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Modultion of the Golgi apparatus in stimulated and nonstimulated prolactin cells of female rats.

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THE ANATOMICAL RECORD 235:353-362 (1993)
Modulation of the Golgi Apparatus in Stimulated and Nonstimulated
Prolactin Cells of Female Rats
A. RAMBOURG, Y. CLERMONT, M. CHRgTIEN, AND L. OLIVIER
Dkpartement de biologie cellulaire et molkculaire d u CEA, Centre d‘ktudes nuclkaires de
Saclay (A.R., M.C.), and Facultk de mkdecine Pitik-Salpktri6re (L.O.),Paris, France;
Department of Anatomy, McGill University, Montreal, Quebec, Canada (Y.C.)
ABSTRACT
The three-dimensional structure of the Golgi apparatus
and its compartments in prolactin cells has been examined in lactating rats
in which secretion of prolactin was suppressed by removing the litter or
stimulated by allowing the pups to suckle again. As soon as 2 hr after
removal of the litter, large irregular progranules and numerous large pale
vesicles accumulated in the trans-Golgi area together with vesicular or
tubular fragments. The cis-tubular network was no longer recognizable on
the cis-face of the Golgi ribbon; the saccules of the midcompartment were
partitioned by narrow fissures and also became perforated in register by
numerous fenestrations of various sizes and irregular contours. The concomitant appearance of numerous vesicles in the cavities thus formed as
well as in the surrounding cytoplasm indicated that they probably arose by
the progressive cavitation and fragmentation of saccules of the mid compartment. Such a process, which reached a maximum between 4 and 6 hr
after removal of the litter from the mother, was no longer observed at 8 and
12 hr, at which time intervals the Golgi apparatus was reduced in size with
no cis-tubular elements and progranules on its trans-aspect and few vesicles in its surroundings. When mothers, separated from their litters for a
period of 12 hr, were returned to their pups for 20 min, the cis-tubular
network reappeared on the cis-aspect of the Golgi stacks and presumably
formed by fusion of vesicles and anastomosed tubules located next to the
cisternae of the rough endoplasmic reticulum. In addition, the structure of
the midsaccules returned to the stimulated condition, and early progranules were again segregated within the trans-most saccules of the Golgi
stack. Hence, the Golgi apparatus of prolactin cells was rapidly and deeply
modified in the presence or absence of stimulation. o 1993 Wiley-Liss, Inc.
Key words: Progranules, Tubular network, Lactation
Electron microscopic stereoscopic studies of the Golgi
apparatus revealed that in most cells studied this organelle was composed of a continuous ribbon-like
structure, which branched and anastomosed to form a
loose juxtanuclear spheroidal mass (Rambourg and
Clermont, 1990). In transverse sections of this ribbon
and in the cis-trans axis as defined by Ehrenreich et
al. (1973), the stacked elements observed have been
subdivided into three compartments. The first or ciscompartment is formed by a network of osmiophilic
membranous tubules referred to as the cis-element
(Rambourg et al., 1974). Subjacent to it, several superposed flattened elements constitute a mid-compartment. Continuous along the whole Golgi apparatus,
this compartment presents compact zones made up of
stacked, poorly fenestrated saccules alternating with
highly fenestrated or tubular elements forming less
compact intersaccular zones. A third or trans-compartment consists of one or more sacculotubular elements.
In contrast to the elements of the midcompartment,
0 1993 WILEY-LISS, INC.
these are not continuous along the Golgi ribbon, but
usually show a “peeling-off” configuration. In some
glandular cells, the elements of this trans-compartment are involved in the segregation and packaging of
secretory proteins and the formation of prosecretory or
secretion granules (Rambourg and Clermont, 1990).
In a recent steroscopic study of the Golgi apparatus
of prolactin cells of lactating female rats, it was observed that large, elongated, polynodular progranules
form within trans-saccules, which progressively became extensively porous or fenestrated. These progranules, with some tubular residues still attached to them,
Received June 3, 1991; accepted July 16, 1992.
