BIOLOGY

OF

REPROI)UCTION

FSH

13,

and

19 5-202

LH Release

During

Immature
SHUJI

SASAMOTO

and

Ovulation
with

TETSU

of Veterinary
of Agriculture

Fuchu,
Tokyo
of Physiology,
National
Chiba
280,

Department

First

Rats Pretreated

Laboratory
University

Tokyo

the

Period

of

PMS

JOHKE

Physiology
and Technology

183, and
Institute
Japan

of Animal

Industry

ABSTRACT
Plasma
and pituitary
FSH and
LH were measured
by nadioimmunoassay
at different
times
during
the first ovulation
period
(from
1100 h on day 28 to 0800
h on day 29) of immature
rats
pretreated
with
3 IU PMS on day 26. Gonadotropin
changes
were compared
with those of normal
4-day
cyclic
rats during
the peniovulatory
period
(from 1100 h on proestrus to 0800
h on estrus) to
elucidate the mechanism
responsible for the delay of the second estrus in these PMS-primed
rats.
A rapid
increase
of plasma
FSH and LH occurred
after
the critical
period
in the immature
rats
as well
as in the normal
cyclic
rats; the peak level of FSH was higher and that of LH lower in
immature
rats than in adults.
Plasma
FSH in the immature
rats decreased
after 2300 h on day 28,
contrary
to the adults
in which
elevated
plasma
FSH persisted
until
0500
h of the morning
of
estrus. The LH surge in the adult rats was transitory showing a different pattern from FSH. In the
immature
rats the LH surge lasted 3 h longer
than in the cyclic
rats.
Pituitary
FSH and LH contents
of cyclic
rats rapidly
decreased
after
the critical
period.
FSH
was replenished
during
the night
(from proestrus to estrus) despite persisting high plasma
FSH
levels.
In contrast
there
was only
a slight
restoration
of pituitary
LH during
this period.
On the
other
hand,
in immature
rats pretreated
with
PMS, pituitary
FSH and LH contents
revealed
the
same
pattern:
after
the critical
period
there
was a gradual
decrease
until
0200
h on day 29,
followed
by a slight
increase
in the morning.
These results indicate that the duration of the FSH surge during
the first ovulation
period
in
these PMS-pnimed
immature
rats may not sustain
the threshold
time necessary
for the initiation
of
follicular
maturation
of the new set of follicles
for the succeeding
estrous
cycle.
The regulatory
pattern
of pituitary
gonadotropin
and/or
the sensitivity
of the pituitary
to gonadotropin
releasing
factors
in the immature
rats have not yet differentiated
into the regulatory
pattern
in the adult.
INTRODUCTION
A single
in

injection

rats

immature

treatment.
tion

day

ministration
1963;

and

1963;

et

al.,

1971

nous

gonadotropin

maturation

PMS

has

1965;

Klawon

et

et al.,

1972).

maintenance

and

been

(Sasamoto
the

biological

determined

for
in

these
and
active

(Sasamoto

not

adult

1971;

rats

tant

Endoge-

1973).

role

follicles

follicular

has
in

(Schwartz,

animals

The

Kennan,
life

of

et

al.,

release

of
during

Received
Accepted

delay

August 28, 1974.

May 16, 1975.

rats.

195

rats.

of

Welschen,

The
the
the

to

patterns

the

first

the

FSH

ovulation
rats
of

estrus

in

imporset

succeeding

of
with
the

mechanism

second

an

of

cycle

1973).

the

purpose

during
cycle

play
of

the

to
and

estrous

is concerned

immature

elucidate

the

development

paper

due

surge

of

in

corpora

but

(Sawamoto

assumed

ovulate

1969;

PMS-treated

to

the

present

comparison

cyclic

been

which

of

gonadotropin

period

et

which

1969),

growth

periovulatory

Taya

anestrus,

Wiest,

first

the

1969;

activation

to

follicular

PMS-primed
after

Wiest,
were

and

of

these

9 days

animals

attributable

failure

in

and

(1-lashimoto

the

Wagner
al.,

was

the

Sasamoto,

and

1964;

