreta

Recov­
ered
from
Skel­
eton

T ota l
Recov­
ery

7.5 . .

.

34

22

12

0

1

89

15 . . .

.

44

27

17

0

2

85

30 . . .

.

49

31

18

0

3

85

. .

60

48

12

0

7

95

60 . .

57

46

11

0

6

95

90 . .

72

56

16

22

14

rg
o
o
i—
i

. .

78

51

27

5

22

100

180 . .

91

13

78

64 2

14

100

240

92

8

84

69

15

100

100

0

100

762

24

100

45

120

. . . .

(24 h o u r s )

^ Total

r e c o v e r y = u p p e r gut + l o w e r gut + e x c r e t a +

skeleton.
Z

Calculated.

2

2

73
dehydration.
differences
w hether
large,

F urther,

the t r a c e r

i n c e r t a i n gut r e c o v e r i e s .

s ta tis tic a lly the n u m b e r

This

It m a y be q u e s t i o n e d , t h e n ,

of c h i c k s p e r g r o u p w a s

s u f f ic i e n tl y

statistical

"negative"

gr oups.

s i t u a t i o n is by no m e a n s u n c o m m o n .

results arise,

creased num bers,
conclu sions.
sented a r e

It m u s t be r e m e m b e r e d ,

consistent effects
th e

T he d a t a a r e

of l a c t o s e

only va lid

of l a c t o s e ,

s to ck , i n ­
d e fi n it e

h o w e v e r , t h a t the d a t a p r e ­

s i g n i f i c a n t to a l l o w d r a w i n g

p h y sio lo g ical conclu sio ns.

Therefore,

c h o ic e of g e n e t i c

u n i f o r m body w e i g h t - - m a y a l l o w m o r e

sufficiently

some

not c o m p l e t e

in in te s tin a l Sr

90

-Y

90

very basic

e no ug h to show

recoveries.

c o n c l u s i o n h a s to be t h a t t h e s e

latter e f­

if in de ed t h e y e x i s t , m u s t be q u a n t i t a t i v e l y

w i t h r e s p e c t t o i n d i v i d u a l d i f f e r e n c e s in i n t e s t i n a l e m p t y i n g .
c o r o l l a r y to th is

is that although la ctose

t h r o u g h t h e b o w e l , i t s e f f e c t on S r

90

-Y

90

T he p e r

c e n t Sr

90

in the l a s t c o l u m n of T a b l e

-Y
VI.

90

sm all
The

increased w ater tran sit
t r a n s i t was m uch less.

V a r i a t i o n . i n e x t r a c e l l u l a r flu id a n d soft t i s s u e Sr
injection.

W hen­

it m a y be p o s s i b l e t h a t a d d i t i o n a l

technical r e fin e m e n ts - - such as b etter

fects

significant

c o n s i d e r i n g i n d i v i d u a l v a r i a b i l i t y , to p i c k up s m a l l but s i g n i f i ­

c a n t d i f f e r e n c e s b e t w e e n the

ever

d a t a do show s t a t i s t i c a l l y

90

with t i m e a f t e r

p e r m i l l i l i t e r of whole b lood i s g i v e n
If th e h e m a t o c r i t in c h i c k s of t h i s

74
age

is ta k e n a s

m illiliters

0.29 (0.71 p l a s m a ) and the p l a s m a v o lu m e a s

(Sturkie,

1954), t h e t o t a l S r

g i v e n by ( p e r c e n t p e r m i l l i l i t e r )
unilliliter).

f lu id v o l u m e

(per

of 25 m i l l i l i t e r s

(Manery,

1950).

The to ta l Sr

be e s t i m a t e d

90

-Y

m a y be

reported interstitial
10 m i l l i l i t e r s

X 25 = 29.92

(per cent p e r m i l l i -

90 ,
i n th e e n t i r e e x t r a c e l l u l a r fluid could t h e n

No S r

v e r y low.

cent p e r

—
Y

(35 p e r c ent of 100 m i n u s

X 0.85/0.71

90

In g e n e r a l , t h e p e r

the

(0.85) and the

by m u l t i p l y i n g t h e p e r

w h o l e b lood by 44,

blo od w a s

= 14.08 ( p e r

will be

T h e t o t a l i n t e r s t i t i a l fluid S r 9 ° - Y 9 ° i s then:

cent p e r m illilite r)

liter).

X 1 0 /0 .7 1

in the p l a s m a

S i m i l a r l y , t h e t o t a l i n t e r s t i t i a l fluid S r

c a lc u la te d f r o m the Donnan Ratio

plasm a)

- Y^

10

-Y

90

c en t Sr

90

-Y

90

cent r e c o v e r y

of S r

90

-Y

(about 0.04 m i c r o g r a m s

c a lc u late d to p a s s th ro u g h the

of

e n t e r s the e ry th ro c y tes.

A l t h o u g h u l t i m a t e l y , at l e a s t

injected t r a c e r atom s

per m illiliter

circulatory

90

f r o m whole

59 p e r

c ent of

of s t r o n t i u m ) w a s

s y s t e m , the e n t i r e

extra­

c e l l u l a r fluid p r o b a b l y n e v e r c o n t a i n e d m o r e t h a n 1.74 m i l l i - m i c r o g r a m s
a t any one t i m e
understo od

(about 2.5 p e r

only if t h e

Sr

90

ce nt of the i n j e c t e d d o se ) .

were

tim e
the

c a n be

(in g e n e r a l , and w i th c e r t a i n e x c e p ­

t i o n s ) f ol l o w i n g th e v e r y m u c h l a r g e r
apparently

This

c a l c i u m p o o ls .

The

soft t i s s u e s

c o n t a i n e d a s u b s t a n t i a l a m o u n t of t r a c e r d u r i n g the e a r l y

periods

s i n c e the t o t a l r e c o v e r y f r o m the d i g e s t i v e t r a c t and

s k e l e t o n did not a c c o u n t f o r th e a m o u n t of Sr

90

injected.

For

75
e x a m p l e , at

7.5 m i n u t e s t h e

jected

( T a b l e IX), t h e

dose

c e l l u l a r f l u id a c a l c u l a t e d

b o w el c on ta in e d
s k e l e t o n only

Z per

c ent.

88 p e r

1 per

c ent of the i n ­

cent, and the e x t r a ­

Since e x c r e t i o n of S r ^

w a s h i gh ly i m p r o b a b l e t h i s e a r l y , th e u n d e t e r m i n e d a c t i v i t y
probably p r e s e n t
From
in the f i r s t
low er

in s o m e u n s a m p l e s

W o lterin k 's

soft t i s s u e s

be c a l c u l a t e d t h a t a c t u a l l y ,

7.5 m i n u t e s t w o - t h i r d s

of t h e

Sr

(th u s,

Z3 p e r

of t h i s ,

in t u r n ,

initial

'' s p u r t ' 1 of S r o f f e r e d to the bowel f o r

b o th i n t h e

m echanism .

r a p i d m o v e m e n t of the

custom arily

tion--that

of l a r g e

num bers

i s , n et l o s s

th e

here

ionic

s m a l l n u m b e r of r a d i o ­
is

a s an " e x c h a n g e "

re­

of the e x c h a n g e p r o c e s s e s being lef t open f or
What i s

strated

''a b s o rp tio n '1 m ust

i n t e s t i n a l m u c o s a in both d i r e c t i o n s

f u r t h e r investigation.

c o u ld not p o s s i b l y

T h u s the

(at l e a s t a s f i r s t a b s o r b e d ) .

s o m e w h a t l o o s e l y in t h e l i t e r a t u r e

a c t i o n , th e n a t u r e

had b e e n p a s s e d b a c k

on t h a t a c c o u n t w a s p r o b a b l y in s o m e

a t o m s a c r o s s the

designated

''exchange''

b o w el a nd i n the blood

extrem ely

w h ic h s p u r t e d to t he

c e n t of t h e i n j e c t e d d o s e ) w a s a c t u a l l y

l o w e r b o w e l by

be h ighly m o b i l e and

some

90

to the

active

of the body.

it m a y

segm ent

This

w as

equation,

a b s o r b e d a nd t h a t f o u r - t e n t h s

form ,

-Y ^^

c ontinue at t h i s

rapid

called in testin al a b s o r p ­

of a t o m s f r o m the bowel,
rate.

What w a s d e m o n -

w a s p r o b a b l y t h e i n i t i a l e q u i l i b r a t i o n of S r

90

between

gut p oo l ( o r p o o ls ) a nd t h e pool (or pools) in t h e e x t r a c e l l u l a r

76
body w a t e r
process
the
as

com partm ents.

intestinal

w h i c h a p p a r e n t l y d e p e n d s not

sim ple

ability

on t h e e x i s t e n c e

of m e c h a n i s m s

which a r e

"trap "

fo r the alkaline

first

in testin al contents
of e n d o g e n o u s

then,

so m u c h on e x c h a n g e

is a

rates

or

w i t h i n t h e body f or t r a p p i n g m a ­

"exchanged"
e a rth s is,

into the p l a s m a .

of c o u r s e , the

strontium

as a "sin k "

which d iv e rts

c hie f
The

of s e c r e t a i o n

b a c k into t h e bowel, a nd t h e
reabsorb

The

skeleton.

w h i c h bind c a l c i u m , the p r o c e s s

r e n a l t u b u l e s to c o m p l e t e l y

regarded

"absorption/'

of a m a t e r i a l to t r a v e r s e t h e i n t e s t i n a l m u c o s a ,

terials

the

True

i n a b i l i t y of

s t r o n t i u m m a y e a c h be

strontium from

its

s ite

of

d e p o s i t i o n i n bone.
A fter
blood S r
at

90

7.5 m i n u t e s , no

occurred.

30 m i n u t e s ,

initial

90

90

became

w a s what one would e x p e c t if

rapidly absorbed
available

and t h e n t h e m o r e

for ab so rp tio n somewhat la te r.

of the p i c t u r e d r a w n f r o m th e d a t a f o r

s e g m e n t of the b ow el a p p e a r e d a l s o i n the blood

a l t h o u g h to a
Since th e e n t i r e

contain s

in

w a s, h o w e v e r , a s i g n i f i c a n t d e c r e a s e

sequence
was

e s s e n t i a l points

the a b so rp tiv e
picture,

This

s p u r t of S r

slowLy m o v i n g S r
T hus the

significant in c re a s e

f o l l o w e d by a s ec on d i n c r e a s e to the e a r l i e r l e v e l

and a g r a d u a l f a l l .
the

There

statistically

sm a lle r degree.
e x t r a c e l l u l a r fluid (including th e p l a s m a )

only a f r a c t i o n of t h a t a m o u n t

of c a l c i u m found i n the

lower

77
bowel c o n te n ts , it is
m e n t is
lated

easily

to

the

a p p a r e n t t h a t the

saturated

bowel

pool

and

behaves

and

is

e x t r a c e l l u l a r f lu i d c o m p a r t ­
as

if

it

exchangeable

w ere
with

m echanism s

which re g u late

b e t w e e n the

gut a n d p l a s m a p o o l s a r e by no m e a n s

W h e t h e r the
m ucosa

"exchange”

w ater

are

processes

ratio n had very little

h a v e no d i r e c t

e f f e c t of i t s

ents

of t h e

90

from

im m ediate
VI,
in

IX).
the

the

upper

amount
are

of

90

is h ig hly q u e s t i o n a b l e .
e f f e c t, a lt h o u g h the n e t

P r a c t i c a l l y , th e n, i t i s c o n ­
excreta from

the c h i c k n e e d

own on the a b s o r p t i o n of o t h e r c o n s t i t u ­

in th e s k e l e t o n .

gut

depositio n
There

differences

the i n t e s t i n a l

ration.

D e p o s i t i o n of Sr
Sr

of a w a t e r y

earths

( eve n though w a t e r - s o l u b l e )

l a r g e l y i n d e p e n d e n t of e a c h o t h e r .
existence

The

A ga in, i t i s i n d i c a t e d t ha t

of w a t e r a n d s t r o n t i u m

c l u d e d t h a t the

re­

understood.

going on a c r o s s

m o v e m e n t w a s g r e a t l y m o d i f ie d .

t he m o v e m e n t s

i t.

the p a r t i t i o n i n g of the a l k a l i n e

c a n b e a l t e r e d by p h y s i o l o g i c m e a n s

C e rta in ly , the la c to s e

closely

of

are

segment
Sr

in

the

its

Sr

90

consistently

s i n c e th e a m o u n t of S r

90

rapid

skeleton

d ifferences between

skeletal
not

90

and

A f t e r the i n i t i a l s p u r t of

the

deposited.
in

the

absorption,

took
two

place

feeding

However,
same

d e p o s i t e d is f a i r l y

an

(Tables
regim es

these

direction,

and

sm all, p e rc e n ta g e -w ise ,

78
one

c a n n o t d r a w any d e f i n i t e

ference

b etw een the

conclusions

lactose

and c o n tr o l gro ups.

C o m b i n i n g the two g r o u p s
done w i t h o t h e r d a t a ) , it w a s
very

the

of bone d a t a

occurs

control group

th a n the
the

tw o

45 m i n u t e
60 m i n u t e

( T able IX).

w h ic h w a s p r i m a r i l y

(Table

VI).

lactose

At 45 m i n u t e s ,

a tw o­

due t o the high va lue

Since t h i s g r o u p w a s

g r o u p and w a s a l s o

groups,

(as w a s p r e v i o u s l y

o b s e r v e d t h a t bone d e p o s i t i o n p r o c e e d e d

slo w ly a f t e r t h e i n i t i a l u p ta ke

fold i n c r e a s e

concerning a significant d if­

of

significantly h ig h er

significantly h i g h e r th an

it c a n be a s s u m e d t h a t t h i s p a r t i c u l a r g r o u p

of c h i c k s a c t e d d i f f e r e n t l y t h a n a l l o t h e r g r o u p s up to t h i s t i m e .
T h erefo re, this
true

picture

c o n t r o l v a l u e a t 45 m i n u t e s p r o b a b l y d i s t o r t e d the

somewhat.

