BIOLOGY
OF
REPRODUCTION
Sertoli
Cell
31,
565-577
Development
C.J.A.U.V.
VAN
(1984)
of Pig Testis
in the
VORSTENBOSCH,’
E. SPEK,
and
C.J.G.
of
of
Period
B. COLENBRANDER
Anatomy
Veterinary
State
Utrecht,
and Neonatal
WENSING
Department
School
Fetal
University
The
Medicine
Utrecht
Netherlands
ABSTRACT
The
Sertoli
cells
of pig fetuses
from
35 days postcoitum
until
1 mo after
birth
have been
investigated
by light and electron
microscopy
in decapitated
animals
and their control
littermates,
as well as in untreated
animals.
Until
52 days postcoitum,
Sertoli
cells change
in form
during
the
formation
of sex cords
but from
then onwards
they are rather
uniform.
They primarily
display
an
elongated
nonindented
nucleus
with one or more
prominent
nucleoli,
a well-developed
Golgi
apparatus,
and in the basal compartment
below
or beside
the nucleus,
a large lipid
droplet.
There
are large quantities
of rough endoplasmic
reticulum
(RER)
from
52 days postcoitum
onwards,
often
with
complex
whirl
forms
and a parallel
arrangement
of profiles
with
relatively
few ribosomes.
After
birth
their
numbers
seem
to be somewhat
less, and by 1 mo after birth
the RER profiles
are
often
shorter
and almost
free of ribosomes.
Clustered
ribosomes
are found
in large quantities
throughout the period
under
inyestigation.
Especially
in the early fetal period,
the endoplasmic
reticulum
(ER)
profiles
show prominently
filled cisternae.
Mitochondria
are mostly
long and slender,
or small
and ovoid.
Most
have lamellar
cristae,
but mixed
lamellar-tubular
cristae
can also be seen. Between
decapitated,
control
and untreated
animals
no obvious
ultrastructural
differences
could
be observed.
The peritubular
cell sheath
surrounding
the sex cords
did not show
signs of differentiation
into a
layer of myoid
cells.
Morphological
cell are
INTRODUCTION
In the pig
the gonad
becomes
as a testis
at
recognizable
days
postcoitum
dig cells
(Pelliniemi
(p.c.)
cannot
et
already
(Stewart
concentration
and
(Pelliniemi,
be identified
al., 1979),
capable
of
producing
Raeside,
peaks
at
of
ogenesis,
Leydig
match
cells,
the
lasting
until
is pituitary
a second
wave
which
This
days
period
begins
and
lasts
until
latter
p.c.
testosterone
(FSH)
in
data:
after
birth
(term
is 114 days
tenbosch
et al., 1982,
1984).
60
and
p.c.)
(van
studied
The fetal
of
duct.
days
birth,
but
present
at
70
1 mo
on
intermediate
Received
with
the
Vors-
time
1981).
the
‘Reprint
Vakgroep
TD Utrecht,
May
29,
1984.
6, 1983.
requests:
C.J.A.H.V.
Funktionele
Morfologie,
The Netherlands.
September
brander
van Vorstenbosch,
Valelaan
1, 3508
exception
Sertoli
p.c.
hardly
produce
at
hormone
in male
sexual
regression
of
and
90
high
cells
are
estrogens
little
acquire
high
dethe
levels
days
not
p.c.,
(Iran
et
synthesize
protein
capable
is
are
afterwards.
are
Data
available,
at
which
al., 1977,
proteins
(ABP)
and
of converting
(Dorrington
is known
about
these
functions
et
al.,
at what
(Colen-
1982).
In the pig, fetal hypophysectomy
tation
does
not seem
to effect
565
has
cells
decline
of
is also
Sertoli
into
et al.,
hormone
cells
period
such
as androgen-binding
inhibin.
In vitro
they
Accepted
(sustenta-
In the
pig AMH
is produced
p.c.,
a high
concentration
days
1978).
However,
age Sertoli
cells
con-
number
of pri197 5a,b).
The
anti-Mullerian
33
testosterone
pig
cords
cells
these
which
is important
because
it causes
level
Mature
sex
a follicle-stimulating
protein,
by
the
developing
early
Sertoli
at all.