1974);

the

1962,

estrus

delayed

was

(Hashimoto

al.,
lutea

ad-

Zarrow

Mahesh

clarified

1973)

Zarrow,

on

PMS

Meyer,

1962;

requirements

and
been

and

ovulasurge

second

however,

ovulation

after

that
LU

following

and

Brown-Grant,

1972,

indicate

Meyer,

Quinn

ovulation
morning

endogenous

afternoon

and

Ving

have

day

(McCormack

Strauss

Quinn,

an

produces

third

studies

on

second

PMS

on the

Recent

depends

the

of

The

1972).

rats,

the

and

LU

period
those

of

these

in
adult

investigation
responsible

in

with

was
for

PMS-primed

the

SASAMOTO

196

MATERIALS

AND

AND

METHODS

Twenty-four-day-old
(day
24) female
Wistan
rats,
weaned
on day 21, and adult female rats of the same
strain
were
obtained
from
the Laboratory
of Animal
Reproduction,
Omiya,
Japan.
The
animals
were
housed
in an air conditioned
room
(22
to 28 C)
illuminated
from
0500
to 1900 h and provided
food
and water
ad libitum
throughout
these
experiments.
Usually.
10 immature
rats or 5 adult rats were housed
pen steel cage (21 X 26 X 40 cm). Vaginal
opening
was
normally
observed
on 36.8 0.3 days of age.
Immature
rats were given 3 IU of PMS intravenously at 0900
h on day 26. In the adult
rats, vaginal
smears
were
examined
daily
and only
rats showing
3
on more consecutive
4-day
cycles were used.
Four
to 7 rats were
killed
by bleeding
via the
abdominal
aorta
using
a hepaninized
syringe
under
light ether anesthesia
every
3 h from
1100 hon
day 28
to 0800
h on day 29 in the PMS treated
immature
rats, or 1100
h on proestrus
to 0800
h on the day of
estrus
in adult
cyclic
rats.
Individual
plasma
samples
were
obtained
after
centnifugation
and stored
at -20
C until
assayed.
At
autopsy,
the pituitary
was removed
within
5 mm of
exsanguination
percent
the

and

NaCI.

After

supennatant

Plasma
Pituitary

in

5 ml

of

the

homogenate,

stored

at

was

until
assayed.
Wet
drainage
of luminal
were examined
for
Whitney
(1941).
measured

homogenized
centnifugation
obtained

and

of

cold

weights
of ovaries
and uteri
(after
fluid)
were obtained,
and oviducts
ova by the method
of Bundick
and

and

pituitary

using

the

levels
RIA

Distribution

kit

Program,

of

FSH

and

supplied

LH

by

were

the

NIAMDD,

TABLE
1. The time of ovulation
rats pretreated
with
PMS.

Time of
autopsy

Day

28

29

No.
Mean
PMS

of

the immature
rat was diluted
50-fold,
adult
was diluted
40-fold,
with
0.5
percent
bovine
serum
albumin
in 0.01
M phosphate
buffered
saline (0.5 percent
lISA).
These diluents
were
assayed
in duplicate
in 100 and 200 l aliquots.
The lowest measurable
value of LH RIA was 8 to
20 ng/tube.
Serum
samples
were assayed
in duplicate
in 100
and 200 j.tl aliquots.
The
supernatant
of the
pituitary
homogenate
of the immature
rat was diluted
50-fold,
and that
of the adult
was diluted
100-fold,
with
0.5 percent
BSA.
These
diluents
were assayed
in
100 and 200 ml aliquots.
Experimental
group
data were analyzed
for significant differences
by means
of Duncans
multiple
range
test (Steel
and Torrie,
1960). A value for P of less than
0.05 was taken as significant.
of

RESULTS

The

Time

of Ovulation

Neither

PMS-treated

ovulated

by 2300

Rat

NIH

IRat

and

changes

of ovarian

next

morning.

was

very

similar

The

occurred
adult

those

of

value

at

ence

1100

as

was

weight

changes

probably

and

5
5
5

4/5
5/5
5/5

29.8
31.8
28.3

26.