B e t w e e n 60 a nd 90 m i n u t e s a l a r g e
Sr

90

deposited

in the

skeleton o c c u rre d .

increase

i n the a m o u n t of

Later, there are

s o m e d i f f e r e n c e s b e t w e e n th e two f e e d i n g g r o u p s w h ic h a r e
the

same

d i r e c t i o n a nd f r o m w h i c h no c o n c l u s i o n s

R e f e r r i n g n o w to t h e
th at the l a r g e s t i n c r e m e n t s
m inutes

of t h e

Sr

90

c o m b i n e d d a t a ( T a ble IX), it a p p e a r s
of Sr

90

t a k e p la c e b e t w e e n 60 and 90
A m ore

r e m a i n i n g i n the l o w e r gut to the

o c c u r r e d during th is l a t t e r period.
excreta

not in

can be d r a w n .

an d t h e n a g a i n b e t w e e n 4 h o u r s a nd 24 h o u r s .

ficient t r a n s f e r

again

A l s o , the

loss

of Sr

ef-

skeleton

9° .
m the

b e t w e e n 4 a n d 24 h o u r s w a s m u c h s l o w e r t h a n t h a t w hic h

79
took p lace

earlier.

a b ly l o c a t e d

Since a l a r g e p e r c e n t a g e

in t h e

p o s te r io r portions

of the S r ^

of th e a l i m e n t a r y t r a c t at t h i s

tim e,

it wo uld be i n t e r e s t i n g to i n v e s t i g a t e the

these

processes.
These

the

results

process.

The

with e i t h e r i n c r e a s i n g

c e c a in

The d a t a

(T able

X) a l s o

Thus the

skeleton may

o r d e c r e a s i n g blood l e v e l s .

s i m p l e s t way to d e s c r i b e t h e s e f l u c t u a t i o n s i s in t e r m s

of " c l e a r a n c e s , "

by t h e

skeleton,

a g i v e n i n t e r v a l of t i m e .
sim ply the am ount

of t h e blood a v a i l a b l e t o i t d u ri n g

A s u s e d h e r e , the

of blood

(in m i l l i l i t e r s )

up a l l i t s t r a c e r to th e t o t a l

consideration.

blood p e r m i n u t e .

skeletal clearance

w h ich would have to give

s k e l e t o n in the t i m e p e r i o d

T h i s f i g u r e w ill be in m i l l i l i t e r s
It i s

is

s k e l e t o n to a c c o u n t f o r th e a m o u n t of

t r a c e r a c c u m u l a t e d , p e r m i n u t e , by the
under

of the

of d e p o s i t i o n w a s not d e p e n d e n t only upon the

r „ 90
90
c o n c e n t r a t i o n of S r
-Y
in t h e blood.
sh o w i n c r e m e n t s

role

i n d i c a t e d t h a t th e d e p o s i t i o n of a m i n e r a l into

s k e l e t o n w a s not a u n i f o r m

show th a t the p a t t e r n

was p r o b ­

c u s t o m a r y to

co rrect this

of whole

clearance for

body w e i g h t d i f f e r e n c e s by c o r r e c t i n g i t to m i l l i l i t e r s p e r m i n u t e
per

square

M

=

m eter

of s u r f a c e

w h i c h g i v e s th e

body w e i g h t in k i l o g r a m s .
c o l u m n of T a b l e

X.

a r e a , u si ng t h e

square

surface a re a formula,

m e t e r s a s the

2 / 3 p o w e r of the

Such c l e a r a n c e s a r e t a b u l a t e d in the

last

80
TABLE X
SK ELE TA L CLEARANCE OF WHOLE BLOOD S r9 ° - Y 9 °

M ean B lo o d Sr
(p e t./m l.)
7. 5 m in u te s

90

S k eleton Sr
(pet.)

90

m l. B l o o d /m in .
/S k e le t o n /M 2

0.0457 ± 0.004

1.5 ± 0.2

42.7 ± 6.9

15 m in u t e s

.

0.0365 ±

0.004

1.3 ± 0.1

- 2.8 ± 5.6

30 m in u t e s

.

0.0239 dh 0.003

2.9 ± 0.3

14.9 ± 3.5

45 m in u t e s

.

0.0340 ± 0.003

7.4 ± 0.5

42.3 ± 7.5

60 m in u t e s

.

0.0236 ± 0,003

5.7 ± 1.0

90 m in u t e s

.

0.0249 ± 0.003

13.8 ± 0.8

48.2 ± 7.2

120 m in u t e s

.

0.0491

±

0.004

22.0 db 1.5

35.0 ± 8.7

180 m in u t e s

.

0.0 164 ±

0.003

14.0 ± 0.8

-1 9 -9 ± 5.1

240 m in u t e s

.

0.01 5 4 ± 0.002

15.8 ± 1.2

9.2 ± 7.0

0.0

24.1 ± 3.4

3.8 ± 1.3

24 h o u r s

. .

- 1 6 .7

2

7.1

B lo o d c l e a r a n c e defined as the num ber of m i l l i l i t e r s of w hole
b lo od w h ich c on tain the am ount of Sr^° taken up by the sk e le to n in
one m in u te .
2

90
___________________
.
Skeleton Sr % _______________
m e a n b lood Sr^^% X tirhe (m in.) / body s u r fa c e a r e a (M^)
^

The stan d a rd e r r o r w a s c a lc u la te d by com bining the standard e r r o r
of g r o u p s r a th e r than d e te r m in a tio n of individual d if f e r e n c e s .

From
90

T able

X, it i s e v i d e n t t h a t a bout t h e m o s t r a p i d u pta ke

90

of S r

-Y

by bone t o o k p l a c e

T h e n b e t w e e n 7.5 a n d
if anything,
ance

15 m i n u t e s , the a v e r a g e blood l e v e l fe ll and,

so d id t h e

skeletal strontium

was actually negative,

the beh av io r
it d u r i n g

of t h e

b e t w e e n 0 t i m e and 7.5 m i n u t e s .

a l t h o u g h not

s k e l e t o n toward, t h e

successive

recovery.
significantly

T h u s the
so.

Sim ilarly,

cha nging blood l e v e l s

p e r i o d s w a s g i v e n by th e

successive

clear­

offered

clearance

value s .
Obviously, the
accum ulates m o re
reverse

was tr u e .

succeeded
rapid

Sr

90

-Y

90

Further,

by a c o n s i d e r a b l y

uptake

actu ally

r e s u l t s indicated that

c a n not be

increase.

T his

s o m e t i m e s the

t h a n it r e l e a s e s ,
a very

rapid

and

so m e tim e s the

clearance

was usually

s l o w e r one, a s if the m e c h a n i s m f o r

sustained,

e v e n t h o u g h the blood l e v e l m a y

suggested

(though it did not p r o v e ) t h a t t h e r e

m i g h t be m o r e t h a n one m e c h a n i s m w h e r e b y the
rem ove

skeleton

s k e l e t o n i s a ble to

s t r o n t i u m f r o m t h e blood.
In i n t e r p r e t i n g t h e s e

standards for

com parison.

clearance
Two

standards are

b e i n g t h e o r e t i c a l in n a t u r e and th e
f r o m th e

values,

one o bviously n e e d s
a v a i l a b l e , the f i r s t

second based

on v e r y

Limited d a t a

literature.

1.
of a n o r m a l

It m a y be
100 g r a m

c o m p u t e d f r o m the e x p e r i m e n t a l g r o w t h r a t e

c h i c k t h a t a bo u t 0.14 m i l l i g r a m

of c a l c i u m is

82
d ep o site d in the
the

skeleton,

sk ele to n p e r m inute.

on t h e

average,

b lood e v e r y m i n u t e .
wou ld be

slightly

square m e t e r
average
about

The

should c l e a r about 2 m i l l i l i t e r s

clearance p e r

over 9 m illiliters

of body s u r f a c e a r e a

clearance,

s q u a r e m e t e r of body

surface

by th e e n t i r e

skeleton.

Th e

w e i g h t e d f o r t i m e , f r o m t h e d a t a of T a b l e X, w a s

11 m i l l i l i t e r s

of whole blood p e r m i n u t e p e r
(the f i g u r e

square m e te r

w as a c t u a l l y

11.3 ±

If t h e m e a n blo od l e v e l b e t w e e n 4 a nd 24 h o u r s w a s half the

4 h o u r fig u re , th e n the av e rag e
tim e,

clearance,

calculated over a d a y 's

is about 7 m i l l i l i t e r s p e r minute p e r

7.1 ± 0.8).

It w a s e v i d e n t , t h e n , t h a t t h e

clearances,

when a v e r a g e d

g row th r e m a r k a b ly

w h ole blood m u s t be
norm al

s i z e h a v e only a bo ut

s t i t i a l flu id .

clear.
To

is

100 g r a m

strontiu m

re flecte d the

skeletal

c hick, about 2 m i l l i l i t e r s

rem arkable

constant,

in i t s e l f ,

sin c e

chicks

of t h i s

of whole blood (10 m i l l i l i t e r s

In a d d i t i o n , t h e r e
c l e a r a l l the

of

of c a l c i u m e v e r y m i n u t e to s u p p o r t

14 m i l l i l i t e r s

e x t r a c e l l u l a r f lu id can be
rate

observed

o v e r s u f f ic i e n t t i m e ,

cleared

s k e le ta l grow th is

p la sm a ) to

s q u a r e m e t e r ( ac tu al l y

well.

T h e f a c t t h a t in a

the

of whole

of whole blood p e r m i n u t e p e r

calcu lated th ro u g h the f i r s t 4 h o u rs
1.1).

To do t h i s would r e q u i r e t h a t

i s about 25 m i l l i l i t e r s

of

of i n t e r ­

o r i g i n a l c a l c i u m f r o m the e n t i r e

c o m p u t e d to r e q u i r e

o r a b o u t 90 m i n u t e s

only

10 m i n u t e s if

if a l l o w a n c e i s m a d e f o r

83
exponential
atom

w ash-out.

Thus,

at

least

s i x t e e n t i m e s d a il y, e v e r y

of e x t r a c e l l u l a r fluid, c a l c i u m w a s w a s h e d out into the

At le a s t

six t i m e s

every

hour,

a n a m o u n t of c a l c i u m e q u a l to t h a t

i n t h e t o t a l e x t r a c e l l u l a r f lu id i s laid down i n t h e
this,

it i s

readily

seen that

S in c e t h e d a t a

pass from

e nough, f a i l u r e

d e v e l o p p r o p e r l y m u s t u s u a l l y be due to

c o u rse the
in a f o r m

The

case w here

From

c om in g to it f r o m the

sh o w t h a t t h e a l k a l i n e e a r t h s

gut to b lo od r e a d i l y

no t i n t h e b ow el.

sk e le t o n .

s k e l e t a l g r o w t h i s not so m u c h d ep e n d e nt

u po n b lo o d c a l c i u m a s it i s upon c a l c i u m
bowel.

s k e le t o n .

c a n and do

of the

skeleton to

s o m e t h i n g w r o n g in t h e bone,

c h ie f e x c e p t i o n to t h i s g e n e r a l i z a t i o n i s

of

c a l c i u m i s a c t u a l l y bound by bowel c o n t e n t s

w h i c h i s not a v a i l a b l e f o r a b s o r p t i o n .

P h y t a t e binding i s

a typical exam ple.
2.

The

clearance

values is

an a b stra c t

second

s t a n d a r d f o r c o m p a r i s o n of t h e s e

g i v e n by t h e

sin gle

reference

in the l i t e r a t u r e ,

of a p a p e r p r e s e n t e d b e f o r e t he San F r a n c i s c o m e e t i n g

of t h e A m e r i c a n P h y s i o l o g i c a l S oc ie ty i n A p r i l ,
Honour,

observed

a nd Copp (1955).

These

investigators

c l e a r a n c e s i n s e v e n young an d s ix a du lt
a f t e r injection.

The

w hole

i n young

skeleton:

p e r m inute p e r

valu es

s t u d i e d Calcium.’-45

r a t s i n th e f i r s t

obtained were,

rats,

1955, by F r e d e r i c k s o n ,

f o r the

127 ± 10 m i l l i l i t e r s

s q u a r e m e t e r , a nd in a d u lt r a t s ,

5 m inutes

clearance

by t h e

of whole blood

71 ± 7 m i l l i l i t e r s

84
of whole blood
corresponding
chicks,

from

p e r m in ute
value

this p re se n t

b lood p e r m i n u t e
The
after

(first

per

p e r square

s tu d y , i s 4 2.7 ± 6.9 m i l l i l i t e r s

rapidly as

calcium .

b e t w e e n t h e two

studied a fte r

T he
the h ig h e s t

50 p e r

cent

points, the

cle ara n c e

values

i n good g e n e r a l a g r e e m e n t a s f a r a s

concerned.

im plications
clearance

7.5 m i n u t e s i s

rats

i s p r o b a b l y not c l e a r e d a s

Considering these

is

of whole

o r a l a d m i n i s t r a t i o n , the

Also,

and. s t r o n t i u m

species are

o r d e r of m a g n i t u d e

The

m eter.

intravenous adm inistration.

longer than 5 m inutes,

surface.