(precursor)
one
reached
dependent.
at approximately
approximately
of
Mullerian
from
27
a devel-
approximately
independent,
precursor
dependency
(AMU),
velopment
steroid-
the
The
and a relatively
small
gonocytes
(Pelliniemi,
development
least
on
scarce.
of
cells)
mordial
(Ford
et al., 1980).
the changes
in mor-
is pituitary
mainly
cular
been
biochemical
wave
which
sist
30 days p.c.
the testis
is
important
opmental
days
p.c.
26
Ley-
Testosterone
37 days
1976).
approximately
data
Sertoli
1975a).
before
although
p.c. and declines
afterwards
From
35 days p.c. onwards
phology
ultrastructurally
approximately
Sertoli
by decapicell de-
VAN
566
velopment,
level,
at least
during
pituitary
trast
to
not
the
activity
Leydig
later
cell
at the
light
microscopic
fetal
period
when
is high.
This
development
tary
dependent
Vorstenbosch
(Colenbrander
et al., 1982,
studies
of sex
cord
shown
decrease
1979).
a significant
in cord
The
latter
sectomized
VORSTENBOSCH
Materials
et al., 1979;
1984).
Quantitative
however,
to
monkeys
et
al.,
The
present
study
light
electron
microscopic
development.
Fetal
were
carried
out to
izing
hormone
this
p.c.
descent.
up
to
and
provide
qualitative
and
FSH
dependency
The period
choosen
postpartum)
covers
the
second
developmental
velopment,
which
organelles
set
data
on pig Sertoli
cell
decapitation
experiments
determine
a possible
lutein-
(LH)
development.
until
1 mo
cular
was
microscopic
Attention
involved
has
in
been
protein
and
of
(30 days
first and
waves
of Leydig
includes
the period
cell deof testi-
focused
on
steroid
syn-
thesis.
AND
12 nm.
METHODS
Testes of fetuses
of Yorkshire
and Dutch Landrace
crossbred
pigs were studied.
At 42 days p.c. a number
of fetuses
was decapitated
according
to the method
of
Stryker
and
Dziuk
(1975):
littermates
served
as
controls.
Moreover,
a number
of fetuses
of normal
sows
(untreated
animals)
were
used
to check
the
reliability
of the control
fetuses,
since the handling
of
the uterus
for the decapitation
procedure might have
affected
the
normal
development
of these controls.
The number
of animals
used, sample
data and the
number
of litters
are given in Table
1. The
period
investigated
ranged
from
35 days
p.c.
to 30 days
postpartum
(pp.).
TABLE
design: the
1. Experimental
number
of animals
RESULTS
Light
Microscopy
At
35 days postcoitum
the sex cords
formed
an anastomosing
network
consisting
of cell
plates
and cord-like
structures
of
2-3
cell
layers
thickness.
The
primitive
sex cords
were
surrounded
in
the
observed
Normal
Litter
a
One
bTerm
day
animals
variation
at 113
days
in the
in each
at 42 days
dLitter
of the neonatal
origin
Post coitum
(p.c.)
in days
large
interstitium
After
80
90
100
Term
5
10
5
4
5
5
2
6
3
6
9
2
5
10
5
6
3
6
4
4
6
4
5
7
7
12
3
3
sampling
data;
p.c.
piglets
unknown.
but
were
( ±) 1 daya
75
the age of the animals
cells,
cells
group.
62
3
Leydig
these
52
p.c.
cDecapitation
numerous
35
animalsC
(untreated)
by
peripheral
Age
Decapitated
Controls
as reported
cleared
and
MATERIALS
processed
aperture.
On
semithin
sections
of true
cross-sectioned
seminiferous
tubules,
diameters
were measured
and
the number
of nuclei of both germ cells and Sertoli
cells were counted
in normal,
control
and decapitated
animals at the ages shown in Table 1. Results were analyzed with the one-way
analysis
of variance
(ANOVA
Fl) at a level of significance:
P<0.05.
The correlation
coefficient
was calculated
between
the number
of nuclei and the surface
of the tubular
cross sections.