1 and

and

to

weight
SD)

1.2
.8
1.4
2.4
1.3
1.7
2.5
.5

ovarian

29.

weight

In

1700

compared

h-groups,

of

but

the

presence

of

day

28

h-group;
of

periovulatory

the

differ-

1400

the

a
with

nonsignificant
that

the

showed

during

(mg)

surge

of

tendency

during

rats

LH

day

general

weights

hr
hr
hr

hr on day

h on

ovulation

immature

2300

with

no

uterine

02.00
05.00
08.00

iv at 09.00

0200

the

h on

h showed

due

was given

at

compared

there

34.6

(3 lU)

surge;

of

elevation

ovarian

h-

2300

0/4

examined.

rats

LU

(Tables

when

0800

increase

rat.

by
time

the

of

This

rats,

significant

22.5
22.0
27.0
26.7

of animals

adult

of the

groups

h,

until

cyclic

Ovarian
(mean

per ovulating

both

3).

persisted

0/5
0/5
0/6
0/7

of ova

the

1700

(Table

weight

Ovulation

ovulating/No,

ovulated

weight

at

5
5
6
7

number

had
Thus,
in

ovarian

hr
hr
hr
hr
hr

of animals

nor

day

2).

No. of
animals

(7.8)
(8.2)
(8.8)

rats

h on the

(4/5)

most

the

14.00
17.00
20.00
23.00

11.00

the

increased

LH-l-2,
Rat LH-RP-1
(0.03 X NIH-LH-S1),
A-Rat
LHS-1:
Rat FSH-I-1, Rat FSH-RP-1
(2.1 X NIH-FSH51), A-Rat
FSI-IS-4l
RIA results were expressed
as ng
of NIAMD
Rat-RP-1/ml
plasma,
or g
of NIAMD
Rat-RP-1 /pituitary.
The lowest
measurable
value
of FSH
RIA
was 6
ng/tube.
Serum
samples
were
assayed
in duplicate
in
50 and 100 zl or in 100 and 200 iI aliquots
depending
on their
potency.
The
supernatant
of the pituitary

Day

homogenate
and that

whereas

0.9

-20

JOHKE

ovarian
period

several

sets

of

to 29 in immature

Uterine
(mean

107.6

100.8

113.3
112.7
126.3

113.4
103.6
98.0

weight
SE)
6.5
6.8
6.2
5.1
8.5

3.3
5.0

6.5

(mg)

PERIOVULATORY

TABLE
cyclic

2. The
rats.

time

of

ovulation

Time of
autopsy

11.00

Proestrus

14.00

17.00
20.00
23.00
Estrus

02.00
05.00

08.00

No.

lutea.

The

uterine

ly

increased

had

at

0/5
0/4
0/5
0/5
0/5

69.8
75.5
67.4
85.2
89.4

hr
hr
hr

5
5
5

4/5
5/5
5/5

of ova

of animals

per ovulating

(7.5)
(12.8)
(13.8)

2300

on

day
day

occurred.
those

rats

28,
29

gradual-

followed
when

These

observed

by

the

Plasma
day

28

FSH

cyclic

but

FSH Levels
During
of Immature
Rats
(Table
3, and Fig.

levels

showed

were

low

at

Uterine

(mg)

weight
SE)

to estrus

(mean

in adult

weight

368.4
380.5
353.4
380.6
364.6

5.2
2.1
7.1

334.6
314.8
290.2

(mg)

SE)

4.1
3.9
2.0
4.1
4.1

marked

on

15.4
16.8
20.0
23.4
12.0

7.5
10.4
4.0

day

on

and

observed

15

FSH
percent

gradually

h on

similar

at

content
greater

(Fig.

at

29.