Address reprint requests to Dr. A. Rambourg, Departement de biologie cellulaire et molkculaire, Centre detudes nucleaires de Saclay,
Gif-sur-Yvette 91191 Cedex, France.
354
A. RAMBOURG ET AL.
Fig. 1. Low-power photograph of the Golgi region of a prolactin cell
from a lactating rat. Several stacks of saccules cut transversely or
obliquely are seen at the periphery or in the center of the Golgi region
(S).In such a thin section, the peripheral stacks of saccules are long
and separated from each other by narrow gaps. On their cis-face, they
show a linear array of membranous profiles corresponding to the cisGolgi network or cis-elements (CE). Underlying these there are some
four or five flattened saccules. On the trans-aspects of the stacks,
there are some “peeling o f f saccules (arrows), some showing distended portions filled with secretory material (arrowheads). Progranules or mature granules (g) are present. ER cisternae are seen outside
and inside the Golgi region ER (ER). Vesicles are associated with
stacks; some are small and within the 80 nm range (v). A few larger
vesicles (V) are seen in the central portion of Golgi region between
mitochondria (m) and lysosome-like bodies (L). N, nucleus. x 31,000.
detach from the trans-aspect of the stacks and eventually transform into polymorphous and then mature
granules (Rambourg et al., 1992). In the pituitary
glands of lactating females, the prolactin cells exhibit
an enlarged Golgi apparatus associated with numerous
polynodular tubular progranules, polymorphous and
mature secretion granules. In nonlactating animals, in
contrast, the prolactin cells are smaller and show a
MODULATION OF THE GOLGI O F PROLACTIN CELLS
Fig. 2. Portion of the Golgi of a prolactin cell from a lactating rat
ribbon showing the Golgi stacks in side view on the left 611, in
oblique view on the right (S2), and in face view in the center 631, all
continuous with each other. In the face view of the Golgi ribbon, the
saccules show some small pores (arrows) but are free of large fenestrations. Some small, 80 nm vesicles are seen along the saccules.
Numerous ER cisternae are seen both along the cis and trans aspects
of the stacks. x 39,000. Abbreviations for this figure and for subsequent figures are as given for Figure 1.
355
Fig. 3. Low magnification of the Golgi apparatus of a prolactin cell
2 hr after the separation of the pups from their lactating mothers. The
stacks of saccules are small and disorganized. The cis-elements of the
stacks are unidentifiable. On the cis- but even more so on the transaspects of the stacks, there are numerous vesicles of various sizes
(straight arrows). “Wells” or small vesicles containing cavities in the
stacks are indicated by curved arrows. The asterisk indicates a cluster
of small vesicles and short tubules between the ER and the Golgi
stack. Progranules were not seen associated with the cis-face of the
stacks, but polymorphous granules (g)are present in the Golgi region.
ER, cisternae of endoplasmic reticulum. x 31,000.
Figs. 4 , 5 . Stereopairs of small portions of the Golgi apparatus from
prolactin cells 2 hr after separation of the litter from their mother. A
single magnified stereoscopic image may be obtained by utilizing a
properly adjusted (at 65 mm) binocular lens.
Fig. 4. Numerous small vesicles are seen in the vicinity of the Golgi
stack, some being enclosed within multivesicular-like bodies (arrows).
Polynodular or polymorphous granules are seen at proximity (g).
x 2,900.
Fig. 5. Face view of a midsaccule, showing numerous irregular pores
or fenestrations. Some are linearly arranged. A larger fenestration
corresponding to a face view of a well (w) is indicated. Numerous
small vesicles or tubules are seen in proximity (arrow). x 40,000.
Fig. 6. Low magnification of the Golgi ribbon in oblique view of a
prolactin cell 6 hr after the separation of the mother from her litter.
The saccules show numerous pores of various sizes, some being linearly arranged (arrows),while other perforations are slit-like (arrowheads). Other larger fenestrations in register associated with small
vesicles form numerous “wells” (w). Numerous vesicles (v) are seen in
the Golgi region. g, polynodular or polymorphous granules; m, mitochondria. x 40,000.