7.5 m i n u t e s ) f o r S r 9 ° - y 90 in 100 g r a m

square

chicks w ere

m e t e r of body

of t h e s e f in d i n g s a r e

values

(those

a s follows:

o b ta in e d v e r y e a r l y

F irst,

in the e x ­

p e r i m e n t s ) m u s t r e p r e s e n t a m i n i m a l flow of blood ( o r e x t r a c e l l u l a r
fluid) t h r o u g h t h e

skeleton.

d o s e into t h e w hole
average

blood

T h u s , to get

s k e l e t o n of the

1.5 p e r

c h i c k i n 7.5 m i n u t e s ,

c o n c e n t r a t i o n of a p p r o x i m a t e l y

m i l l i l i t e r of whole

b lo od ( l / 2

cent of the i n j e c t e d

of 0.0457)

0.025 p e r

of blood had t o flow t h r o u g h t h e

A sm aller

blood flow would not get e n o u g h blood S r

m ore
per

ce nt p e r

r e q u i r e d t h a t at l e a s t 40

m illiliters

s k e l e t o n to a c c o u n t f o r th e

with a n

s k e l e t a l u p ta ke .

skeleton e v e ry minute.
90

Probably ,

-Y

90

to the

considerably

blood a c t u a l l y t r a v e r s e d the b o n e s but at l e a s t 40 m i l l i l i t e r s

m inute

had to get t h e r e .

In o t h e r w o r d s , th e fluid flow t h r o u g h

85
the

skeleton was actually

very great.

T h e r e i s no e x p e r i m e n t a l

r e a s o n t o b e l i e v e t h a t t h e blood flow w a s any l e s s a t l a t e r t i m e s .
C onsequently, the high in itial c le a ra n c e
sociated with the p r o c e s s e s
flo w in g t h r o u g h t h e

f o r e , t h e a m o u n t of S r

of t h e

of s k e l e t a l g ro w th .

have b e e n a s ­

T he a m o u n t of blood

s k e l e t o n d u r i n g t h i s e a r l y p e r i o d would

enough c a lc iu m to g r e a tly

t a k e n into t h e

could s c a r c e l y

90

exceed that re q u ired

f o r g r ow th .

supply
The r e -

c le a re d during this period was apparently

s k e l e t o n by e x c h a n g e b e t w e e n bone

blood r a t h e r t h a n by n e w g ro w th .

c a l c i u m and Sr

In o t h e r w o r d s ,

clearances

so m u c h g r e a t e r t h a n 9 m i l l i l i t e r s m a y be u s e d a s i n d i c a t o r s
large

quantities

indicators

of f lu i d flowing t h r o u g h t h e

required

processes

fact th a t the a v e r a g e

a r e t r a n s i e n t p h e n o m e n a (and t h u s
as transients).

fle cted the net

skeletal

a

clearance

f o r bone g r o w t h i n d i c a t e d t h a t th e

m athem atically

ured

of the

but not a s

of s k e l e t a l g r o w t h p r o c e s s e s .

Second, th e
to t h a t

skeleton,

90

rapid

c lo se

"exchange"

should be h a n d le d

However, the m e an c le a ra n c e s

re­

c a l c i u m i n c r e m e n t , a nd t h u s p r o b a b l y m e a s ­

sig n if ic a n t b io lo g ic a l phenomenon.

ex p erim ental dem onstration
s k e l e t a l g r o w t h in t r a c e r

of the

studies

is evident fro m the foregoing,
tra n sie n ts from

was very

T h i s i s a c t u a l l y the f i r s t

r e l a t i o n of c l e a r a n c e
of t h e

s k e le to n .

v a l u e s to

It w a s b a s e d , a s

on d i s t i n g u i s h i n g the e f f e c t s

t h e g e n e r a l l y m u c h s l o w e r and p r o b a b l y

of

steadier

86
biological grow th p r o c e s s e s .
have m a d e

Bauer,

is

(1955)

a s i m i l a r d i s t i n c t i o n and have p r e s e n t e d a d i f f e r e n t

m ath em atical approach based
Their

C a r ls s o n and Lindquist

on C a l c i u m - 4 5

s t u d i e s in the

rat.

a p p r o a c h i s not a s d i r e c t a s t h a t u s e d h e r e i n , h o w e v e r ,

still r e g a r d e d a s untested.

extend t h e i r
clearance

and

H o w e v e r , t h e y c l a i m to be a ble to

c o n c l u s i o n s f u r t h e r t h a n s e e m s to be w a r r a n t e d by a

approach.

F in ally , the diffe ren ce between c le a ra n c e s

c a l c u l a t e d f o r the

d i f f e r e n t t i m e p e r i o d s w a s v e r y l a r g e - - m u c h l a r g e r t h a n c a n be
a c c o u n t e d f o r by e x p e r i m e n t a l e r r o r .
parent

r e l a t i o n t o a n y th i n g

tio n data.

s e.

be r e g a r d e d
were,

a specific m e a s u r e

as a v e ry dynam ic

or a b so rp ­

clearances,

of t h e m e t a b o l i c a c t i v i t y of

If t h i s i s e v e n a p p r o x i m a t e l y t r u e , t h e
system , turning

skeleton m u st

on and off a s it

but a l w a y s i n a c h a n g in g b a l a n c e b e t w e e n i t s a n a b o l i c and

catabolic p r o c e s s e s .
since

noted in the intestinal t r a n s i t

One i s f o r c e d t o c o n c l u d e , t h e n , t h a t i n th e

one p r o b a b l y h a s
bone p e r

T h e f l u c t u a t i o n s have no a p ­

In th e g r o w i n g

chick, t h a t i s p a r t i c u l a r l y t r u e ,

a s t h e b o n e s g r o w , m u c h of the bone f i r s t l a i d d own h a s to

be r e m o v e d an d i n e f f e c t ,
The

’’l a i d down s o m e p l a c e e l s e . "

statistical aspects

of t h e m e a s u r e d

s h o u l d be m e n t i o n e d .

Mean clearances

le s s than

standard e r r o r

10 p e r c e nt

skeletal c le aran ces

c a l c u l a t e d o v e r 4 h o u r s had

(thus

11.3 ± 1.1) s in c e a l l of

87
the

blo od v a l u e s t h r o u g h f o u r h o u r s

The

skeleton values, however, are

group

(a bo u t 20 o b s e r v a t i o n s ) .

counts
error

was about

8 per

c o n t r i b u t e d to the
b a s e d on t h e

The

s in g le 240 m i n u t e

standard e r r o r

c e n t of the m e a n ,

In b r ie f , the m a j o r

ce nt of the

The fa irly larg e p e rcen tag e

of l e s s e r

standard

v a r i a t i o n i n t h e fi n al f i g u r e a r i s e s

v a r i a t i o n i n bone a c t i v i t y .
b lood l e v e l a r e

of t h e f e m u r

w h e r e a s the

of t h e i n t e g r a t e d blood l e v e l s w a s only 5 p e r

mean.

f in al f i g u r e .

from

v a r i a t i o n s in

consequence.

Taking the individual tim e p e rio d s

s e p a r a t e l y , the

standard

e r r o r i n e a c h p e r i o d t e n d s to be r e a s o n a b l y u n i f o r m a t a bout 6 to
8 m illiliters
w ere

2
p e r m inute p e r M .

observed.

w h eth er the

However,

clearance

norm al,
of a bo u t
P

but f a i r l y

genetic
et al.

standard,

This

selection.

The

wa s a

v a r i a t i o n i n t h i s p o p u la t i o n of c h i c k s
2,

(2 X s t a n d a r d e r r o r =

substantially

r e d u c e d by

s t r a i n u s e d by T i t u s et a l . (1953) and Kennedy
Vitam in D a s s a y s ,

p e c t e d to be m u c h m o r e
w i t h i n the

standard e r r o r

This im plies that th e re

v a r i a t i o n m i g h t be

(1953) f o r t h e i r

viduals

same.

o r negative, its

12 m i l l i l i t e r s p e r m i n u t e p e r M

of 0.0455).

low er e r r o r s

one g e t s t h e i m p r e s s i o n t h a t no m a t t e r

was positive

w i l l p r o b a b l y be a bo u t t h e

In a few c a s e s ,

in p a r t i c u l a r , m i g h t be e x ­

u n i f o r m , p a r t i c u l a r l y if s e l e c t i o n of i n d i ­

stra in were practiced.

T h e e v a l u a t i o n of s t r a i n

88
d ifferences

in s k e le ta l

calcium

c l e a r a n c e s would be a m o s t

valu­

a b l e e x t e n s i o n of t h i s p r e s e n t w o r k.

V a r i a t i o n due to h a n d l i n g .
citable

Since the c h i c k e n i s a r a t h e r

a n i m a l a n d t h e Single C om b White L e g h o r n i s

ex­

thought to

b e p a r t i c u l a r l y so , i t s e e m e d a d v i s a b l e to c o m p a r e d a t a c o l l e c t e d
w it h d i f f e r e n t m e t h o d s

of h a n d l in g .

T h e tw o e n d p o i n t s

c o m p a r e d w e r e the

rate

m o v e d f r o m t h e u p p e r gut and the a m o u n t of Sr
skeleto n

90

at w h ic h Sr

90

d e p o s i t e d in t h e

( T a b l e XI).

The ex p erim en tal differences
birds

in g r o u p s of t e n ,

large

groups

in groups

(ten b i r d s ) w e r e

w ith the isotope

s o lu t i o n ,

sacrificed.

sm all groups

The

consisted

s i m p l y i n ha ndling the

of f o u r , and a s i n d i v i d u a l s .

r e m o v e d f r o m the b a t t e r y ,

T he

injected

and r e t u r n e d to th e b a t t e r y u nt i l t h e y w e r e
(four b i r d s ) w e r e

re m o v e d fro m the

b a t t e r y a nd a c c l i m a t e d in s m a l l c a g e s f o r one day p r i o r to i n j e c ­
t io n .

T h e y w e r e t h e n i n j e c t e d and r e t u r n e d to t h e

sacrificed.

T he i n d i v i d u a l b i r d s w e r e

to which they w e re

s m a l l c age u nt il

a c c l i m a t e d in i n d i v i d u a l c a g e s

r e t u r n e d a f t e r inje ctio n until they w ere

F e e d a nd w a t e r w e r e

sacrificed.

s u p p l i e d ad l i b i t u m and t h e t e m p e r a t u r e

m a i n t a i n e d at a p p r o x i m a t e l y
that the e n v iro n m e n t was

90

F.

by i n f r a r e d h e a t in g l a m p s

c o m p a r a b l e to t h a t of th e b a t t e r y .

w as
so

89
T A B L E XI
EFFECT

O F CHICK HANDLING ON S r ^ M O V E M E N T
AND A B SO R P T IO N
(all v a l u e s g i v e n as m e a n ± sta.nd-a.rcl e r r o r )

Treatm ent

Method, of
Handling

90
P e t . Sr
UPP' e r Gut

120 m i n .
control
. .

groups

of t e n

23

4

120 m i n .
lactose
. .

groups

of t e n

21

3

120 m i n .
control
. .

individual

23 ± 4

120 m i n .
lactose
. .

individual

32 dh 3

240 m i n .
control
. .

groups

of t e n

6 ±

4

240 m i n .
lactose
. .

groups

of t e n

10 ±

3

240 m i n .
control
. .

groups

of f o u r

5 ±

2

240 m i n .
lactose
. .

groups

of f o u r

2 ±

1

240 m i n .
control
. .

individual

24 ±

5

240 m i n .
lactose
. .

individual

35 ±

9

S ta tistic a l pro bability.

2

NSD = no s i g n i f i c a n t d i f f e r e n c e .

P

1

q 90
P e t . Sr
Ske leton

P

18 ± 1

NSD2

26 ± 3

1.1%

12 ± 1

7.2%

11

1

NSD

16 d: 2

NSD

14

2

NSD

31 da 3

NSD

19 at 1

1%

20 ± 1

NSD

12 ± 2

1%

90
H a n d l e d i n a g r o u p of t e n , t h e r e
lactose

a n d c o n t r o l g r o u p s , e x c e p t t h a t at

tion, m o r e
group.

90

was

c e n t of t h e

segment

is

Sr

recovered

T his difference

22 p e r

only

w a s no d i f f e r e n c e

120 m i n u t e s a f t e r i n j e c -

s k e l e t o n of t h e l a c t o s e - f e d

w a s gone a t 240 m i n u t e s ,

injected dose

of t h e b o w e l of t h e

8 per

in the

w as

b et w e en

however.