At

were

the

1400

than

h on

Thereafter,

FSH

to 0200

latter

higher

(nonsignificant)

1).

decreased

day

values

levels were still 3 times

at 1100 h on day 28.

FSH
amount

1100

at

28)

time,

was

increase

was

at 0800

Pituitary

I)
h

increase

maintained
the

the

1100

a nonsignificant

h.

2300

were

adult

rats.

Plasma
and Pituitary
Periovulatory
Period
Pretreated
with
PMS

proestrus

1700 h and plateau

levels were maintained
until
2300
h on
day
28. Thereafter,
FSU
levels
declined
until
0500
h (P<0.05
vs the value
at

ovulation

changes

in

during

rat.

immature

h on
h

197

examined.

1400
of

76.8
77.4
81.2

RATS

weights

5
4
5
5
5

0800

to

uterine

hr
hr
hr
hr
hr

weight

already

similar

and

IMMATURE

Ovarian
(mean

ovulating/No,

to

decrease

of ovarian

LH:

Ovulation

number

corpora

changes

AND

No. of
animals

of animals

Mean

and

FSH

h on

day

28

than

at

content

day

29 when
Cs)

()

IMMATURE

RATS
ON

PETREATED

DAY

WITH

PMS

ADULT

CYCUC

RATS

26

H
Avug.

14

MY

FIG.
standard

1. Changes
errors

of the

of

FSH

ot

11.00

at

Iv

00%

20

23

02

06

061v

II

14

25

of

LH

I?

I
II

QmoIt

DAY

pituitary

means.

FSH-

and

PROOSTRUS

29

LH-contents

17

during

the

periovulatory

20

I
period.

02

06

OS

ESTRUS

Vertical

bars

indicate

198

SASAMOTO

the

lowest

value

vs the

P<0.05
was

followed

value

that

and

by

still

the

before

observed

at

an

was

was

was

value

72

1400

increase

by

percent

of

significantly

critical

(59

h on

less

0800

28).

This

h to

FSU

1100

than

the

content

Fig.

Plasma

of

162

than

350

persisted
The

FSH

1400

marked

o
u-

,,,

&

t-

*O
r.-ox

C(M
Or-!-.
-

1)

h,
the

(nonsignificant),

decrease

at

until

hand

this

the

next

of

still

3 times

>,-3c

#{176}
*1

+1

*1

r-Z

higher

o +1

+1

os

r-.i

esi

,t

Os 00

10

percent

followed
h to

00

58

*1

*1

Os

00

c!

by

only

1!

00

period.

increased

1700

--

significantly

critical

content

low

1700

decreased

0800

before

Pituitary
at

at
0500

levels

at

value

to a maximum

ng/ml
until

FSH

ng/ml

the

were

increased

approximately

morning.

which

levels,

proestrus,

elevation
to

FSH Levels
During
the
of Adult
Cyclic
Rats

00

1)

FSH

h on

1400

period.

Pituitary
Period

(Table4,and

h levels

Plasma
and
Periovulatory

JOHKE

percent;

day

the

AND

0a
>

percent
C

of

the

1100

levels
of

h values

persisted

until

pituitary

night

FSH

and

there
FSU

by

was
which

1).

2000

Although

h,

contents

0500

compared

(Fig.

a gradual

h on

the

the

initial

during

morning

the

of
of

amount

increase

occurred

90
percent
recovery
was
a nonsignificant

to

similar

estrus,

pituitary
difference

at

1100

r-

.9

r-

hon

LH Levels
of Immature

During
Rats

N*e-I

+1

*:

..

Pituitary
Period

dr-

.
...

proestrus.

Plasma
and
Periovulatory

.9

the

._.