For
the calculation
of the surfaces
it was assumed
that the
true cross sections
were
circles.
Reference
standards
of negative-stained
crystals
of
catalase
showing the 8.12 nm periodicity
(Polysciences,
Warrington,
PA) were used
to estimate
the diameters
of cytoplasmic
filaments
and to classify
the microfilaments
if their
diameter
was less than 8 nm and as
intermediate
filaments
if their diameter
was between
8
1977).
quantitative
and
dehydration
were
have
(Gulyas
handled
in graded
series
of acetone.
Materials
in propyleneoxide
and
embedded
in
either
DER I-mixture
(Lockwood,
1964)
or in Durcupan ACM
(Fluka
Chemical
Co., Hauppauge,
NY). Semithin sections
were
stained
with either
paragon
or toluidine
blue. Thin sections
were stained
with
lead citrate
(Venable
and Coggeshall,
1965).
A Philips
201G
microscope
was used at 60 kV with a 30-tm
objective
van
increase
in diameter
and a
length
(Colenbrander
et a!.,
is also observed
in hypophy-
rhesus
were
elsewhere
(van Vorstenbosch
et al., 1982). In brief: a
double
fixation
in 2.5% glutaraldehyde
followed
by
1% 0s04
according
to Sabatini
et al. (1963)
was
succeeded
by block staining
in 2% uranyl
acetate
prior
the
is in sharp
conwhich
is pitui-
development,
ET AL.
was precisely
known.
10
birthb
20
30
5d
SERTOLI
FIG.
plate-like
FIG.
1. Thirty-five
network
days
(asterisk).
postcoitum
CELL
DEVELOPMENT
(p.c.)-untreated
animal.
IN THE
The
early
567
PIG
sex
cords
formed
an
anastomosing
X 500.
2. Fifty-two
days
p.c-control
animal.
Sex
cords
were
clearly
of the testis. Numerous
germ cells could be observed. X 200.
FIG.
3. Fifty-two
days p.c-control
animal.
Sex cords varied
in diameter.
could be observed. X 500.
formed
but
still
anastomosed
at
(single,
pairs
clusters)
periphery
Germ
cells
and
the
568
VAN
scarce.
In the sex
were
precursor-Sertoli
cords
and
only
gonocytes
(Fig.
1).
At
cords
cords
a few
52
days
was
and
changed.
plates
straight
periphery
number
membrane
were
now,
by
and sometimes
be observed
could
be
the
of the
and
these
period.
and
cord
organization
p.c.
onwards
cords
was stable
period.
However,
control
of
of
Sertoli
the sex
animals
differ
The
sex
the
cells
cords
increased
animals
at
from
those
could
Leydig
birth
the
62 days
in controls
the
controls
cord
p.c.
decapitated
organization
rapidly
there
of
the
in
differed
and normal
in
control
and gonocytes.
of both
normal
diameter
difference
and
from
until
the end of the
there
was a shift
and 52 days p.c.; thereafter
any increase
until
the end
cells
in
days
not
35
Pairs,
of gonocytes
number
of
62
in the number
The
diameter
the
testis
(Fig.
2). A
with
the
spermatogonia.
small groups
(Fig.
3). The
sex
did
animals
of the sex
observation
anastomosing
into
single
towards
existed
in contact
cords
definition,
Sex
decapitated
animals.
From
observed
organization
but
still
were
of
decreased.
.uI
p.c.
the
ET AL.
majority
of the cells
(sustentacular
cells)
Centrally,
the
were
transformed
solid
cords
anastomoses
of gonocytes
basal
cells
p.c.
the
cells
VORSTENBOSCH
between
was hardly
observation
decapitated
significantly
animals.
diameter
animals
At
between
was
(59.7
pm
I
FIG. 4. Term-control
X 500.
FIG.
5. Term-decapitated
almost
disappeared
and
animal.