MODULATION OF T H E GOLGI OF PROLACTIN CELLS
poorly developed Golgi apparatus, which, however, included some polymorphous and mature secretion granules. Smith and Farquhar (1966) have reported that
removal of the suckling youngs from their mother induced a marked regression of the Golgi apparatus and
the elimination of secretion granules by the lysosomal
system of the prolacting cells. The purpose of the
present study is to investigate in electron microscope
stereopairs the modifications of the various components of the Golgi apparatus at short time intervals,
i.e., from 2 to 12 hr, following the separation of the
litter and the mother. Furthermore, the Golgi apparatus of prolactin cells has been examined in females that
had been separated from their litters for 12 h r and then
returned to their suckling pups for a period of 20 min.
This permitted a n analysis of the modifications of the
Golgi apparatus compartments in stimulated and nonstimulated prolactin cells in situ.
357
sues postfixed with reduced osmium, it formed a ribbon-like structure made up of stacked elements
delimiting a roughly ovoid area (Fig. 1). When the
plane of section through such a ribbon was perpendicular or slightly oblique, a distinct cis-trans polarity
was easily observed through the Golgi stacks. On the
cis-aspect of the ribbon facing the rough endoplasmic
reticulum (ER), the first or cis-element consisted exclusively of anastomosed membranous tubules (Fig. 11,
which corresponded to the cis-osmiophilic network described in other cell types (Rambourg et al., 1974; Rambourg and Clermont, 1990). Underlying this ciselement were three to five slightly dilated and parallel
saccules (Fig. 1). When seen in front views, these elements making up the midcompartment were perforated with pores of various sizes (Fig. 21, but they were
usually deprived of large fenestrations in register or
“wells” already described in the Golgi apparatus of
many other cell types (Hermo et al., 1980; Rambourg
MATERIALS AND METHODS
and Clermont, 1990). On the trans-aspect of the Golgi
Lactating female rats used in this experiment were ribbon, two or three flattened and curved elements
first allowed to feed their pups normally. On the fifth were separated from each other in the cis-trans axis by
day postpartum, the females were separated from their cytoplasmic areas and thus displayed a “peeling off”
pups and sacrificed 2 , 4 , 6 , 8 and 12 h r after removal of configuration (Fig. 1). They usually showed along their
the suckling youngs. Prior to sacrifice, the females length or a t their extremities dilatations containing
were anesthetized by a n intravenous injection of pen- nodular masses of various electron densities, corretothal in a saline solution containing heparin. Addi- sponding to early prosecretory granules. Besides these
tional females separated from their litters for a period trans-most saccules, the cytoplasm of the trans-Golgi
of 8 h r were allowed again to feed their pups for 20 min region contained polynodular tubular progranules and
just before sacrifice. Four females were utilized for a few large pale vesicles. Small vesicles, 80 nm in dieach time interval selected. Females were fixed by per- ameter, were seen next to and on all sides of the Golgi
fusion through the left ventricle with a fixative con- ribbon (Fig. 1). On the cis-aspect of the stacks, between
taining 3% glutaraldehyde in 0.1 M cacodylate buffer, the proximal ER cisternae and the cis-element, they
pH 7.4, with 5 mM/liter Cacl,, 0.5 g/liter picric acid, were seen in small clusters and intermingled with a
and 4% (v/v) hydroxyethyl starch solution Haes accord- few narrow, branching tubules. The diameter of the
ing to the method of Kaissling et al. (1985). Following latter tubules was similar to that of the anastomotic
1 min of perfusion, the pituitary glands were removed tubules forming the cis-element (Fig. 1; Rambourg et
and stored at 4°C in buffer. On the following day, they al., 1992). Some small vesicles were also present along
were postfixed for 1 h r a t room temperature in a 1 :l the edges of the stack or in the trans-region of the
mixture of 2% aqueous osmium tetroxide and 3% organelle. In this region, occasional larger and paler
aqueous potassium ferrocyanide (Karnovsky, 1971). spherical vesicles were also visible (Fig. 1).