Since

r e c o v e r e d f r o m th e u p p e r

c o m b i n e d g r o u p s at

c e n t a t 240 m i n u t e s , the d i f f e r e n c e

12 0 m i n u t e s and

between these tim e s

r - 0 . 00843t
6 0 . 5e

g i v e n by:

W hen p l a c e d i n i n d i v i d u a l c a g e s , n e i t h e r l a c t o s e - f e d n or
c o n t r o l c h i c k s l o s t any Sr
120 and 240 m i n u t e s .
l a t i o n of t h e
em ptying.
m inutes
ner

c an be li tt l e doubt the n, t h a t i s o ­

eac h other
90

a m o u n t of S r

seriously

i n t e r f e r e d w ith c r o p

d e p o s i t e d i n th e

s k e l e t o n at

w a s l o w e r in i n d i v i d u a l b i r d s in a highly

in b o th l a c t o s e

a nd c o n t r o l g r o u p s .

c o n t r o l d a t a a t 240 m i n u t e s
v id u a ls and b i r d s
handled in groups

In g e n e r a l ,

retention

of

S r 9°

due to the

However, those

90

c o n t r o l g ro u p .

the

and

90

birds

r e t e n t i o n value

T h i s g r o u p was

r e t e n t i o n t h a n any o t h e r g r o u p m e a s

how ever, these
by

significant m a n ­

Combining l a c t o s e

of f o u r had a m u c h h i g h e r Sr

m u c h h i g h e r in s k e l e t a l S r

120

r e s u l t e d i n no d i f f e r e n c e b e t w e e n i n d i ­

f r o m th e l a r g e r g r o u p .

which w as p r i m a r i l y

ured.

f r o m the u p p e r d i g e s t i v e t r a c t b e tw e e n

Thus, there

chicks fro m

The

90

skeleton

r e s u l t s i n d i c a t e d t h a t the net
was

not

affected

by

in di vi du a l

91
handling
segm ent.

despite

the

effect

of

slo w er

Sr

90

transit

from

the

upper

92
P enicillin E xperim ents

B ody Weight

The
will

be

second phase

discussed
general,

in

of

the

results

this

same
of

t h is

experim ent
order
series

as

(the

the

will

penicillin

lactose

be

study)

study

com pared

was,

to

that

s tu d y .
T h e i n c l u s i o n of p r o c a i n e p e n i c i l l i n G in the r a t i o n at a
l e v e l of
Single

15 p a r t s

grow th ra te
are

a nd N o r r i s
was m o s t

not t h o s e

have been

(1952) sh o w e d t h a t the

(1952)

of c h i c k e n s

e f f i c i e n t i n the f i r s t f o u r w e e k s , t h e s e

expected.

However,

m ore

w orkers

chickens

are

that

reared

response

of

th e

environm ent.

example

i n c r e a s e d the g r o w t h

to

It h a s a l s o b e e n shown by

antibiotics

in old, c o n t a m i n a t e d a r e a s

that r e s p o n s e

For

w h e n c h i c k s w e r e k e p t on the f l o o r t h a n

i n c l e a n a r e a s ( B i r d et a l . ,
appears

re­

e n v i r o n m e n t a l c o n di ti o n s

s h o we d t h a t a n t i b i o t i c s

w h e n r e a r e d in w i r e - f l o o r e d b a t t e r i e s .
various

s t i m u l a t i o n of

s h o w n to a f f e c t r e s p o n s e to a n t i b i o t i c s .

Saxena et al.
rate

r a t e of g r o w t h of

C o m b W h i t e L e g h o r n c h i c k s to 2 w e e k s of a ge ( T a b l e XII).

S in c e H e u s e r

sults

p e r m i l l i o n did not a f f e c t the

is

greater

t h a n when r e a r e d

1952; a n d C o a t e s et a l ., 1951).

to a n t i b i o t i c s
S in c e

these

m a y be
birds

w h en

Thus it

r e l a t e d to c o n t a m i n a t i o n

were

reared

in

electric

93
TABLE

XII

E F F E C T O F P E N I C I L L I N ON BODY AND G U T W E I G H T
(all v a l u e s g i v e n a s m e a n ± s t a n d a r d e r r o r )

Tim e

and

N o . of
Chicks

T reatm ent

Body
Weight
( g m s .)

F

1

7 .5 m i n . c o n t r o l g r o u p ................
. . .
7 .5 m i n . p e n i c i l l i n g r o u p

10
10

165 ± 3.4
168 rfc 4.2

NSD

15 m i n . c o n t r o l g r o u p ....................
15 m i n . p e n i c i l l i n g r o u p ................

10
10

131 rfc 3.3
137 ± 5.0

NSD

30 m i n . c o n t r o l g r o u p ...................
30 m i n . p e n i c i l l i n g r o u p ................

10
8

144 ± 3.8
1 43 rfc 4.3

NSD

50 m i n . c o n t r o l g r o u p .......................
50 m i n . p e n i c i l l i n g r o u p ...................

11
11

131
138

2.8
4.1

NSD

rfc

90 m i n . c o n t r o l g r o u p ................... . .
90 m i n . p e n i c i l l i n g r o u p ................ . .

9
9

165 ± 6.8
164 ± 5.2

NSD

120 m i n . c o n t r o l g r o u p ................... . .
120 m i n . p e n i c i l l i n g r o u p ................

8
9

173 rfc 4.7
170 ± 4.5

NSD

10
9

93 rfc 2.7
99 rfc 2.8

NSD

240 m i n . c o n t r o l g r o u p ....................
240 m i n . p e n i c i l l i n g r o u p ............... ...

10
10

87
99

1.7%

X control group
X p e n i c i l l in g r o u p ......................................

78
76

180 m i n .
180 m i n .

c o n t r o l g r o u p ....................
p e n i c i l l i n g r o u p ...............

rfc
rfc

2.7
4.0

137 ±
X' 43
140 ± 1.5

NSD

S tatistical probability.
2
3
errors

NSD = no

significant d ifferen ce.

T he s t a n d a r d
of e a c h g r o u p .

error

was

c o m p u te d by com bining

standard

2,

94
TABLE

Uppe r
G ut Wt.
p e r 100
gms .

9.6
9.9

rfc

0.1
0.2

9.2 ± 0.2
8.8 rfc 0.4
9.7 rfc 0.2
9.0 ± 0.2
9.6
9.0

rfc

8.7
9.0

rfc

9.9
9.1

rfc

rfc

rfc
rfc

0.6
0.4
0.2
0.2
0.3
0.3

10.0 ± 0.4
9.2 rfc 0.1
9.0 ± 0.2
10.3 ± 0.2
9.7 ± 0.1
9.4 ± 0.1

P

XII ( C o n t i n u e d )

Lowe r
Gut Wt.
p e r 100
gms.

NSD

8.7 ± 0.2
7.6 ± 0.2

NSD

10.2 ± 0.4
8.6 ± 0.5

1%

8.8 rfc 0.2
8.8 rfc 0.2

NSD

9.7 ± 0.7
8.8 rfc 0.6

NSD

8.2 rfc 0.2
8.7 rfc 1 .0

5.7%

8.2
8.5

rfc
rfc

0.3
0.2

5.1%

1 1 .3 rfc 0.1
10.6 ± 0.2

1%

10.5 ± 0.3
11.0 rfc 0.3
0.1
±
0.2
9-1

9-2
3.2%

P

T otal Gut
p e r 100
g m s . Body
W e ig h t

P

1%

18.3 ± 0.2
17.9 rfc 0.3

NSD

1 .2%

19.4 rfc 0.4
17.4 rfc 0.6

1%

NSD

18.4 rfc 0.3
17.7 db 0.3

NSD

NSD

19.3 ± 0.9
18.8 ± 1.0

NSD

NSD

17.3
17.5

0.3
1 .0

NSD

NSD

18.1 ± 0.4
17.2 rfc 0.4

NSD

1%

21.3 rfc 0.4
19.8 ± 0.2

1%

0.4
0.4

1%

0.2
0.2

NSD

rfc

rfc

NSD

19.5
21.3

rfc

NSD

19.0
18.5

rfc

rfc

95
batteries

with r a is e d

s c r e e n flo ors that were frequently

s t i m u l a t i o n of g r o w t h would not n e c e s s a r i l y

cleaned,

be e x p e c t e d .

Gut W eights

The

w e i g h t of b oth u p p e r a nd l o w e r gut s e g m e n t s w as d e ­

t e r m i n e d w i t h i n g e s t e d f e e d and. o t h e r
sults

(Table

XII)

c o n t e n t s i n c lu de d.

s h o w e d t h a t the p r e s e n c e

The

of p e n i c i l l i n i n the

re­
ration

r e d u c e d t h e w e i g h t of b oth u p p e r and l o w e r gut s e g m e n t s in s e v e r a l
cases.

C o m b i n i n g th e d a t a at d i f f e r e n t t i m e s into one f i g u r e

XII), a s i g n i f i c a n t
associated
peared

r e d u c t i o n of the u p p e r gut s e g m e n t weight w as

w i th p e n i c i l l i n f e e din g.

b e t w e e n the l o w e r
Although th e s e

No s ig n i f ic a n t d i f f e r e n c e a p ­

se g m e n t weights.

results

d u c e t h e w e i g h t of t h e

s u g g e s t t h a t p e n i c i l l i n t e n d e d to r e ­

c h i c k ' s gut, th e

in m a g n i t u d e t o t h o s e

of C o a t e s

approxim ately

cent with s i m i l a r

t o note t h a t
m arked

17 p e r

Coates

obtained

results are

as

not c o m p a r a b l e

(1953), who showed d i f f e r e n c e s of
groups.

It is i n t e r e s t i n g

s u c h d i f f e r e n c e s with c h i c k s

s t i m u l a t i o n of g r o w t h due to p e n i c i l l i n .

weight fails to i n c r e a s e
the

(Table

r a p i d l y a s the

showing

P e r h a p s the gut

body we ight.

However,

only g r o u p in t h i s e x p e r i m e n t w h ic h showed a s i g n i f ic a n t weight

increase

due to p e n i c i l l i n ,

digestive

tract.

also p o s s e s s e d a h ea v ier than control

96
Bone

The

rig ht fe m u r

W eig hts

of b i r d s f r o m e a c h fe e d i n g r e g i m e

at each

tim e p e rio d was

c l e a n e d and w e ig h e d when r e m o v e d f r o m the

T h e wet w e i g h t s

of t h e

cept in the

c l e a n f e m u r w e r e not u s u a l l y d i f f e r e n t , e x ­

l a s t two g r o u p s

Both the

(T able XIII).

n o n a s h e d a nd the a s h e d weight of the f e m u r f r o m

thep e n ic illin s u p p le m e n te d
c o n t r o l (highly

chick.

180

minute

g r o u p w a s h e a v i e r t h a n it s

s i g n i f i c a n t ) a s w a s a l s o t he

c a s e at 240 m i n u t e s

(significant).
T he
those

r e s u l t s f r o m th e

of R o s s

and Yacowitz

g r a m s p e r t o n of r a t i o n
are

also

180 a nd 240 m i n u t e

s i g n i f i c a n t l y i n c r e a s e d the bone a s h .

c o n s i s t e n t w i th t h o s e of M i g ic o v s k y e t al.

However, th e re

in t h i s i n s t a n c e .
creased
was

m inute

group,

not be d e t e c t e d

c e n t and 4.7 p e r

respectively.
w i th the Sr

on s k e l e t a l S r

of Ca

45

s k e le t o n .

XIV)

,

in­

H o w e v e r , the i n c r e m e n t

c e n t i n the

180 m i n u t e a nd 240

Such s m a l l d i f f e r e n c e s p r o b a b l y
90

signifi-

90 ,
(Table

bone a s h c e r t a i n l y i n d i c a t e s

c a l c i u m d e p o s i t i o n by the

They

(1951), who

s k e l e t a l uptak e

w a s no e f fe c t

The i n c r e a s e d

only '7.1 p e r

with

(1954), who showed the p e n i c i l l i n a t 2.4

sh o we d t h a t p e n i c i l l i n i n c r e a s e d t h e
cantly.

groups ag ree

counting t e c h n i q u e s e m p l o y e d .

can

97
T A B L E XIII
EFFECT

T im e and
T reatm en t

OF PENICILLIN ON FEMUR WEIGHT

No. of
C hicks

2

m g . A sh ed
F em u r
p e r 100 g m s .
Body Weight

m g. F em ur
p e r 100 g m s.
Body Weight

10
10

875 ± 17
871 ± 7

NSD

c o n t.
pen. . .

10
10

801 ± 26
824 ± 19

NSD

30 m i n . c o n t.
30 m i n . p e n . . .

10
8

853 ± 18
777 ± 41

NSD

c o n t. .
p e n. . .

11
11

879 ± 11
888 ± 22

NSD

90 m i n . c on t.
90 m i n . p e n . . .

9
9

903 ± 27
910 ± 24

NSD

c on t.
pen. . .

8
9

909 ± 24
907 ± 14

NSD

180 m i n . c o n t. .
180 m i n . p e n . . .

10
9

781 ± 12
840 ± 7

cont.
pen. . .

10
10

770 ± 12
806 ± 13

7. 5 m i n .. c o n t.
7. 5 m i n . p e n . ^
15 m i n .
15 m i n .

50 m i n .
50 m i n .

120 m i n .
120 m i n .

240 m i n .
240 m i n .

S t a t is t ic a l p ro b a b ility .
C ontrol group.
P e n i c i l l i n group.
NSD = no sig n if ic a n t d iffe r e n c e .

A

1%

106.4 ± 1.3
114.0 ± 2.6

1%

5%

104.8 ± 1.9
109.7 ± 1.4

5%

98
TABLE

XIV

DISTRIBUTION OF S r 9° RECOVERED, P E N IC IL L IN E X P E R IM E N T
(all v a l u e s g i v e n a s m e a n ± s t a n d a r d e r r o r )

Per
Tim e

and T r e a t m e n t

No. of
Chicks

Cent S r 9 °

Uppe r
G ut

P 1

7 .5 m i n . c o n t r o l g r o u p .......................
7 .5 m i n . p e n i c i l l i n g r o u p
...............

10
10

71
77

3
2

NSD

15 m i n .
15 m i n .

c o n t r o l g r o u p ...........................
p e n i c i l l i n g r o u p .......................