Except
animal.
the
interstitium
for
The
was
its diameter
sex cord
intruded
the
diameter
by
sex
was
erythrocytes.
cord
larger,
at term
the
X 500.
did
same
not
differ
from
as in controls.
that
Leydig
at 52 days
cells
had
SERTOLI
diameter
seminiferous
CELL
DEVELOPMENT
tubule
IN THE
Electron
).im
35
days
(hereafter
p.c.
a
axis
ficial
30
50
70
90
f
hO
20
FIG.
6. Changes
values
with
standard
mals
(open
circles),
symbols
in sex
cord
days
diameters.
The
average
deviations
are given:
control
normal,
untreated
animals
circles),
and decapitated
are used in Figs.
animals
7 and
(asterisks).
ani(solid
The
were
mostly
and
other
signs
same
8.
ments
could
to
basal
the
cells.
However,
Sertoli
cell.
made
be
observed
membrane
They
reliable
pm
15%
a
approximately
± 7.5,
respectively)
(Figs.
increase
is
which
4,
5 and
6).
a few
gonocytes
per
be
observed
at
35 days
increased
until
52 days
onwards
it
normal
that
as
the
was
rather
well
as
cells
were
normal
of
number
significantly
days
higher
p.c.
The
also
to birth
number
of
increased
pace.
After
cross
section
birth
cells
and
11).
animals
at
those
in
in
Sertoli
cells
birth
controls
section
in
on
number
cross
at
section
differed
and
cells
The
number
in
decapitated
normal
Sertoli
in every
throughout
very
the
tight
packing
impossible.
than
They
intermediate
to originate from
electroncell membrane
(Figs.
10
part of the cells a crisscross
the
intermediate
of apbe observed.
filaments
10 nm
diameter
could
Sertoli
cells
intermediate
showed
filaments
large
quantities
of
(Fig.
12). Some-
they
a close
spatial
showed
the
nuclear
membranes.
nucleus
showed
chondrial
relationship
with
or
membrane
the
mito-
outer
-
form
indentations.
Most
nucleoli
(Fig.
13).
number
of
germ
mostly
a
slightly
deep
cells had one or two
large
The Golgi
apparatus
was
well
but
displayed
seldom
cross-
per
tubule
14
12
per
of
from
animal
sectioned
cells
52
increased
a slower
significantly
in the
adjacent
most
80
section
35
of Sertoli
slightly.
and
cross
their
number
apparently
of
and
was
from
between
but
the
per
times
The
the
animals
scarce.
appear
seen
smaller
In the basal
irregular
birth
controls
Some
these
not
The
seemed
proximately
until
a sudden
7).
dropped
per
was
before
cross
substantially
p.c.;
thereafter
with
time,
Sertoli
than
(Fig.
days
linearly
(Fig.
10
8
6
4
2
8).
Between
the
and
Sertoli
cytes
decapitated
one
and
plaques
dense
with
for
animals
per
in
7 shows
identical
decapitated
gonocytes
then
birth
until
p.c.
After
birth
there
to a level
half
that
from
animals.
In decapitated
drop
number
from
Figure
days
80
could
Their
and
constant
germ
and
normal-control
section
p.c.
p.c.
in controls.
of
number
cross
super-
Coated
were
within
were
be
arrangement
Only
only
parallel
measurements
to
filaments
73.9
and
The
Cell-to-
be observed.
did
they
period.
appeared
and
width.
of pinocytosis
cell
observation
± 4.9
Their
directed.
loose,
could
Sertoli
form.
a variable
a
Between
Sertoli
cells,
and
also
between
Sertoli
cells
and
gonocytes,
gap junction-like
cell
connections
were
scarce
and
of variable
dimension.
Bundles
of tightly
packed
microfila-
40
birth
had
The
conical
radially
space
interdigitations
pits
0
was
upon
sometimes
layers.
or
cells
rested
which
several
polygonal
intercellular
cell contacts
20
of
Sertoli
cells)
lamina
to consist
cells had
longitudinal
precursor
Sertoli
basal
seemed
40
the
termed
well-developed
60
569
Microscopy
At
80
PIG
animals
hand,
sections
coefficient
of
cells
at
the
of
the
numbers
normal,
various
of gonocontrol
ages
30
50
70
90
f
110
20
40
and
on
the
the
surface
of the sex cord cross
the other,
there
was a correlative
of 0.96
(Fig. 9).
and
on
mean
birth
FIG.
cross-sectioned
animals.