*00NX

(I

,,

u
C

Pretreated
Plasma
h

on

day

than

40,

less

than

at

with

PMS

LH

values

28

were

andat
40

1100

(Table

h or

47,

per

40,

and

at 1100

one

50,41

plasma.

h was

rats

40

60,

ml

1400

1)

of individual

1400h,
ng

3, Fig.

of

mean

value

only

from

calculated

6.

less

andtwoof

The

>,

.#{149}
*

#{176}.

*
*

O00rs1

these

samples.

low

at

1100

marked

and

increase

by

a slightly

LH

further

2300

Plasma

LU

1400
at

levels,

h on

1700

h,

day

level

of

decreased

to

a mean

day

28,

28,

400

ng/ml.

though

>5

3 h later

of

NO-

were

showed

followed

lower

on

which

0C

C
0

Plasma

189

ng/ml

at

was

no

there

C
0

Os

significant

at

difference

2000

compared

h, probably

due

with

to the

the

wide

values

range

of

..

variation.

At

morning,

plasma

mean

of

slightly

0500

or

0800

LU

further

approximately
higher

than

h of

the

the

following

decreased

.2

to

70

ng/ml,

which

level

before

the

a
was

critical

,.

.0

.C

.C

00000

period.
Changes
in Fig.
value

.C

.C

.C

.0

1,
at

pituitary

expressed
1100
h

LH

contents

as percentages
on
day
28.

are

shown

of the
Pituitary

#{149}j

,,.,

mean
LH

00

only

a nonsignificant

decrease

at

1700

C-

.0
0
C

000

of

C- a

a
V

00

showed

.0

PERIOVULATORY

FSH

AND

IMMATURE

LH:

h,

though

there

plasma

LI-I

occurred
2
+1

ac

rsl

ri

t-

*1

*1

*1

so

+1

E
C
0

0.

E
.

r-!

*
,s

!-4

50
Os

00

o-

so

+1

*1

the

h on

*1

so

1400

C-!
ri

r-!

V,

(s
O5

contents

day

29,

with

the

was

at

1700

until

day

was

at

1100

h on day

Pituitary

LH

in

decrease

h on

0200

Afterwards
79 percent
nonsignificant

to

which

increase

further

(60
percent
(P<0.05
vs

28).

recovered

and

marked

h.

lowest
value
was
observed

level

Plasma

>5

199

gradually

when
values)

+1

RATS

day

of

29,

1100
value

the

h
at

putuitary
LU
at 0800
h on
compared
28.

Levels

During

the

Periovulatory

-oa

(Table

._

rslr!N

2
3

2 E
.

+1

r-

rl-.s
r-

critical

Cse-ir-0

to

._.

-0

r-

4,and

of Adult

Fig.
levels,

Plasma

Period

period

a peak

1)
which

(123

of

Cyclic

were

ng/ml),

1144

Rats

low

before

strikingly

ng/ml

at

the

increased

h and

1700

this

was

00-N

r-l

followed

by

2300

level

the

a marked
LU

found

Pituitary

significantly

before
LH

at 2000

decrease

plasma

the critical

contents

h.

By
to

decreased

period.

showed

a nonsignifi-

be

.2

cant

decrease

-5
.9C

*1

Os

*1

55
*1

+1

*(

*1

,s
u
C00*N
o-s V

so

Crsr!
C-i *

50

'5

'5

*1

._.

(41

until

1100

the

65

a marked

percent

of

significant

the

h on

lowest
content
was observed.

morning

of

estrus,

was still
initial
level

but

with

period

1100
decrease

significantly

compared

critical

by

h, when the
h values)

increased

58 percent

before

2300

content

only

to
further

to 0800

0200

LU

of

h, followed

1).

percent

From
*1

1400

1700

(Fig.

occurred

Os

at

at

values

C!

!l)

to
2a0!J

increase

the

(P<0.05).