10
10

55 ± 3
55 ± 2

NSD

30 m i n .
30 m i n .

c o n t r o l g r o u p ...........................
p e n i c i l l i n g r o u p .......................

10
8

44
41

3
4

NSD

50 m i n .
50 m i n .

control group . . . . . . . .
p e n i c i l l i n g r o u p .......................

11
11

36 ± 5
34 rfc 4

NSD

90 m i n . c o n t r o l g r o u p ........................... . .
90 m i n . p e n i c i l l i n g r o u p ....................... • -

9
9

25 ± 3
17 rfc 3

5%

120 m i n . c o n t r o l g r o u p ........................... . .
120 m i n . p e n i c i l l i n g r o u p ....................... . .

8
9

29
33

180 m i n .
180 m i n .

c o n t r o l g r o u p ...........................
p e n i c i l l i n g r o u p ....................... • .

240 m i n . c o n t r o l g r o u p ...........................
240 m i n . p e n i c i l l i n g r o u p .......................

1

10
9
10
10

rfc

rfc

rfc

rfc

rfc

5
2

15 ± 5
2 db 1
2
2

rfc

1
1

NSD

1%

NSD

S tatistical probability.

C a l c u l a t e d v a lu e p e r c e n t S r
in s k e l e t o n =
(c s p / f e m u r ) / (8 .84 ) / s t a n d a r d .
F a c t o r 8.84 is e x p l a i n e d i n th e a p p e n ­
dix .

4

99
T A B L E XIV (C o n tin u ed )

P e r Cent Sr
Low e r
Gut

P

20 ± 3
14 ± 2
29
35

±

48
54

±

±

±

3
2
3
5

39 ± 5
48 ± 5
55 ± 3
61 ± 9
39 ± 3
55 ± 2
21
16

±

3

±

2

14

±

2

11

±

2

-a,

S k eleton

2

0.2
0.2

90

P

NSD

1.5
1.3

NSD

1.1 db 0.1
1.3 ± 0.2

NSD

0.092 db 0.007
0.097 ± 0.012

NSD

3.7
3.6

0.3
0.3

NSD

0.101 ± 0.009
0.090 ± 0.008

NSD

NSD

3.7 ± 0.4
3.6 ± 0.3

NSD

0.078 ± 0.008
0.067 ± 0.004

NSD

NSD

4.9 ± 0.2
6.2 db 0.4

5%

0.056 ± 0.005
0.053 ± 0.004

NSD

1%

15.9 ± 1.2
13.1 ± 1 .3

NSD

0.057 ± 0.005
0.044 ± 0.002

1.6%

NSD

16.6 ± 1.5
16.5 dk 0.8

NSD

0.020 ± 0.002
0.012 ± 0.001

1%

NSD

18.4
16.5

NSD

0.018 ± 0.003
0.013 db 0.002

NSD

±
±

±
±

±
±

1.3
1.2

NSD

0.051
0.040

0.003
0.002

1%

NSD

90

C a lc u la te d value p e r c e n t Sr
( c p s / m l . ) X 0.0 0 3 1 5 .
F a c t o r 0.00315 is
4

p e r m l.
Blood^

P

NSD = no sig n ific a n t d iffe r e n c e .

dz
±

p er m i l l i l i t e r of blood =
exp la in ed in the appendix.

100
The

e x p l a n a t i o n of t h e s e

l a t t e r two g r o u p s being d i f f e r e n t in

f e m u r a s h w e i g h t m i g h t be a c c o u n t e d f o r by a d i f f e r e n c e
weight.

A l l o t h e r g r o u p s i n the p e n i c i l l i n s e r i e s

2 -l/Z

weeks

weeks

old.

of a g e ,

w hile th e

of t h e

injected

scribed, p rim a rily from
ever,
m ents
be

s i n c e th e d a t a
w ere very

Table

were approxim ately

180 a nd 240 m i n u t e g r o u p s w e r e

Movement

The fate

in age and

2

of S t r o n t i u m - 90

radioactive

s t r o n t i u m w ill be d e ­

XII, in c h r o n o l o g i c a l o r d e r .

c o l l e c t e d f r o m the p e n i c i l l i n s e r i e s

s i m i l a r to the la c to se

chiefly a c o m p a r i s o n b etw een th e s e

How­

of e x p e r i ­

s e r i e s , the d i s c u s s i o n w i l l
e x p e r i m e n t s a nd the l a c t o s e

experim ents.

Loss
c e n t of S r

f r o m the u p p e r gut.

90

in m u c h the

f r o m the

5 and

u p p e r gut o c c u r r e d in the f i r s t

of l o s s i s not quite a s

7.5 m i n u t e s

great

(Table

XIV,

6).

In only two i n s t a n c e s
significant d iffe re n c e s
T herefore,

of a p p r o x i m a t e l y 2 6 p e r

sam e m a n n e r a s that previo usly d isc u sse d for lactose;

how ever, the m agnitu de
F igures

A lo ss

out of e ig ht c o m p a r i s o n s w e r e t h e r e

betw een p en icillin —
fed and c o n t r o l g r o u p s .

th e d a t a w e r e

c o m b i n e d a nd u s e d to c o n s t r u c t F i g u r e 6.

101

bJ
CL
UJ

<2>

^

UJ

UJ §

XO

K-J

UJ

<0

io

ro

f\i

J.N3WQ39 d 3 d d O Nl * d S %

O

102
SINCE DIFFERENCES WERE INFREQUENTLY NOTED BETWEEN
CONTROL AND PENICILLIN TREATED GROUPS, THIS FIGURE
IS CONSTRUCTED FROM COMBINED MEANS. THE STANDARD
ERRORS OF EACH POINT INCREASES FROM 2.4 AT 7.5
MINUTES TO 29.4 AT 180 MINUTES. THE CORRELATION
COEFFICIENT WAS CALCULATED FROM THE MEAN POINTS
ON THE FIGURE AND NOT FROM INDIVIDUAL BIRDS.

%

SRWREMAINING IN UPPER

SEGMENT =

73.8 e

T\/z - 53.4

|txy = 0.965

J20
>80
MINUTES
FIGURE 6. RATE OF SR90 PASSAGE FROM THE UPPER TO
THE LOWER BOWEL. f PENICILLIN AND CONTROL GROUPS
ARE COMBINED')

103
A g a i n it i s n o te d t h a t t h e

rate

of e m p t y i n g f r o m the u p p e r to the

lo w er

seg m e n t took place

at an expon ential r a te ,

h ig h l y

c o r r e l a t e d with t im e .

The m o v e m e n t f r o m the u p p e r gut in

this

series

of e x p e r i m e n t s a l s o

T he

initial

11s p u r t ' '

by t h e e q u a t i o n :

o c c u r r e d a t two d i f f e r e n t r a t e s .

which was very

Sr

rapid

u p p e r = 26.2e

i s g iv e n a p p r o x i m a t e l y
, and the

w a s a s s u m e d to be due to n o r m a l m u s c u l a r
seg m en t is
tim e
per

g i v e n by:

p e r cent Sr

i n m i n u t e s in b o t h c a s e s ) .
, ^
c ent S r m u p p e r
While t h e

portant basic principle
u p p e r to the

w a s h ig hl y

rates

in t h e

f e e d in g

(t is

- 0 .0 1 2 97t

,
- 0. 6t
+ 2 6.2e

s lightly d i f f e r e n t in t h i s

series

l a c t o s e e x p e r i m e n t s , the i m -

o 90
b r o u g h t out in both c a s e s i s t h a t Sr
moves
low er

s i m i l a r to those

different, the
data

- 0 0129 7t

s e g m e n t at a n e x p o n e n t i a l r a t e

which

c o r r e l a t e d w ith t i m e .

t h a t w hile the

these

which

c o m b i n e d e x p r e s s i o n , th en , is:

M o v e m e n t t h r o u g h t h e l o w e r gut.
a nd f i g u r e s

slo w r a t e

c o n t r a c t i o n of the u p p e r

u p p e r = 7 3 . 8e

T he

of t r a n s i t a r e

of e x p e r i m e n t s t h a n t h e

f r o m th e

90

s e g m e n t = 7 3 . 8e

rates

w hic h was very

values

A construction

of the l a c t o s e

series

regim e

showed a g a i n

o b t a in e d in the p e n i c i l l i n study a r e

g e n e r a l p a t t e r n i s v e r y m u c h the

c a n be

of t a b l e s

same.

slightly

T herefore,

c o m p a r e d with t h o s e d i s c u s s e d u n d e r the l a c t o s e

(Figures

7 a nd

8, T a b l e s XV and XVI).

104

o

o

o

if)
^
fO
J.N 3W 03S U3MOT Nl *H S %
< 0

105

CALCULATED

jO

70

40

OBSERVED

DIFFERENCE

COMBINED
ABSORPTION
AND DEFECATION
US
%

ABSORPTION

OF

INITIAL SLUG

180

240

MINUTES
FIGURE 8. SR90RECOVERIES FROM LOWER BOWEL
(BASED ON COMBINED MEANS
FOR COMPARISON
WITH FIGURE 6)

106
TABLE

XV

C O M B IN E D C ON T R O L AND P E N I C I L L I N DATA,
BOWEL R EC O V ER IE S

Tim e
(m in.)

0
7.5

Pet. S r9°
in U pper
G ut

100
74 ±

P et. S r9°
in Lower
Gut

0

Calculated
P e t , S r 90
in L o w e r
G ut

0

Loss from
L o w e r Gut
(pet.)

0

2

17 ± 2

26 ±

2

9 ± 3

15

55 ± 2

32 ± 2

45 ±

2

13 ± 3

30

43 ± 3

51 ±

57 ± 2

6 ± 4

50

35 ± 3

44 ± 4

65 ±

3

21 ± 5

90

21 ± 2

58 ± 5

79 ± 2

21 ± 5

120

31 ± 3

42 ±

2

69 ± 3

27 ± 4

180

9 ± 3

19 ± 2

91 ± 3

72 ± 4

240

2 ± 1

13 ± 1

98 ± 1

85 ± 1

3

107
TABLE

XVI

M EAN P E R CE N T R E C O V E R IE S OF S r 9 ° - Y 9°
A F T E R CROP INJECTION
( c o m b i n e d d a t a f r o m c o n t r o l a nd p e n i c i l l i n - f e d g r o u p s )

T im e
( m i n .)

7.5 . .

.

15 . .

Upper
to
Lowe r

L ow e r

26

17

9

0

1

92

45

32

13

0

1

88

Dif­
fer­
ence

Feces

S k e l­
eton

Total
Recov­
ery^

30 . . .

.

57

51

6

0

4

98

50 . . .

.

65

44

21

0

4

83

90 . .

79

58

21

152

6

100

120 . .

69

42

27

122

15

100

180 . .

91

19

72

552

17

100

240

98

13

85

682

17

100

. . . .

1

90
D o e s n o t i n c l u d e p e r c e n t Sr
which m a y have
p r e s e n t in the sof t t i s s u e s a nd e x t r a c e l l u l a r fluid.

C alculated.

been

108
Again,
with tim e

a m ultiple t e r m

e q u a t i o n f o r the b e h a v i o r of s t r o n t i u m

in t h e l o w e r i n t e s t i n a l s e g m e n t m a y be c a l c u l a t e d ,

f lo w s h e e t

c o n s t r u c t e d w ith t h e

( se e A p p e n d i x ) .

same

g e n e r a l conclusions as before

How ever, in this penicillin s e r ie s

it m a y be c a l c u -

l a t e d t h a t d e f e c a t i o n did not b egin u n ti l 74 m i n u t e s
whereas
of the

in the la c to s e

Sr

series,

90

s e r i e s , the

a t the v ent w a s

em ptied

lo w er bowel was

somewhat

a f t e r i nj e c ti o n ,

c a l c u l a t e d t i m e for the a r r i v a l

57 m i n u t e s .

som ewhat m o re

and a

T h u s the c r o p , in t h i s

rapidly,

slower.

but t r a n s i t t h r o u g h the

C o n s e q u e n tl y , m o r e

of the

t r a c e r w as available to the

s k e l e t o n and f o r a l o n g e r t i m e .

th e le s s , in th is

s k e l e t o n in 240 m i n u t e s a c c u m u l a t e d

17.5 ±

0.9 p e r

series,

s e r i e s , the
cent

of th e i n j e c t e d S r

15.9 ± 1.3 p e r

90

-Y

90

w h e r e a s m the l a c t o s e

cent w a s a c c u m u l a t e d .

e v i d e n c e t h a t t h e m a j o r f a c t o r w h ic h r e g u l a t e s
not so m u c h th e q u a n t i t y
when larg e

differences

None­

This is very

clear

s k e l e t a l uptake i s

o f f e r e d (although t h i s i s c e r t a i n l y a f a c t o r

o c c u r ) , but the i n h e r e n t a b i l i t y of the

skeleton

i t s e l f to a c c u m u la te the t r a c e r .

U p t a k e of S r
level
as

of

the

S r 90

90

f ound

in

by

the

the

blood

blo o d

and

at

7.5 m i n u t e l e v e l i n the l a c t o s e

bone

7.5

depositio n.

minutes

experim ents.

p e r c e n t of S r 9 ° i n t h e b l o o d i n c r e a s e d a t

was

the

The
same

H o w e v e r, the

15, 30, and 50 m i n u t e s

109
to l e v e l s
groups

of t h e

le v els
were

w h i c h w e r e tw o to f o u r t i m e s t h o s e
lactose

of b lood Sr
again the

90

experim ents

of t r a c e r

vent.