7. Changes
cord
in
the
in normal,
number
control
of
days
gonocytes
and
decapitated
per
VAN
570
number
of
sertoli
cells
cross-sectioned
VORSTENBOSCH
per
ET AL.
bodies
tubule
to
35
could
the
be observed,
lysosomal
lipid
most
family.
droplets
per
cell
likely
One
were
or
belonging
a
few
present
in
large
its
basal
compartment.
30
At
Sertoli
25
not
20
52 days
cells
at
as profound
as one
rearrangements
15
10
surrounded
Sertoli
cells
30
50
70
90
f
110
20
(Fig.
40
birth
days
cells,
cytes,
in the number
of Sertoli
cells per
in normal,
control
and decapitated
FIG.
8. Changes
cross-sectioned
cord
animals.
and
tyosomes.
present
consisted
Coated
and
varying
quantities
in
Golgi
area
whole
observation
cilium
a
much
single
wider
space.
poorly
mon
finding
the
between
lamellar
the
mitochondria
form;
The
spatial
there
seen:
were
then
very
relationship
filled
the
only
strongly
but otherwise
to those
at
still
a regular
the
apparatus
9+0
coated
a
were
the
free
Sometimes
and
a
a
the
filled
(Figs.
this
Sertoli
cells
vesicles
appeared
more
no
elon-
clear
Large
differnucleoli
and
The
vesicles
and
Sertoli
joining
developed.
the
Golgi
quantity
was
of
considerable
to cell. Mitochondria
and slender
forms,
mitochondria
at
a com-
membranes
were
age
mainly
also
displayed
frehow-
common.
a lamel-
close
total
number
seminiferous
of cells/diameter
tubules
peculiar
membranes
mem-
with
finely
15
profiles
of
very
and
ER
16).
disIf
present,
an intimate
Sometimes
large,
and
each
showing
of
such
mitochondrion
(Fig.
may have originated
single
ER-mitochondrial
from
of
the
electron-dense,
cristae
were
cell
form
ribosomes
ER
was
formed
5
10
stacks
of
surface
seminiferous
FIG.
lysosome-like
ovoid
bodies,
membrane-bound
15
20
not
profiles.
few
ovoid
The
Interdigitations
pits
displayed
35 days
p.c.
observation
cell
long
gono-
Pinocytotic
uncoated
but varied
from
quently
showed
Ser-
and
with
ribosomes,
was smooth.
The
a
of
ER
a
and
projections
between
cells
basal
nuclei
well
and
mostly
(ER)
often
on
became
complexes
process
an
showed
very
a few
cisterns
parallel-arranged
Besides
the
gated,
ences
were
In
tubular-lamellar
wide
and
material
contained
observed.
The
displayed
frequently
each
showed
with
a mitochondrion.
complexes.
a
juxta-mitochondria!
invaginations
These
cristae
occupied
membrane
invaginations,
17).
infrequent.
rare.
by
reticulum
“pouch”-like
a fusion
Their
and
were
flocculent
were
was
cell
between
very
The
gonocytes
Sertoli
cell
the
thin
between
they
be observed.
often
ribosomes
was
usually
the opposite
cisternae
persed
cases
and
however,
were
die-
In
surrounded
frequently
of
be
during,
very
the
relationship
brane
while
seen
could
endoplasmic
arrangement
could
14).
cells
the
cells
were
(Fig.
rare
were
form.
rare.
the
tunnel
of
the
Coated
on
the
cords
elongated
uniform.
both
from
lamina.
more
Sertoli
mostly
from
were
The
spaces
developed.
Sertoli
ever
elongated-ovoid
the
In
cilia
mitochondria
was
stacks
vesicles
was
period.
cases
The
several
uncoated
centriole
in a narrow
most
of
expect
basal
be
by very
and
differed
changes
cords.
a single
intercellular
gonocytes;
developed
sex
to
The
were
were
of
by
seemed
gonocytes
18).
toli
would
the
deep
pouches
in which
embedded.