Dehydration was carried out in ethanol followed by
Ultrastructural Modifications After Removal of
embedding in Epon. Thin sections were cut with a n
Suckling Young
LKB automatic ultramicrotome and counterstained for
2 min with lead citrate prior to examination with a
As soon as 2 h r after removal of the young, signs of
CM12 Philips electron microscope at 80 kV.
ultrastructural disorganization were observed in the
For stereoscopy, grids were placed on the goniomet- Golgi apparatus of prolactin cells. Large irregular proric stage of the electron microscope, and stereopairs granules accumulated in the trans-Golgi region among
were obtained by taking pictures of the same field after a large variety of saccular and tubular fragments. The
tilting the specimen - 15” and + 15” from the 0” posi- whole area was filled with vesicles of various sizes and
tion. A three-dimensional magnified image of the densities, while the number of large pale vesicles instructures was obtained by looking at properly ad- creased significantly (Figs. 3-6). At this time, numerjusted pairs of such photographs with a stereoscopic ous multivesicular bodies with a smooth membrane,
binocular lens.
frequently associated on their cytoplasmic side with
short membranous tubules, were seen (Fig. 4). In secRESULTS
tions perpendicular to the Golgi ribbon, the slightly
Structure of the Golgi Apparatus in Control Animals
distended saccules of the midcompartment were perfoIn the pituitary gland of control, lactating females, rated in register to form cavities or “wells,” with their
the prolactin cells were numerous and large. As previ- mouths directed toward the cis-face of the golgi stack
ously described from thick sections treated to demon- and their bottoms closed by a trans-saccule (Fig. 3).
strate TPPase activity (Rambourg et al., 19921, they Such cavities were usually associated with numerous
displayed a large Golgi apparatus consisting of a con- small vesicles. Occasionally, only one or two saccules
tinuous, twisted ribbon-like structure that branched located deep in the stack of saccules were perforated,
and anastomosed to form a hollow spheroidal mass lo- and the cavity thus formed contained a few vesicles
cated in the juxtanuclear area. In thin sections of tis- (Fig. 4). Face views of saccules showed, in addition to
358
A. RAMBOURG ET AL.
Fig. 7. Low magnification of the Golgi region from a prolactin cell 8
hr after the separation of the mother from her litter. In this Golgi
apparatus, the stacks of saccules are small, the cis-elements are not
identifiable, and polynodular granules are absent. However, some secretion granules are present (g), and membrane bound bodies com-
posed of small vesicles and secretory dense material, presumably lysosomal in nature, are identified (curved arrows). Vesicles of various
sizes are present, but they are less abundant than at earlier time
intervals. ER, cisternae of endoplasmic reticulum; m, mitochondria.
x 30,000.
“wells,” pores of various configurations, some suggesting that pores coalesced to form larger perforations
(Fig. 5). On the cis-aspect of the Golgi ribbon, the ciselement with its characteristically anastomosed membranous tubules was hardly recognized. The groups of
vesicles facing cisternae of the rough ER were less homogeneous in appearance and usually consisted of vesicular and tubular elements of various sizes and densities (Fig. 3).
At 4 and 6 h r after removal of the litter, the number
of saccular and tubular fragments showed a tendency
to decrease in the trans-Golgi region, while the saccular elements of the midcompartment were still heavily
perforated by cavities (Fig. 6). In face views, the presence of narrow linear fissures suggested a partitioning
of the midsaccules (Fig. 6). The cis-element was no
longer visible, and only small groups of small vesicles
and tubules were observed proximal to cisternae of the
rough ER (Fig. 6).
At later time intervals, e.g., 8 and 12 h r after removal of the litter, saccular and tubular fragments
had decreased markedly in the trans-Golgi region.
Prosecretory granules were replaced by multivesicular
bodies containing nodular masses of various electron
densities (Fig. 7). The parallel saccules of the midcompartment were poorly perforated. A limited number of
small vesicles and occasional large pale vesicles were
encountered mainly at the edges of the midsaccules or
next to the trans-elements (Fig. 7). The overall structure was that of a quiescent Golgi apparatus reduced in
size and seemingly not producing secretory granules.