( T a b l e s VI and XIV).

th e n fell until (from

w a s p r o b a b l y due to the

f r o m the u p p e r

s e g m e n t and i t s

T h e s e high

soo n e s t a b l i s h e d .

series.

somewhat m ore

r a p id

s l o w e r t r a n s i t to t h e

I n t e s t i n a l a b s o r p t i o n w a s e v i d e n tl y v e r y

b e t w e e n gut a n d blood w a s

comparable

1Z0 to 240 m i n u t e s ) they

s a m e a s t h e l e v e l s in t h e e a r l i e r

This difference
loss

found in th e

r a p i d , and e q u i l i b r i u m
The

sk e l e to n ,

however,

a p p a r e n t l y f a i l e d to c l e a r t h e blood a s w e l l a s in the e a r l i e r l a c t o s e
experim ents.

Since the p e n i c i l l i n t r e a t m e n t w as without e ff ec t , it

s e e m e d likely that th e se differen ces were

a s s o c i a t e d w ith the

what g r e a t e r body w e i g h t s due to o l d e r age of the

som e­

b i r d s in t h i s

serie s .
A s p o i n t e d out a b o v e , the p e r c e n t of S r
s k e l e t o n i n t h e two
The

sequence

of Sr

series
90

d e p o s i t i o n in the

60 and

w a s t h a t the

cillin t r i a l s ,
m inute

the l a r g e s t

period.

s e e m e d t o be

Sr

90

sim ilar

The

chief

while in the p e n i ­

i n c r e m e n t w a s d e l a y e d until the

"slow er"

same.

d e p o s i t i o n t o ok p l a c e b e tw e e n

lactose ex p erim en ts,

A g a i n , t h e n , the
somewhat

s k e l e t o n was v er y

study ( T a b l e s VI a nd XIV).

greatest

90 m i n u t e s i n the

r e c o v e r e d f r o m the

of e x p e r i m e n t s w as e s s e n t i a l l y the

to t h a t found in the l a c t o s e
difference

90

s e c o n d (penicillin)
t h a n the f i r s t

series

( l a c to se )

90 to

120

of c hi c ks
series.

110
It m i g h t be
flected t h e i r

s t a t e d t h e n , t h a t , p h y s i o l o g i c a l l y the

''o ld e r''

sa m e , the

Table

XVII

calculated

h i g h e r while the

of t h e t r a c t .

H o w e v e r , the

m ean clearance
These

w as

clearances

s p r e a d t h r o u g h the a b s o r p t i v e

T h e f i n a l (240 m i n u t e )

seg­

c le a ra n c e s were again

2
of 4 - 5 m i l l i l i t e r s p e r m i n u t e p e r M , and the

in t h e n e i g h b o r h o o d

2
per M .

case.

out high, t h e n f e l l p r e c i p i t o u s l y and r o s e a g a i n a s the

m a j o r p o r t i o n of th e i s o t o p e
m ents

s k e l e t a l uptake i s about

c l e a r a n c e s m u s t be l o w e r t h a n b e f o r e .

s h o w s t h i s to be the

again sta rte d

re­

a ge.

If b lo o d l e v e l s a r e
the

c h ic k s

f o r the whole 4 h o u r s i s
values a r e

b e e n s a id a b o u t t h e

6.1 m i l l i l i t e r s p e r m in u te

low and only e m p h a s i z e

' ' o l d e r ' 1 b e h a v i o r of the

what h as a l r e a d y

c h i c k s in the p e n i c i l l i n

serie s .
Although th is
a significant effect

second

series

of e x p e r i m e n t s fa iled to r e v e a l

of p e n i c i l l i n , they do, e v e n with t h e i r d i f f e r e n c e s ,

c o n f i r m t h e p h y s i o l o g i c p o i n t s c l a i m e d e a r l i e r f r o m the d ata
lacto se t r i a l s .
which some
t he

of t h e

skeleton.

passage
in t h e

from

strict

There
Sr

There

w a s no doubt of the e x t r e m e
90

r a p id i ty with

le ft the bowel and t r a v e r s e d the blood to

w a s , th e n , v e r y

l i ttle h i n d r a n c e

of s t r o n t i u m

gut to blood in the m u c o s a of the bowel.
sense,

p a s s into t h e blood,

was

of the

very

rapid.

Absorption,

If s om e a t o m s of s t r o n t i u m

why d o n ' t a l l of t h e m

get t h e r e ?

The d a t a a r e

Ill
T A B L E XVII
S K E L E T A L C L E A R A N C E OF S r 9 ° - Y 9 ° IN
T H E P E N I C I L L I N SERIES
90
B l o o d Sr
(pet./m l.)

7. 5

minutes . .

0.0454

'

S k e le to n
S r ^ 0 (pet.)

0.002

1. 4

±

0. 1 4

15

minutes

. . .

0.0942

±

0.007

1.2

±

0. 10

30

minutes

. . .

0.0958

±

0.006

3. 7

±

0.20

50

minutes

. . .

0.0725

±

0. 005

3. 6

±

0.30

90

minutes

. . .

0.0545

db 0 . 0 0 3

5. 5

120

minutes

. . .

0.0510

d= 0 . 0 0 3

14. 5

±

180

minutes

. . .

0. 0161

±

0. 0 01

240

minutes

. . .

0.0154

±

0. 003

ard

27. 1

±

0. 5

6. 6

±

2. 6

0. 2

d: 0. 4

0.20

2. 7

rfc 0. 4

0. 90

18. 7

16. 5

d: 0 . 9 0

3. 8

rfc 1.2

17. 4

±

0.90

4. 6

±

-

-

r

=b 2. 6

1. 3

1 T h e a m o u n t of whole b lood c l e a r e d by the
m inute p e r

ml. Blood/m in.
/ Skeleton/

±

2. 6

1.6

skeleton p e r

s q u a r e m e t e r of body s u r f a c e .

2 T h e s t a n d a r d e r r o r w a s c a l c u l a t e d by com bin in g the s t a n d ­
e r r o r of g r o u p s r a t h e r t h a n d e t e r m i n a t i o n of i n d iv id u a l d i f f e r ­

ences .

112
c o n s i s t e n t with the h y p o t h e s i s that, in e ffect, many do, but there
being

"no p l a c e to g o M a m ajor ity of them come

the bow el.
essarily

Then, if the

be d e f e c a t e d .

scribed as

right back into

sk e l e t o n can't acce pt them, they must n e c ­
The net e ffe ct i s what i s c l a s s i c a l l y d e ­

’ ’poor a b s o r p t i o n . ”

Actually, it s e e m s to be poor

skeletal m etabolism .
To r e t u r n to the a b s e n c e
pointed out that in t h e s e t r i a l s ,

of a p e n i c il l i n effect, it may be
15 p a rt s p e r m i l l i o n of pe nic illin

did not e x e r t an e ff ec t on growth.
not sh o w a b e t te r Sr
is true.

90

In that c a s e , the

skeleton should

in c orporation, if what i s conjectured above

T h u s, e v e n t h is fa cto r i s c on siste n t with the g en er al p i c ­

ture w hic h h a s e m e r g e d .

SUMMARY AND CONCLUSIONS

The
rate
th e

e f f e c t w h i c h two d i f f e r e n t f e e d in g r e g i m e s have

of i n t e s t i n a l t r a n s i t ,
s k e l e t o n of S r

90

-Y

90

blood c o n c e n t r a t i o n , a nd net r e t e n t i o n
has

b e e n s tu die d.

o th e r p h ysiological effects w ere
clusions

o b s e r v e d with the

ration

following c o n ­

c a n t m a n n e r a t tw o w e e k s
fem ur were

in o rganic a s h was noted.
and e x c r e t e d

m ore

chicks also

the evidence
2.

of age .

reduced

cent l e v e l , in a

r e d u c e d live weight and i n c r e a s e d the

w e ig h t of t h e a l i m e n t a r y t r a c t and i t s

rate

In ad dition , c e r t a i n

S u b s t i t u t i o n of l a c t o s e a t the 25 p e r

n o r m a l chick sta rtin g

These

by

resulting.

1.

of t h e

on the

c o n t e n t s in a highly

The m o i s t u r e

signifi­

and o r g a n i c m a t t e r

s i g n i f i c a n t l y ; h o w e v e r , no eff ect on the
C h i c k s on t h i s f ee ding r e g i m e

consumed

w a t e r t h a n c o n t r o l c h i c k s in a 4 h ou r p e r i o d .
r e t a i n e d m o r e w a t e r d u r i n g t h i s p e r io d d e s p it e

of d e h y d r a t i o n of bone t i s s u e .

L actose,

at 2 5 p e r

cent of the

r a t i o n , did not a ff ect the

of S r 9 ° - Y 9 ° t r a n s i t t h r o u g h th e d i g e s t i v e t r a c t e v e n though it

did i n c r e a s e
on t h e n e t

th e flow of w a t e r .

retention

of S r

90

T h i s l e v e l of l a c t o s e

by the

113

s k e le t o n .

had no e ffect

114
3.
starting

Penicillin,

diet,

at

15 p a r t s p e r m i l l i o n in a n o r m a l chick

had no e f f e c t on live body weight o r the weight of the

a l i m e n t a r y t r a c t with the

contents intact.

I n tw o c a s e s w h e r e the

chicks w ere

2 weeks

c i l l i n i n c r e a s e d t h e i n o r g a n i c a s h of t he f e m u r
highly
were

significantly.
approxim ately

w e t w e i g h t of t h e

of a g e , p e n i ­

si g n i f i c a n t l y and

H o w e v e r , in s ix o t h e r c a s e s w he re the
2~l/2

chic ks

w e e k s old, p e n i c i l l i n had no e f f e c t on the

fem ur.

A s h e d w e i g h t s of t h e s e

g r o u p s w e r e not

determ ined.
4.
transit

Penicillin,

like l a c t o s e , ha d no e ffe ct on the

o r n e t r e t e n t i o n by t h e
5.

s k e l e t o n of Sr

B a s i c a l l y , in b ot h s e r i e s

90

-Y

rate

of

90

of e x p e r i m e n t s , it w a s found
90

t h a t a f t e r i n j e c t i o n into t he

c r o p , the t r a n s i t of S r

gut o c c u r r e d a t two r a t e s .

The f i r s t r a t e , whic h o c c u r r e d b e t w e e n

0 a nd 7 - l / Z

from the upper

m i n u t e s a f t e r i n j e c t i o n , w as v e r y r a p i d a nd a c c o u n t e d

for approxim ately
was a sim ple

30 p e r

c ent of the i n j e c t e d d o s e .

The

s e cond

e x p o n e n t i a l r a t e w h ich w as highly c o r r e l a t e d w ith

tim e.
6.
described.
f o ll ow i ng
Sr^^ fro m

The

r e c o v e r y of Sr

From

90

from

.
the l o w e r i n t e s t i n e was

the m a t h e m a tic a l analy sis

interpretations

a p p e a r to be

of the t i m e c u r v e , the

required;

The a b s o r p t i o n of

t h e l o w e r b o w e l o c c u r r e d in two p h a s e s which w e r e

115
r e l a t e d t o t h e two m o v e m e n t s f r o m

the u p p e r bowel.

The in i t i a l

90
’’s p u r t 11 of S r
rapidly.

into the l o w e r gut w as p r o b a b l y a b s o r b e d

A l t h o u g h m u c h of t h i s

Sr

90

extrem ely

w as c l e a r e d f r o m the blood by

r a p i d e x c h a n g e w ith t h e s k e l e t o n , a c o n s i d e r a b l e p o r t i o n was not
accounted fo r.
sam pled
large

T h i s m i g h t h a v e b e e n t e m p o r a r i l y d e p o s i t e d in un­

s o ft t i s s u e s ,

p a r t of t h i s

o r it m i g h t r e p r e s e n t a c c u m u l a t e d e r r o r s .

e a r l y a b s o r b e d Sr

the l o w e r bowel and so m e was
second Sr

90

’ ’p o o l ”

was a b s o r b e d v e ry

90

was a l s o

r a p i d l y r e t u r n e d to

e x c r e t e d i n the u r i n e .

(that w hich m o v e d slowly f r o m
slowly f r o m the bowel.

A.

Much of the

the u p p e r gut)

Up to 40 p e r c e n t (of

the i n j e c t e d d o s e ) m a y h a v e b e e n e x c r e t e d w ithout being a b s o r b e d .
7.
ferent

The u p t a k e of S r

rates

m inutes

at least.

be distinguished.

in the

s k e l e t o n o c c u r r e d a t two d i f ­

T h e v e r y r a p i d r a t e ( m e a s u r e d within 7 - 1 / 2

a f t e r i n j e c t i o n of Sr

change phenomenon.

effects

90

90

into the c r o p ) was a p p a r e n t l y an e x ­

A. s l o w e r s k e l e t a l d e p o s i t i o n of t r a c e r m a y a l s o

T h is was

a p p a r e n t l y an o v e r - a l l a v e r a g i n g of the

of ne w s k e l e t a l g r o w t h and of the r e s o r p t i o n p r o c e s s e s

s o c i a t e d w it h s k e l e t a l
8.