Frequently,
separated
0
of
were
showed
were
5
of
p.c. Sertoli
cells
35 days
p.c. The
some
sections
cytes
9. Correlation
of the cords
and Sertoli
cells
and
per
tubules
between
the
the average
age group.
(.103))12
surface
of
number
of gono-
cross
SERTOLI
CELL
DEVELOPMENT
IN THE
FIG.
10. Fifty-two
days p.c.-control
animal.
Sertoli
cells adjacent
to
intercellular
space
(arrowheads).
Bundles
of microfilaments
(small
arrows)
dense plaques
on the cell membrane.
A large lipid droplet
(L) can be seen.
FIG.
11. Fifty-two
days
p.c.-decapitated
animal.
A section
parallel
plate-like
layer
of microfilaments.
X21,000.
FIG.
12. Fifty-two
days
p.c-control
animal.
A crisscross
arrangement
served
in the
basal
cell compartment
of the Sertoli
cell.
X 35,000.
PIG
571
one
another
showed
a uniform
narrow
seemed
to originate
from
electronX21,000.
to the basal cell membrane
showing
a
of intermediate
filaments
could
be ob-
572
FIG.
13. Thirty-five
Sertoli
cells. X 21,000.
FIG.
14. Fifty-two
uncoated
vesicles
could
area (arrow).
X21,000.
VAN
days
days
p.c.-untreated
p.c.-control
be observed.
VORSTENBOSCH
animal.
animal.
A cilium
Highly
The
Golgi
in a narrow
ET AL.
complex
organized
apparatus
tunnel
nucleoli
were
was well developed.
(cell
invagination?)
common
Numerous
could
be seen
in early pig
coated
in the
and
Golgi
SERTOLI
CELL
DEVELOPMENT
IN THE
PIG
573
-
.‘.-
‘a.
17
..-,,
FIG.
chondrial
FIG.
15. Thirty-five
membranes
16. Thirty-five
chondrial-ER
FIG.
large
cistern
mitochondrial
membrane
17.
Thirty-five
within
its
-
was occupied
days
membrane
ER membrane
with
p.c.-untreated
pocket-like
contained
-
____
animal.
A peculiar
arrangement
in the Sertoli
cells. X 36,000.
animal.
Higher magnification
days p.c-untreated
could
be observed
days p.c.-untreated
ribosomes,
animal.
pouches
ribosomes.
while
which
X 25,000.
-
the
ER
membrane
complex.
mitochondria
was smooth.
The
are
on the ER-juxta-mitocomplex.
of a similar
the opposite
Mitochondrial
in
of ribosomes
cisternae
located.
Note
The juxta-mitoX 55,000.
fused,
creating
that
the
juxta-
a
574
VAN
VORSTENBOSCH
ET AL.
FIG.
18.} .. y.two p.c.-decapitated
animal.
y thin cell processes
of the Sertoli
cells enveloped
the gono(C). X 25,000.
FIG.
19. Fifty-two
days p.c.-control
animal.
The ER was organized
in a loose
concentric
whirl.
The membranes
were smooth
over
large
areas but still contained
patches
of ribosomes.
At the bottom
left is a gonocyte(G).
X21,000.
FIG.
20. Fifty-two
days p.c.-decapitated
animal,
the ER was more parallel
arranged.
Also one could
observe
transitional
ER. X 25,000.
cytes
SERTOLI
lar form.
The ER
arrays
of profiles
are
a regular
tity
varied
but
seen
days
The
p.c.
DEVELOPMENT
(Fig.
sys-
sex cords
however
relationship
between
not so evident
at 52
(Christensen
testicular
19,
extremely
mostly.
and ER
ER
20).
Their
quan-
well-developed
The
was
was
irregularly
covered
with
patches
of ribosomes
but areas
completely
devoid
of ribosomes
could
also be observed.
The
cisternae
were
filled
with
finely
dispersed
floecular
material.