.2
C
u
V
C
0

DISCUSSION

the
to

3ct

In

the

present

adult

the

the FSH
surge
in
from
late proestrus

experiments,

cyclic

rat

persists

of

morning

estrus

which

agrees

the

with

*NN
-

>5

pattern
observed
by
and
Parlow
(1971),

(1972),

+t

0..

Gay

et

and

Daane

al. (1970),
Taya
and

Igarashi

L.

00V

.-

0
C

E
-

but

is contrary

to

the

results

during
the
rats pretreat-

ed

have

with

PMS,

plasma

half

FSU

as much

levels

at 0200

to

compared
although

to the levels
at 2300
h on day 28,
peak
levels
of FSU
are higher
than

those

in

of

the

5-

.0

5..
.

00000

o#{176}

000
C

---C-sIr-I

000

by

These

changes

rats

Mahesh

of

are
et

similar
al.
(1972),

h on day

plasma
to

those
who

30-day-old
rats after
PMS
administration;
their study
a significant
increase
in plasma
an dLUhadalreadyoccurredatl400honday

29

FSH
obused

in
FSU

32.
In

rats.

immature

served

>5

already

decreased

08

observed

by Ayalon
et al. (1972).
However,
first ovulation
period
in immature

rats,
the
plasma

LU

respectively
Therefore

in

half-life
of
were
reported

endogenous
as 2.5

FSH
and
h and 0.5 h,

C-

C-

0.

it

(Bogdanove
is assumed

and
that

Gay,
1969).
pituitaries
of

SASAMOTO

200

immature

rats, pretreated

lower

steady-state

by

0200

h on

period;

the

whereas

h of

the

morning

FSH

influence

of

an

by
androgen

tion

during

pituitaries

of

been

rat

the

and

the

ovulation

to

the

rate

of

secreted

-,

has

The

the

in adult

be

2000

present

study,

cyclic

rats,

h on

pituitary

FSU

increased

ceeds

proestrus

to

0800

h on

the

the

In

immature

second

is

due

opment.

By

at

1800

immature

ovulation
These

to

the

the

the

next

rats

maturation

of the

have

reported

during

rats

may

be

rats.

The

amount

for

follicles

et
Plasma

a transient

initial

of
rat

of
The

adult

cyclic

re-

(PMS)
of

for

in an

the

ovulable

peak

levels
after

in

adult
the

cyclic

critical

29
in

still

in

is

ovulation,
h in

completion

the

of

Raj

FSU

LU

be

circuis

ovulation

and

Moudgal,

and

(1973)

and

observed

from

Parlow

to

et

al.

and

(1974).

of

maximum

by

Igarashi

Aiyer

FSH

secretion

administration

release
different

as suggested

Taya

pituitary

LH-RH

FSH
by

LU,

(1971)

Aiyer
that

determinations

that

controlled

mechanism

Daane

and

indicate

may

In

et

al.

in

re-

is highest

proestrus

as

well

at

as

LU.

to LH, the plasma

level of
at 0500
h of estrus,
11 h

FSU

immature
surge

LH

it
in

lasted
the

decrease

response

to

longer

adult

of

LU-RU

until

pituitary

rats,

show

a different

pattern

sensitivity
into

LU

0200

which

peak

contents

on

day

observed.

the

factors

may

adult

type.

FSH

29,
These

release

pattern
with

pituitary
of

the

in

pretreated

of

a lower

contrast

cyclic

the

the

period

are similar
to those
of
with the results

LU

of adult

tory

with

is

PMS,

critical

Pituitary

content

is in striking

rats

the

but

rats.

lowest

This

with

after

synthesis
immature

tiated

greater

pretreated

gradually
the

changes
FSH.

rats
occurred

pin-releasing

secondary

PMS,

FSH
pituitary
not

from

and
to

yet

and
LH.

the

regula-

LU

and/or

In

gonadotro-

have

differen-

maintenance
state

(Sasa-

1972).