Effects of a Short Stimulation on the Structure of the
Golgi Apparatus
When the prolactin cells were restimulated by allowing the pups to suckle again for a period of 20 min, the
following characteristic modifications of the Golgi ap-
MODULATION OF T H E GOLGI O F PROLACTIN CELLS
paratus were observed. The anastomotic tubules making up the cis-element, which had disappeared after
removal of the pups, were again present on the cis-face
of the Golgi stacks (Fig. 8). They were frequently associated with homogeneous groups of vesicles and small
anastomosed tubules located between the cisternae of
the rough ER and the cis-elements (Fig. 8). The parallel saccules of the midcompartment were poorly fenestrated and slightly dilated (Figs. 8, 9). On the transaspect of the Golgi ribbon, the “peeling-off” saccules
contained electron-opaque nodular masses and became
more fenestrated (Fig. 9). Polymorphous and polynodular tubular progranules reappeared in the trans-Golgi
area, where they were interspersed among a few saccular and tubulovesicular remnants (Fig. 9).
359
males (e.g., at 2, 4, and 6 hr). It is therefore suggested
that most of the small vesicles present in the Golgi
area, at these early time intervals, were originating
from the midsaccules of the Golgi ribbon as they perforated.
Besides cavities, face views of the involuting Golgi
apparatus showed a partitioning of the midsaccules by
narrow fissures (Fig. lo), resembling those previously
described in CMPase-reactive trans-saccules in the
Golgi apparatus of rat spermatids (Thorne-Tjomsland
et al., 1988). In this cell type, the formation of fissures
resulted from the fusion of linearly arranged minipores
and was seemingly followed by a fragmentation of the
whole trans-most saccules into large vesicles. Such a
fragmentation is also likely to occur in the mid-Golgi
saccules of the prolactin cell and thus to contribute,
DISCUSSION
with the formation of the above-mentioned larger fenSmith and Farquhar (19661, studying the modifica- estrations, to the reduction in size of the Golgi apparations of prolactin secreting cells after removal of the tus observed after removal of the suckling young from
suckling pups from lactating rats, reported a progres- thc ir mothers.
sive involution of the protein synthetic apparatus and
Modulation of the cis-Compartment
sequestration of secretory granules into multivesicular
and dense bodies. They noted that the content of imIn most cell types, the first element of the Golgi stack
mature granules was preferentially segregated within is :i network of anastomosed tubules, which is usually
multivesicular bodies. Indeed, in the present study, selectively stained with osmium after prolonged osmimultivesicular bodies containing progranule secretory cation (Rambourg et al., 1974; Noda and Ogawa, 1984;
material were also found in the trans-Golgi region as Lindsey and Ellisman, 1985; Rambourg and Clermont,
soon as 6 hr after removal of the litter, yet the main 1990) and has been recently referred to as the cis-Golgi
finding of the present study was the rapid and exten- network, or CGN (Huttner and Tooze, 1989; Mellman
sive alterations of the Golgi apparatus itself, which did and Simons, 1992). Overlying this network of anastonot form secretory granules any longer and progres- mosed tubules, numerous clusters of small vesicles are
sively reduced its size within the first 12 hr after in- frequently seen, in close association with the cisternae
terruption of lactation.
oft he rough ER and are currently said to form by budding from the ER cisternae. They are considered as
Modulation of the Mid-Golgi Compartment
carriers of proteins from the ER to the Golgi apparatus
A striking feature observed during the progressive (Pa lade, 1975; Goldfischer, 1982; Farquhar, 1985; Pfefinvolution of the Golgi apparatus of the prolactin cells fer and Rothman, 1987). In some cell types, such as
was the formation of large fenestrations in register in spermatids (Hermo et al., 19801, fibroblasts (Marchi
the parallel saccules making up the midcompartment and Leblond, 19831, pancreatic acinar cells (Merisco et
of the Golgi ribbon and the concomitant appearance of al., 1986; Rambourg et al., 19881, plasma cells (Ramnumerous vesicles within the Golgi area (Fig. 10).Such bourg et al., 19891, and the prolactin cells in the
cavities or “wells,” which are always associated with present study, vesicles are intermingled with branched
small, 80 nm vesicles, were first described in the Golgi and sometimes anastornosed membranous tubules (Fig.