The
number
of free
ribosomes
and
polysomes
seemed
free
of ribosomes
In all Sertoli
was
cells
rather
bodies
be
membrane-bound
Lysosomes
(identified
criteria)
a very
were
also
large
lipid
present.
droplet
sheath
At
of
around
62 days
cells
From
was
62
sex
cords
hardly
or
Cisternae
were
and polysomes
end
of
in the
the
animals,
obvious
The
remained
p.c. the
of
this
cells
sex
Free
period
of
into
cords.
a
the
arstacks.
no
signs
peritubular
myoid
Between
were
cell
layer
untreated
animals
were
whole
no
ap-
DISCUSSION
the
Leydig
between
for
the
very
pig
early
ultrastructurally
(1975a,b,
endoplasmic
the
testis
until
before
logically
testis
Leydig
(Raeside
can
cells
gonad
have
been
27 days p.c.
1976).
During
reticulum
pig
during
to
that
(SER)
period
can
exception
which
reticulum
produce
can
and
be
are
investigated
(Pelliniemi,
no smooth
be observed
of a very few
continuous
(RER).
The
testosterone
identified
Sigman,
and
1975).
even
steroids
steroid
synthesis
end of the
the
find
that
p.c.
can
The
fetal
(Dallner
form
is also
observed
guinea
et al.,
pig
a
as was
1966).
fetal
period
Possibly
sharply
for
6-wk-old
sizing
Our
important
and
then
rises
Sertoli
of the
apparatus
observed
again
male
those
of
quite
occurs
(Tran
et
cells
AMU
is
RER
(Tran
and
seems
not to
(Tran
and
observations
it seems
in the
pig
in
process.
Secretion
in protein
synthe-
secretion
cells,
are absent
ultrastructural
with
relatively
hormone
(Tran
et al., 1977).
This
at high concentrations
after
birth
the concentra-
al., 1981).
In the bovine
localized
in the cisternae
Josso,
1982).
The Golgi
be involved
in the
granules,
frequently
in
period.
Until
we did not
observed
glycoprotein
sexual
differentiation
protein
is also produced
at 90 days p.c. Shortly
drops
(Black
mouse
ER can
is involved
in the early
observation
SER
ER
transitional
piglets
(Chevalier,
1978).
The
early
Sertoli
cell
produces
high
concentrations
of anti-Mtillerian
(AMH),
Sertoli
onwards.
A
be observed
period.
a
This
the
the
of
this
of Pelliniemi
of mitochon-
mitochondrion-ER
as much
the
The spatial
relaand endoplasmic
in the early
fetal
this
be
RER.
in
in
nor those
evidence
1969)
and in the fetal
Singh,
1980).
Transitional
complex
Tran
Josso,
are
et
possible
during
al.
1982).
fully
(1981).
that
the
AMH
entire
period.
Intermediate
much
lower
filaments
were
found
quantities
as in Leydig
cells
but
in
of the
same
developmental
period
(van Vorstenbosch
et al.,
1982,
1984).
In the
Sertoli
cells
they
show
the
same
spatial
relationship
with
the
nuclear
and mitochondrial
in Leydig
cells (Aguas,
et
cells
morphoIn
and
cells
the
fetal
undifferentiated
in Sertoli
cells, with the
small
concentric
profiles
with
rough
endoplasmic
early
cells
seems
SER
Sertoli
Therefore
synthesis
In
observed
from
52 days
in relationship
compatible
observation
not
period
similar
in
in
3j3-HSD,
the SER
1969).
In the period
dealt
with
in
tubular
cnistae.
mitochondria
we observed
wanes
change
(HSD)
Raeside,
1972).
be localized
in
dna
with
pure
tionship
between
reticulum
(ER)
tion
ribosomes
the
decapitated
differences
throughout
well
was
ER
of parallel
mostly
filled.
were abundant.
controls
and
morphological
parent
period.
changed.
concentrically
form
of a differentiation
mesenchymal
surrounding
Sertoli
a smooth
appearance
and
of the
adult
Sertoli
cells
The
nucleoli
were
promi-
composed
whirls,
the
was
produce
predominantly
At
SER
the sex cords.
p.c. the ultrastructure
of
and
to
cells.