The

rem odeling.

r e m o v a l of Sr

n ot a c o n s t a n t a n d u n i f o r m
fluctuating

as­

90

from

the blood by the s k e l e t o n was

s in g le p r o c e s s , but an i n t e r m i t t e n t and

r e s u l t a n t of a n u m b e r of p r o c e s s e s .

s i s t e n t d e p e n d e n c e upo n the c o n c e n t r a t i o n of Sr

T h e r e was no con90

i n the blood, o r

116
u po n i n t e s t i n a l

a b s o r p t i o n , w he n s h o r t t i m e p e r i o d s

C o n s e q u e n t l y , the m e t a b o l i c

p a t t e r n in the

were

considered.

s k e l e t o n i t s e l f was c o n ­

s i d e r e d to b e t h e p r i m a r y f a c t o r in the o v e r - a l l m e t a b o l i s m of

APPENDIX

117

118
TABLE

XVIII

90
90
EQUILIBRATION OF Y
DAUGHTER WITH Sr
PARENT
IN S A M P L E S O F D IG E S T E D IN TE S T IN E

Days A fter F i r s t
C ou n t on S a m p l e

Counts p e r Second p e r
4.5 m l . Aliquot of Sample

0

159

1

160

2

161

3

162

4

159

6

162

119
T A B L E XIX
DETERMINATION OF THE FACTOR USED TO CONVERT FEMUR
ACTIVITY TO ACTIVITY OF THE TOTAL SKELETON

Counts p e r Second
p e r A li q u o t of Total
S k e le to n A s h
11.8
17.3
12.7
12.2
10.3
22.8
12.6
13.7
20.3
16.6
17.0
16.3
17.6
15.5
19-7
20.9
11.9
19.2
17.3
19.4
18.7
18.8
26.8
12.9
28.1
27.2
24.6
24.0
32.0
26.4
17.8
26.5
608.9
X = 19

Counts p e r Second
p er F e m u r

104
146
111
111
93
210
122
113
172
150
138
147
148
129
163
163
102
180
142
164
157
160
245
116
277
260
226
226
304
240
162
226
5,407
X = 169

Ratio

8.81
8.84
8.74
9.10
9.03
9.21
9 *68
8.25
8.47
9.04
8.12
9.02
8.42
8.32
8.27
7.80
8.57
9.38
8.21
8.45
8.40
8.51
9.14
8.99
9.86
9.56
9.19
9.42
9.50
9.09
9.10
8.53
283.02
X = 8.84 ± 0.13

120
TABLE

XX

D E T E R M I N A T I O N O F T H E F A C T O R USED TO CONVERT WHOLE
B L O O D A C T I V IT Y F R O M COUNTS P E R SECOND P E R
M I L L I L I T E R TO P E R C E N T OF I N J E C T E D
DOSE P E R M I L L I L I T E R

Sample
Number

Counts p e r
Second p e r
M illiliter

1

137.3

2

131.5

3

124.7

4

130.6

5

123.7

6

129.4

7

122.1

8

124.1

9

122.7

10

124.7
X = 127.1 ± 1.4

F our-tenths

per

c e n t of the i n j e c t e d d os e was u s e d f o r e a c h s a m p l e

d e te r m in a tio n above.
T herefore,
0.4
X % of i n j e c t e d d o s e / m l .
127.1 ~
c p s / m l . blood
P e r c e n t of i n j e c t e d d o s e / m l . b lo o d = ( 0 . 4 /1 2 7 .1 ) X c p s / m l .

whole

b l o o d = 0.00315 X c p s / m l . whole blood.
4.5 m l . (of the 250 m l . s o lu ti on ) c o u n t e d w et a l s o a v e r a g e d 127
c p s / a l i q u o t , t h e s a m e a s the a c t i v i t y of 1 m l . a f t e r d r y i n g and ashing

121
T A B L E XXI
TUBE

CONVERSION F A C T O R S

Blood
1.

T ube

2340K to T u b e 2 - C C - 6 5

2.

T ube

2-CC-65

3.

T u b e 2 - A H - 9 1 to Tube 2 - D I - 2 9 = none ( i . e . , 1.0 X 2 -A H - 9 1 value

to

= 1,68 X 2 3 4 OK value

Tube 2 - D I - 2 9 = 0.83 X 2 - C C - 6 5

v alue

Bone
4.

Tube

2-EE-49

5.

T ube

2-

6.

T u b e 2340K to T u be 2 - C C - 6 5

7.

T ube

to Tub e 2 - D I - 2 9 = 0.86 X 2- E E - 49 v alue

E D - 12 to Tube

2-CC-65

to Tube

2 - D I - 2 9 = 1.96 X 2 - E D - 1 2

value

= 1.68 X 2 3 4 OK value

2 - D I - 2 9 = 0.83 X 2 - C C - 6 5 v alu e

Gut
8.

T ube

2340K to

9.

Tube

2-CC-65

Tube 2 - C C - 6 5

= 1.73 X 2340K value

to Tube 2 - D I - 2 9

= 0.83 X 2 - C C - 6 5 v alue

10.

T ube 2 - A H - 91 to Tube 2 - D I - 2 9 = none ( i . e . , 1.0 X 2-A.H-91 value

1 1.

Tube

2- E E - 49 to

Tube 2 - D I - 2 9 = 1.09 X 2 - E E - 49 value

122
TABLE
LOSS DUE

XXII

TO H O M O G E N I Z A T IO N OF I N T E S T I N A L S A M P L E S

Counts p e r Second
p e r A l i q u o t of
H om ogenized
Sample

Counts p e r Se co nd
of Whole S a m p l e

101.2

124.9

97.6

119.7

92.8

125.9

104.2

127.5

97.3

129.7

99.6

125.1

101.8

115.7

102.6

126.3

99.4

120.9

101.5

132.1

X = 99.8 ± 1 . 0

X = 124.8 ± 1.6

Patio

0.800 ± 0.013

123
TABLE

XXIII

STANDARD A CT I V IT Y VALUES
( c o u n t e d a f t e r s t a n d a r d d ilu ti o n a s
i n f i n i t e l y t h i c k l iq u i d s a m p l e s )

E xperim ent
Number

^ 90
90
Sr
-Y
Activity
(cps / a l i q u o t )

„ 90
90
T ot a l S r
-Y
A c t iv i ty *
(cps / a liq uo t)

1

104

125

2

106

127

3

105

126

4

107

128

5

98

118

6

98

118

7

-

143

C o r r e c t e d for hom ogenization lo ss.

124
T A B L E X XIV
on
qn
CALCULATION OF Sr
-Y
ACTIVITY IN THE BLOOD
(c le fts , taken f r o m the 240 minute control group
in E x p e r i m e n t 7, for ill u st ra t i o n )

Observed
cps / m l .

After
Background
Correction^

A fte r
Tube
Correction^

F inal P e t .
of Injected
D o s e per
m l. Whole
Blood3

1

5.5

5.0

4.3

0.0135

2

6.2

5.7

4.9

0.0154

3

4.7

4.2

3.6

0.0113

4

3.6

3.1

2.7

0.0085

5

7.8

7.3

6.3

0.0198

6

8.3

7.8

6.7

0.0211

7

12.1

11.1

10.1

0.0318

8

9.5

9.0

7.7

0.0243

9

4.2

3.7

3.2

0.0101

10

9.8

9.3

8.0

0.0252

X = 7.2

X = 6.7

X = 5.8

X = 0.0181

Sampl e
No.

1 Background c o r r e c t i o n = 0.5 count pe r second.
2 Tube c o r r e c t i o n = 0.86 x Tube 2 - E E - 4 9 value = Tube 2D I- 29 v a lu e .
3 P e r cent of in je ct e d d o se per m i l l i l i t e r whole blood =
0.00315 x counts p e r s ec on d per m i l l i l i t e r whole blood (after b a c k ­
ground and tube c o r r e c t i o n s ) .

125
T A B L E XXV
90
90
CA LCULATION OF Sr
-Y
PER CENT IN TOTAL SKELETON
(data taken f r o m the 240 minute contr ol group
i n E x p e r i m e n t 7, for i llu st r at io n )

Ob s e r v e d
cps /F e m u r

After
Background
C or re ction^

After
Ab s or b e r
Correction*^

F in a l P e t .
Sr9° - Y 9°
in Whole
Skeleton^

1

26.6

26.0

196

15.5

2

2 5.0

24.4

184

14.6

3

16.9

16.3

123

9.7

4

45.4

44.8

338

26.7

5

2 7.6

27.0

203

16.1

6

26.9

26.3

199

15.7

7

2 1.0

20.4

154

12.2

8

23.7

23.1

174

13.8

9

39.5

38.9

294

23.3

10

23.0

22.4

169

13.4

S a m p le
No.

c p s = counts p er sec on d of Sr

-Y

activity.

Background = 0.6 cps.
3 A b s o r b e r c o r r e c t i o n = 7.54 X c p s / f e m u r .
4 Skeleton per c ent = c p s / f e m u r / 8 . 8 4 / 1 4 3 .

per

143 = the counts

s e c o n d of the standard in E x p e ri m e nt 7.

No tube c o r r e c t i o n required;

counted with Tube 2 - D I - 2 9 .

126
TABLE

XXVI

C A L C U L A T I O N OF U P P E R GUT ACTIVITY
( d a t a t a k e n f r o m the 2 40 m i n u t e c o n t r o l
g r o u p in E x p e r i m e n t 7 f o r i l l u s t r a t i o n )

Final Pet.

O bserved
c p s / ali . 1

After
Background
Correction^

1

1.7

1.1

0.8

2

40.7

40.1

28.0

3

1.3

0.7

0.5

4

1.3

0.7

0.5

5

1.1

0.5

0.3

6

40.8

40.2

28.1

7

3.1

2.5

1.7

8

2.1

1.5

1.0

9

1.1

0.5

0.3

10

1.9

1 .3

0.9

Sample
No.

X

cps/ali.

=

X

9.5

= c o u nt s p e r

-

8,9

X -

6.2

s e c on d .

3 S r 9 ° p e r c e n t of the u p p e r gut a c t i v i t y of 143.

in U pp er
GuP

s e c o n d p e r a li q u ot .

B a c k g r o u n d — 0.6 c o u nt p e r

standard

S r 9 0 _ y 90

gut a c t i v i t y d ivid ed by

127
{% S r 9 °)

=

~ ° ,6t

low er

100 -

3 2 . 9e

- 0 . 0 0 9 66t

-67. le
- 0 . 0 0 5 1 8t

- 1 0 0 + 5 5 . 2e
- 0.08 085t
+ 7 1 6 1 4 2 . 64e

-0.08406t
- 8 6 0 8 8 1 . 64e

- 0 . 1 3208t
+ 3 0 6 5 8 3 . 45e

-0.18801t
- 1 8 8 6 8 8 . 07e

-0.6t
+

2 6 8 5 5 . 52e
- 0.6t

4

3 2 . 9e

100

67.1%
55.2%
57.3
m inutes

11.9%

rapidly absorbed
slowly a b s o r b e d

14.2%

41.0%

Figure

32.9%

9.

W o l t e r i n k e qu a ti o n , l a c t o s e

series

44.8
14,2
59.0
m inim um
to ta l
absorption

128
-

26 . 2e

01297t

- 7 3 . 8e
4->
o
o
1
—
1
o

-0,
+

o
i

low er

vO

-0

(% S r 9 °)

59 •9e
-0, .01686t

+ 156 . 81 e

-0 .0 3 2 0 9 t
-481 . 19 e

-0. 05 067t
+ 228, . 42e

- 0
-0. 01040t

+ 122. 15 e
■0. 10165t
+

13. 91 e

,100
73.8

\L

59.9
7 3.6
m inutes

*
27.9

Figure

26.2

\

\

13.9

32.0

10.

rapidly absorbed
slowly a b s o r b e d

W o l t e r i n k e qu atio n , p e n i c i l l i n s e r i e s .

40.1
32.0
72.1
m inim um
t o ta l
absorption

LITERATURE

CITED

A s h c r a f t , D. W,
E f f e c t of M i l k P r o d u c t s on t h e pH of I n t e s t i n a l
C o n t e n t of D o m e s t i c F o w l .
P o u l t r y Sci.
12: 2 9 2 - 2 9 8 (1933).
Barlow ,

J. S., a nd J.
Young C h i c k s .

F. M anery.
Fed. P ro c.

E l e c t r o l y t e s in the M u s c l e of
No. 1, P t . 1, 9: 8-9 (1950).

B a u e r , G. C. H., A. C a r l s s o n , a nd B. L i n d q u i s t .
E v a l u a t i o n of
A c c r e t i o n , R e s o r p t i o n and E x c h a n g e R e a c t i o n s i n the S k e l ­
e to n .
Kung l F y s i o g r a f s i k a S o l l s k a p e t s I Lu nd F o r H a n d l i n g a r
25: 1 -1 6 (1955).
B e r g e i m , O.
I n t e s t i n a l C h e m i s t r y V,
and P h o s p h o r u s A bsorption.
J.
( 1 9 2 6 ).
Bird,

Carbohydrate
Biol. C h e m .

and C a l c i u m
70: 35-45

H. R ., R. J. L i l l i e , and J. R. S i z e m o r e .
E n v i r o n m e n t and
S t i m u l a t i o n of C h i c k G r o w t h by A n t i b i o t i c s .
P o u l t r y Sci.
31: 907 (1952).

Brown,

E . W.
D i g e s t i o n E x p e r i m e n t s With P o u l t r y .
B u r e a u of A n i m a l I n d u s t r y B u l l e t i n 56 (1904).

U.S.D.A.

B r o w n e , T. G.
Some O b s e r v a t i o n s on the D i ge s ti ve S y s t e m of the
Fowl.
J. C o m p . P a t h . T h e r a p .
35: 12-32 (1922).
Card,

L. E . , M. H. K eith, and H. H. M i tc h e l l.
F a c t s O b ta in e d on
R a t e of D i g e s t i o n of F e e d by He ns.
I l l i n o is Station R e p o r t .
80 (1926).