apparatus of spermatids (Hermo et al., 1980) and aci- 10).These anastomosed tubules closely resemble in size
nar cells of sublingual cells (Ichikawa et al., 1982) and and three-dimensional arrangement the tubular elethen were observed in many other cell types (Ram- ments of the cis-Golgi network or cis-element. They
bourg and Clermont, 1990). In the Golgi apparatus of contain trans-membrane proteins, which are also found
type A ganglion cells, as in the present study, one or in the cis-tubular network (Saraste et al., 1987;
two saccules located deep in the stack showed perfora- Sch weizer et al., 1988, 1990). It has thus been postutions and formed closed cavities, which contained one latcd that the latter are derived from the former and
or two 80 nm vesicles. A close examination of face forin the so-called intermediate compartment (Schweiviews of small vesicles and their relation to saccules zer et al., 1990), which is involved “in transport from
indicated that the small vesicles could originate only in the ER t o the Golgi as well as in the recycling of Golgi
situ and that smaller perforations likely transformed components back to the ER” (Mellman and Simons,
into larger ones with a concomitant production of small 1952). In fact, when prolactin cells lacking a clear-cut
vesicles (Rambourg and Clermont, 1986). Ichikawa et cis-tubular element were again stimulated by suckling
al., (1982) also postulated the formation of small vesi- pups, the reappearance of the latter on the cis-face of
cles in similar cavities in the Golgi apparatus of secre- the Golgi apparatus occurred simultaneously with the
tory cells of the sublingual glands. In the Golgi appa- forination of numerous intermediate clusters of vesiratus of prolactin cells, such cavities, which were clet, and anastomosed tubules. Such an observation
rarely found in lactating animals, were also rare in supports the hypothesis that vesicles derived from the
rats separated from their young for at least 8 hr. In the ER give rise to membranous tubules, which in turn
latter case, there was also a marked reduction in the ana stomose to form the cis-Golgi network or cis-elenumber of Golgi vesicles as compared with that ob- ment, which would thus be continuously renewed.
‘I
he modifications described above were particularly
served a t previous time intervals in postlactating fe-
360
A. RAMBOURG ET AL.
Fig. 8. Low magnification of the Golgi region of a prolactin cell of a
female separated from her litter for 8 hr and then returned to her
suckling pups for 20 min. Note that the stacks of saccules are long.
The cis-tubular Golgi network element is present on the cis-face of the
stacks (CE); few clusters of small vesicles or tubules are seen on the
cis side of the stacks (open arrows). g, polynodular or polymorphous
granules. m, mitochondria. x 30,000.
Fig. 9. Stereopair of an oblique section through a stack of saccules
from a prolactin cell of a female separated from her pups for 8 h r and
returned to her litter for feeding for 20 min. On the cis-aspect of the
stack, a cluster of vesicles and tubules are seen between the ER cisternae (ER) and the Golgi stack (open arrow). The cis-element is difficult to identify in this oblique section of the stack. On the transaspect of the stack, a fenestrated saccule (asterisk) shows a spherical
dense body corresponding to the core of a forming polynodular granule
(arrow). A polynodular granule is seen in proximity (g). x 30,000.
S I T 3 3 NIL3VTOXd
a
do I3103 3 H L do NOILVTnaOW
362
A. RAMBOURG ET AL.
rapid. On the one hand, the disappearance of the ciselement, the resorption through fenestration of the
midsaccules, and the arrest of progranules formation
were already observed 2 hr following the separation of
the mother from its litter. On the other hand, the ciselement as well as the progranules reappeared within
20 min following the suckling stimulus. Hence the kinetic processes observed in the Golgi apparatus of prolactin cells are rapidly and deeply modulated in the
presence or absence of stimulation.
ACKNOWLEDGMENTS
The work done a t McGill University was supported
by a grant of the Medical Research Council of Canada.
The technical assistance of Corinne Le Moal, Centre
d'etudes nucleaires de Saclay, and Beatrice Roncier,
Facult6 de medecine Pitie-Salpetriere, is gratefully acknowledged.
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