LH

synthesis

when

immature

is 10 times

required

follicles

the

Tomacari

gonadotropin
development

and

than

levels

day
when,

(1974).

of

period.

follicles

rats

though

in these
to

of

cyclic

the

female

2),

gonadotropin

Madhwa

after

for

slight

morning

h on

circulating

first

secretion

compared
of

amount

al.,

continued

results

thresh-

the

periovulatory

mature

mature

in

1969;

adult

follicu-

and

the

maximal

needed

for

These

of

of

increased

activities

lower

the

into
the

the

the

secreting

quired

that

proestrus

for

androgen

Gay

the

1970).

the

set of follicles

new

by

surge

induce

no

However,
in contrast
FSU reached
a peak

of

the

during

though

FSI-I

is

con-

period

sustain

to

such

duration

initiation

cycle.

during

responsible

FSH

the

order

thereafter

sponse

1973).

proes-

that

low

estrus,

the

in

the

ovulation
not

for

estrous

testosterone

the

first

may

necessary

after

be

1 and

1700-1800

Sasamoto,

h on

suggests

0200

PMS

with

2000

(Tables

of

period

days

that

first
devel-

ovulation
4

the

follicular

critical

and

suggest

immature
time

of

the

the

injection

after

during

PMS,

after

critical

recovery).

hormone

regulatory

during

with

days

failure

just

results

(1974)

moto

pretreated

delayed

rats,

succeeding

of

secretion

a supplementary

surge

old

than

FSU

estrus
occurs
(Sawamoto

comitant

is

active

rats

estrus

ovulation

lar

of

rate

by

estrus

and

the

steady-state

around

rats,

(Sasamoto,

morning

after

occurs

cyclic

blood

short

occurs

percent

In

that

a rather

may

to

pattern

show

content

contents

of

required

from

period.

these

adult,

lating

contents

steadily

the

day

progress.

cyclic

of estrus
(90
percent
recovery),
contrasting
with
the slow and insufficient
recovery
of pituitary
FSH
in immature
rats. The rate of synthesis of pituitary
FSI-I
in the adult
cyclic
rat ex-

FSH

(58

estrus

ovulating

adult

this

of

the

LH

Naft-

suggest

a lower

release

and

1970;

results

release
reach

proestrus
of

or

LU
they

LH

Ovulation

by the
may

of

rats

synthesis

increase

FSU

adult

Taya

al.,

a different

These

pituitary
LU

night

1971;
et

showing

active

for

by

Parlow,

FSH.

that

condition
The

secre-

rats

of

trus.

ovaries

1972),
of

and

the

et al

and

Brown-Grant

of the

in

period

immature

FSU

al.,
that

period

the

androgen

et al. (1974).

these

et

pituitaries

suggested

secreted

adrenals

first

Daane
1973;

from

0500

(Bogdanove

cyclic

the

1970;

olin

FSU

survive

to

FSH

Falvo

from

In
of

under

al.,

duration

to

the
by

secreted

different
rats.

pituitary
has

adult

both

shown

until

JOHKE

lgarashi,

critical

rat

activated

pituitary

the

from

been

the

cyclic

capacity

androgen-deprived
In

be

the

relative

1974).

adult

FSU

of estrus.

enhanced

circulation

fall into

secreting

after

the
to

secreted

have

PMS,
for

morning

in

is assumed

secretion

with

condition

AND

rats
period

revealed
(Gay

et

ACKNOWLEDGMENTS
The
authors
thank
the
Rat
Pituitary
Hormone
Distribution
Program,
NIAMDD,
NIH,
for supplies
of
RIA
kits
for
FSH
and LH.
PMS (1030
lU/mg)
was

PERIOVULATORY

kindly

provided

by

the

Laboratory

of

FSH

Sankyo

Zoki

Co.

(Tokyo).
This
work
was supported
in part by a grant-in-aid
for Scientific
Research,
Ministry
of Education,
Japan
(1973
and 1974).

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the

S., Fink,
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