C oates,

M. E . , C. D. D i c k i n s o n , G. F. H a r r i s o n , S. K. Kon, S. H.
C u m m i n s , a nd W. F . J. C u t h b e r t s o n .
Mode of A c tio n of
A n t i b i o t i c s in S t i m u l a t i n g G r o w t h of C h ic ks.
Nature.
168.
3 3 2 -3 3 3

C oates

(1951).

M. E .
T h e Mode of A c t i o n of A n t i b i o t i c s in A n i m a l N u t r i ­
tion,
C h e m . Ind.
1333-1335 (1953).

129

130
Com ar,

C. L . , I. B. vVhltney, a n d F. W. L e n g e m a n n .
Comparative
U t i l i z a t i o n of D i e t a r y S r ^ ^ and C a l c i u m by D e ve lo p in g Rat
F e t u s a nd G r o w i n g R a t .
P r o c . Soc. E x ptl. Biol. Med
882 3 2 - 2 3 6 (1955).

Gopp, D. H . , J. G. H a m i l t o n , D. C. J o n e s , D. M. T h o m p s o n , and
C. C r a m e r .
T b e E f f e c t of Age and Low P h o s p h o r u s R i c k e t s
on C a l c i f i c a t i o n a n d t h e D e p o s i t i o n of C e r t a i n R a d i o a c t iv e
M e t a l s i n Bone .
T r a n s , M ac y C o n f e r e n c e on M e ta b o li c
Interrelations.
3: 2 2 6 - 2 5 8 (1951).
C u m m i n s , A. J . , a n d T . P . A l m y .
S tu die s on the R e l a t i o n s h i p B e ­
t w e e n M o t i l i t y a n d A b s o r p t i o n in the H u m a n S m a ll I n t e s t i n e .
G astroenterology.
23: 179-190 (1953).
F e d e r o v s k i i , N. P .
C r o p and S t o m a c h D i g e s t i o n of Hen T u r k e y s .
S o v e tsk a ra Zootekhm ia.
1: 5 0 -5 8 (1951).
Fenton,

P. E.
G lucose

R e s p o n s e of the G a s t r o i n t e s t i n a l T r a c t to I n g e st e d
S o lu t io n s ,
A m . J. P h y s i o l . , 144: 609-619 (1945).

F i s c h e r , J. E . , a n d T. S. Sutton.
E f f e c t s of L a c t o s e on G a s t r o ­
i n t e s t i n a l M o t i l i ty .
J. D a i r y Sci.
32: 139-162 (1949).
F r e d e r i c k s o n , A ., J. H o n o u r , and D. H. Copp.
M e a s u r e m e n t of
I n i t i a l Bone C l e a r a n c e of Ca^^ f r o m Blood i n the Rat.
F ed.
Proc.
14: 49 (1955).
French,

R. B ., and G. R. Cowgill.
I m m a t u r i t y of the O r g a n i s m a s
a F a c t o r D e t e r m i n i n g the F a v o r a b l e Influence of L a c t o s e on
t h e U t i l i z a t i o n of C a l c i u m and P h o s p h o r u s .
J. N u t r .
14:
3 8 3- 3 9 9

(1937).

G a b u t e n , A. R . , a n d C. S. S h a ff n e r.
A Study of the P h y s i o l o g i c a l
M e c h a n i s m s A f f e c t i n g Sp ec if ic G r a v i t y of C h ic k e n Egg.
Poultry

Sci.

33: 4 7 - 5 3

(1954)

G o o d m a n , R. D., A. E . L e w i s , E . A. Schuck, an d M. A. G r e e n f i e l d .
G astrointestinal T ransit.
A m . J. P h y s i o l .
169: 236-241
(1952).

131
Gordon,

H. A.
Colloquium.
S t u d i e s on the G r o w t h E f f e c t of A n t i ­
bio tic s in G e r m - F r e e A n im a ls.
Lobund In stitu te , U n iv e rsity
of N o t r e D a m e , I n d i a n a
(1952).

G r e e n b e r g , D. M.
S t u d i e s i n M i n e r a l M e t a b o l i s m With the Aid of
A r t i f i c i a l R a d i o a c t i v e I s o t o p e s VIII.
T r a c e r E xperim ents
w i t h R a d i o a c t i v e C a l c i u m and S t r o n t i u m on the M e c h a n i s m
of V i t a m i n D A c t i o n in R a c h i t i c R a t s .
J. Biol. C h e m .
157:
9 9 - 1 0 4 (1945).
G r e e n b e r g , D. M . , and F . M. T r o e s c h e r .
Study w ith R a d i o a c t i v e
I s o t o p e s of E x c r e t i o n of C a l c i u m a nd S t r o n t i u m by way of
the bile.
P r o c . Soc. E x p t l . Biol. Med. 49: 4 88 -4 9 1 (1942).
H a m i l t o n , T . S., a n d H. H. M i t c h e l l .
The O c c u r r a n c e of L a c t a s e
in t h e A l i m e n t a r y T r a c t of t h e C h i c k e n .
J. A g r . R e s .
27:
6 0 5 - 6 0 8 (1924).
H a n s a r d , S. L . , C. L. C o m o r , and M. P . P l u m l e e . E f f e c t of
C a l c i u m S t a t u s , M a s s of C a l c i u m A d m i n i s t e r e d , and Age on
C a ^ 5 M e t a b o l i s m in the Ra t.
P r o c . Soc. E x p t l . Biol. Med.
78: 4 5 5 - 4 6 0 (1951).
Henry,

K. M ., A. J. M a c D o n a l d , a n d H. E. M ag ee .
O bservations
t h e F u n c t i o n of the A l i m e n t a r y in F o w l s .
J. E x p t l . Biol.
10:

H euser,

1 5 3 -171

on

(1932).

G. F .
T h e R a te of P a s s a g e of F e e d F r o m the
th e Hen.
P o u l t r y Sci.
24: 2 0 - 2 4 (1945).

C r op of

H e u s e r , G. F . , a n d L. C. N o r r i s .
Some R e s u l t s of F e e d i n g A n t i ­
b i o t i c s to C h i c k e n s .
P o u l t r y Sci.
31: 857-862 (1952).
H i l l e r m a n , J. P . , F. H. K r a t z e r , a n d W. O. Wilson.
Food P a s s a g e
T h r o u g h C h i c k e n s and T u r k e y s and S o m e R e g u la ti ng F a c t o r s .
Poultry
K e i th ,

Sci.

32:

332-335

(195o).

M. H., L. E. C a r d , and H. H. M i t c h e l l .
The R a te
of F o o d T h r o u g h the D i g e s t i v e T r a c t of the Hen.
Res.

34:

7 5 9 -7 70 (1927).

of P a s s a g e
J. Agr.

132
K e n n e d y , G. F . , J. P . M a r v e l , J . W a d de ll, H. W. T i t u s , and A. L.
Me lining, J r .
T h e A s s a y of V i t a m i n D U sing a S t r a i n of
C h i c k e n s S e l e c t e d f o r t h e i r U n i f o r m i t y of R e s p o n s e .
Poultry
Sci.
32: 2 9 - 3 8 (1953).
K l i n e , O. L ., J , A. K e e n a n , C. A. E l v e h j e m , a n d E . B. H a r t .
L a c t o s e in N u t r i t i o n .
J. B io l. C h e m .
98: 121-131 (1932).
L i n d b l a d , G. S., S. J. S l i n g e r , G. W. A n d e r s o n , and I. Motzok.
E f f e c t of A u r e o m y c i n on t h e C a l c i u m a n d P h o s p h o r u s R e ­
q u i r e m e n t s of C h i c k s .
P o u l t r y Sci.
31: 9 2 3 - 9 2 4 (1952).
M a c D o n a l d , N, S., R. E . N u s b a u m , F. E z m i r l i a n , R. C. B a r b e v a ,
G. V. A l e x a n d e r , P . Spain, a n d D. E . R o u nd s.
G astro­
i n t e s t i n a l A b s o r p t i o n of I ons I. A g e n t s D i m i n i s h i n g A b s o r p ­
t i o n of S t r o n t i u m .
J. P h a r m . E x p t l . The r a p .
104: 348-353
(1952).
M i g i c o v s k y , B. B.,
V i t a m i n D.
(1955).

a nd J, W. S. J a m i e s o n .
C a l c i u m A b s o r p t i o n and
C a n a d i a n J. B i o c h e m . P h y s i o l .
33: 202-211

M i g i c o v s k y , B. B ., A. M. N i e l s o n , M. Gluc k, and R. B u r g e s s .
P e n i c i l l i n and C a l c i u m A b s o r p t i o n .
A r c h . B i o c h e m . Biophy.
34: 4 7 9 - 4 8 0 (1951).
P e a r s o n , P a u l B.
M e t a b o l i s m of S t r o n t i u m , C e s i u m and Y t t r i u m by
A nim als.
P r e s e n t e d a t H a r w e l l C o n f e r e n c e (1955).
P echer,

C.
B i o l o g i c a l I n v e s t i g a t i o n s With R a d i o a c t i v e C a l c i u m and
Strontium .
P r o c . Soc. E x p tl . Biol. Med.
46: 86-91 (1941).

P l i m m e r , R. H. A.
On the P r e s e n c e of L a c t a s e in the I n t e s t i n e s
of A n i m a l s a n d on the A d a p t a t i o n of the I n te s ti n e to L a c t o s e .
J. P h y s io l.
35: 2 0 -31 (1906).
R o b e r t s , E . , a nd A. A. C h r i s t m a n .
The In fluence of L a c t o s e and
I t s H y d r o l y s e s P r o d u c t s on the A b s o r p ti o n of C a l c i u m .
J.
Biol.

Chem.

145: 2 67-271

(1942).

R o b i n s o n , C. S., an d C. W. Duncan.
The E ff e ct of L a c t o s e a n d the
A c i d - b a s e Value of the Diet on the H y d r o g e n Ion C o n c e n t r a t i o n
of t h e I n t e s t i n a l C o n t e n ts of the Rat and T h e i r P o s s i b l e

133
I n f l u e n c e on C a l c i u m A b s o r p t i o n .
447 (1931).
Roccuzzo,
Ross,

R utter,

M.

Bol.

Soc. I t a l .

Biol.

sper.

J.

Biol.

20:

Chem.

92: 4 35 -

5 (1945).

F . , and H. Y a c o w i t z .
E f f e c t of P e n i c i l l i n on G r o w t h and
Bone A s h of C h i c k s F e d D i f f e r e n t L e v e l s of V i t a m i n D and
Phosphorus.
P o u l t r y Sci.
33: 2 6 2 - 2 6 5 (1954).
W. J . , P . K r i c h e v s k y , H. M. Sc ott, and R. G. H a n s e n .
The
M e t a b o l i s m of L a c t o s e and G a l a c t o s e in the C h i c k .
Poultry
Sci.
32: 7 0 6 - 7 1 5 (1952).

S a x e n a , H. C., L. R. B e r g , an d J. M c G i n n i s .
F a c t o r s A f fe c ti n g the
G r o w t h R e s p o n s e of C h i c k s and T u r k e y P o u l t s to A n t i b i o t i c s .
P o u l t r y Sci.
31: 1 07 0 -1 0 7 4 (1952).
Shaw, T. P .
D i g e s t i o n in t h e C h i c k .
(1913).
Steinm etzer,

K.

Arch.

f. d.

ges.

Am.

Physiol.

S t u r k i e , P . D., a nd D. P o l i n .
T he E f f e c t s
E g g F o r m a t i o n and Blood CaLcium.
210 (1954).
Titus,

J. P h y s i o l .

206:

500

31: 4 39

(1924).

of I n j e c t e d P e n i c i l l i n on
P o u l t r y Sci,
33: 2 0 9 -

H. W., A. L . M e h r i n g , J r . , J. P . M a r v e l , G. H. Kennedy,
a n d J. W a dde ll.
T he R e s p o n s e of N ew H a m p s h i r e C h i c k s
t o a F i x e d A l l o w a n c e of V i t a m i n D.
P o u l t r y Sci.
32: 3953 (1953).

T w e e d y , W. F .
T h e E f f e c t of P a r a t h y r o i d E x t r a c t Upon the D i s t r i ­
b u ti o n , R e t e n t i o n and E x c r e t i o n of L a b e l e d S t r o n t i u m .
J.
Biol. C h e m .
161: 105-113 (1945).
U y e y a m a , K., J. E . G i o n s i r a c u s a , T. L . A l t h a u s e n , and H. A.
H arper.
A l t e r a t i o n of I n t e s t i n a l A b s o r p t i o n in U l c e r a t i v e
C o l i t i s and in In te stinal. H y p e r m o t i l i t y ; M o d i f ic a ti o n of T h e s e
A l t e r a t i o n s in U l c e r a t i v e C o l i t i s by T w e e n 80, A CT H and
Cortisone.
G astroenterology.
23: 143-154 (1953).

134
Waibel

P . E . , M. L . Sunde, and. W. W. C r a v e n s .
E f f e c t of A d d i t i o n
of P e n i c i l l i n t o t h e H e n ' s R a t i o n on B i o t i n and F o l i c A cid
C o n t e n t of E g g s .
P o u l t r y Sci.
31: 6 2 1 - 6 2 4 (1952).

W o l t e r i n k , L.

F.

Personal

C om m unication,