Neither
this study
(1975a,b,
1976)
showed
described
In most
Sertoli
cells
could
be seen in the
the
Golgi
apparatus
Until
about
90 days
observed
study,
dehydrogenase
histochemically
and Gillim,
differentiation
and Christensen,
(Almond
and
nuclei
showed
mostly
indentations
typical
were
very
scarce.
ranged
ob-
on morphological
similar
to that
at 52 days
p.c.
until
the end of the observation
the
nent,
and
developed.
could
575
PIG
(Moon
is known
the cell.
From
52 days
onwards
cells
formed
a loose
incomplete
very
days
period,
SER
electron-
served.
basal
part
mesenchymal
constant.
not observed.
a few strongly
dense,
IN THE
period,
3j3-hydroxysteroid
has been
demonstrated
was arranged
either
in parallel
or in concentric
whirls;
both
finding
tems
were
mitochondria
CELL
the
al.,
1982).
in this
Whirls,
period,
The
bundles
of
cell
membrane
1981;
outer
membranes
van Vorstenbosch
characteristic
were
never
microfilaments
have
been
for
as
Leydig
observed.
adjacent
described
to
for
VAN
576
Sertoli
In
cells
in
Sertoli
the
cells
microfilaments
1975).
Toyama
for
exist
in
its
the
been
immature
Dym,
pig
FSH
an actin-like
claim
Sertoli
is known
to
cells
in
and,
FSH.
organization
these
actin-like
and
1981),
FSH
the
filaments
and
in this
disorganization
(Chevalier
way
cell
that
actin
plays a role in
et al., 1980)
and in its
secretion
(Welsh
Decapitation
1980).
a deprivation
tropins
of the
and as a consequence
Leydig
cells,
resulting
testosterone
ptuitary
produces
at 60
days
seems
that
(Colenbrander
not
surprising
structural
this
1982,
that
days
dose”
leydig
p.c.
It
might
1970;
The
undisturbed
the
decapitated
tubular
of
The
close
it is
show
no
ultra-
that
level.”
seems
and
is not
Porter,
(cord)
the
fetal
and in the
et al., 1979)
diameter
tubules
and
(cords)
1980).
similarity
in
gonado-
rhesus
monfetal decapian increase
a reduction
can
cellular
be
in
observed.
organization
Sertoli
cells in normal,
control
animals
does
not
give a clue
difference
in sex cord
diameter
and
for
and
of
decapitated
the obvious
the number
of cells per cross section.
The positive
correlation
between
the number
of cells
and cord
surface
per cross
section
indicates
an increase
in cells
but
not
constant
cell
ratio
surface-unit
animals.
cells
in
indications
the
in normal,
increase
An
is unlikely,
is about
animals
dimension,
of
since
equal
in both
(Colenbrander
were
found
since
number
control
in the
the
and
total
number
control
et al.,
of
an
there
of
is
cells
a
per
decapitated
number
of germ
of
cells
and decapitated
1979)
and
increase
of the
the
cords.
from
of the
of
the
a
resulting
Another
reshaping
in a change
resulting
component
Dufaure
caused
of
the
the
intra-
absence
of
testis.
The authors
are most
indebted
to Prof.
Dr. W. A.
de Voogd
van der Straaten,
for helpful
and encouraging
discussions
as well
as for reading
the manuscript;
Dr.
A. A. Macdonald
who
did a number
of caesarian
sections
and corrected
the English;
Mrs. C.M.J.E.
van
Rossum-Kok,
for general
technical
assistance;
Mr. 0.
van der Veen and Mr. H. Otter
for light
micrography
and E. M. printing;
Mr. J. Nokkert,
Mr. G. Hol and Mr.
H. van Dijk for animal
care; and Mr. J. van Vorstenbosch and Mrs. L. Michielsen
for typing
the manuscript.
Aguas,
sub-
by
pressure
cell
for
found
and
have
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key (Gulyasetal.,
1977)
tated
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length
days
development
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fetal
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that
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tropic
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in
70
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Saunders
it
gonado-
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1982),
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1984).
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the disappearance
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Chevalier
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might
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sources,
could
be explained
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in a reshaping
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the
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of
This rearrangement
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ences
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