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| [[File:Mark_Hill.jpg|90px|left]] This historic 1920 paper by Wieman described human adrenal or suprarenal gland development.  
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| [[File:Mark_Hill.jpg|90px|left]] This historic 1920 paper by Wieman described human {{adrenal}} or suprarenal gland development.  
 
+
<br><br>
See [[Endocrine - Adrenal Development]]
+
'''Modern Notes:''' {{Adrenal}}
 
+
<br><br>
 
{{Endocrine Links}}
 
{{Endocrine Links}}
  
 
[https://www.ncbi.nlm.nih.gov/pubmed/?term=adrenal+development Search PubMed adrenal development]
 
[https://www.ncbi.nlm.nih.gov/pubmed/?term=adrenal+development Search PubMed adrenal development]
 
 
|}
 
|}
 
{{Historic Disclaimer}}
 
{{Historic Disclaimer}}
 
 
{{Ref-Wieman1920}}
 
 
Wieman HL. [[Paper - Observations in connection with the early development of the human suprarenal gland|Observations in connection with the early development of the human suprarenal gland]] (1920) Anat. Rec.<noinclude>[[Category:Template]][[Category:Reference]][[Category:1920's]][[Category:Adrenal]]</noinclude>
 
 
 
 
=Observations in Connection with the Early Development of the Human Suprarenal Gland=
 
=Observations in Connection with the Early Development of the Human Suprarenal Gland=
  
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==Introduction==
 
==Introduction==
The purpose of this contribution is to call attention to certain details in connection with the development of the human suprarenal gland, because they happen to be clearly illustrated in the material at hand. This material consists of two human embryos, nos. 1 and 4 of the author’s collection, which were obtained some years ago, freshly killed and fixed, from Drs. H. L. VVoodward and Charles Goosmann, of Cincinnati. No. 1 had been killed in a bichromate-acetic mixture, and no. 4 in Bouin’s solution. The former measured 9 mm., crown—breech, and the latter 12 mm. Both measurements were made in 85 per cent alcohol after fixation, so that the sizes of the living embryos were somewhat larger than these figures. The embryos were embedded in paraffin, cut into serial sections of 10 [1, thickness, and stained on the slide with Delafield haematoxylin and orange G according to the method of Morris (’09). Mitotic figures abound in both embryos, but the fixation is somewhat better in no. 1.
+
The purpose of this contribution is to call attention to certain details in connection with the development of the human {{suprarenal}} gland, because they happen to be clearly illustrated in the material at hand. This material consists of two human embryos, nos. 1 and 4 of the author’s collection, which were obtained some years ago, freshly killed and fixed, from Drs. H. L. VVoodward and Charles Goosmann, of Cincinnati. No. 1 had been killed in a bichromate-acetic mixture, and no. 4 in Bouin’s solution. The former measured 9 mm., crown—breech, and the latter 12 mm. Both measurements were made in 85 per cent alcohol after fixation, so that the sizes of the living embryos were somewhat larger than these figures. The embryos were embedded in paraffin, cut into serial sections of 10 [1, thickness, and stained on the slide with Delafield haematoxylin and orange G according to the method of Morris (’09). Mitotic figures abound in both embryos, but the fixation is somewhat better in no. 1.
  
  
 
==Observations==
 
==Observations==
 +
Embryo no. 1, .9 mm. The suprarenal glands are not recognizable as distinct organs, but consist of a thickening in the mesenchyme on either side of the root of the mesentery, forming a pair of broad ridges projecting into the body cavity from the dorsal body wall. Anteriorly these suprarenal ridges are continuous with the dorsal portions of the pleuroperitoneal membranes, while posteriorly they blend with the genital ridges. Laterally each is separated from the mesonephros by a distinct groove (fig. 1). Occasionally a transverse section of a blood capillary can be seen in the center of each ridge, the beginning no doubt of the central vein of the adult organ.
  
Embryo no. 1, .9 mm. The suprarenal glands are not recognizable as distinct organs, but consist of a thickening in the
+
The ridges are made up of mesenchyme which shows no evidence of differentiation. The ramus communicans divides into two at the level of the sympathetic ganglion where one of the branches terminates, the other passing ventralward into the mesenchyme. The cells composing the sympathetic ganglia stain more deeply than the surrounding cells, but beyond this they show no appreciable differentiation. The ganglia lie on either side of the aorta somewhat dorsal to it. The nerve strands (axis cylinders?) are easily distinguished and followed, owing to the fact that they stain readily with orange G. The ventral branch of the ramus communicans, which is really the direct ventral continuation of the latter, loses itself in the mesenchyme of the suprarenal region. It seems to be unaccompanied by nerve cells (fig. 3). From the picture one gets the impression that the nerve fibers are pushing their way through the mesenchyme, blazing, as it were, a track along which the ganglion cells are to follow later.
mesenchyme on either side of the root of the mesentery, forming
 
a pair of broad ridges projecting into the body cavity from the
 
dorsal body wall. Anteriorly these suprarenal ridges are continuous with the dorsal portions of the pleuroperitoneal membranes,
 
while posteriorly they blend with the genital ridges. Laterally each is separated from the mesonephros by a distinct groove (fig.
 
1). Occasionally a transverse section of a blood capillary can be
 
seen in the center of each ridge, the beginning no doubt of the
 
central vein of the adult organ.
 
 
 
The ridges are made up of mesenchyme which shows no evidence of differentiation. The ramus communicans divides into
 
two at the level of the sympathetic ganglion where one of the
 
branches terminates, the other passing ventralward into the
 
mesenchyme. The cells composing the sympathetic ganglia
 
stain more deeply than the surrounding cells, but beyond this
 
they show no appreciable differentiation. The ganglia lie on
 
either side of the aorta somewhat dorsal to it. The nerve strands
 
(axis cylinders?) are easily distinguished and followed, owing to
 
the fact that they stain readily with orange G. The ventral
 
branch of the ramus communicans, which is really the direct
 
ventral continuation of the latter, loses itself in the mesenchyme
 
of the suprarenal region. It seems to be unaccompanied by
 
nerve cells (fig. 3). From the picture one gets the impression
 
that the nerve fibers are pushing their way through the mesenchyme, blazing, as it were, a track along which the ganglion cells
 
are to follow later.
 
 
 
In the posterior part of the suprarenal ridge, the genital ridge,
 
the subepithelial region of the mesonephros, and in the mesentery,
 
one finds large cells with clear cytoplasm and standing out distinctly from all cells (fig. 3, 4, 5). These 1 take to be germ cells
 
(primary genital cells). Their wide extraregional distribution
 
indicates that these cells undergo a rather extensive migration
 
before reaching the germinal epithelium. This of course is in
 
keeping with what is known about the early development of
 
germ cells in other vertebrates.
 
 
 
Embryo no. 4, 12 mm. The suprarenal glands in this specimen
 
are distinctly marked off from the surrounding tissues (fig. 2).
 
Each one lies with its dorsal surface against the pleuroperitoneal
 
cushion, while in close contact with its median side are bundles
 
of nerve fibers and -ganglionic clumps. Near the ventral border,
 
as shown in figure 2, some nervous tissue is pushed into the substance of the gland, but only to a very slight extent, definite immigration, according to other authors, not taking place until a
 
much later period.
 
  
The prevertebral sympathetic ganglion is now a very distinct
+
In the posterior part of the suprarenal ridge, the genital ridge, the subepithelial region of the mesonephros, and in the mesentery, one finds large cells with clear cytoplasm and standing out distinctly from all cells (fig. 3, 4, 5). These 1 take to be germ cells (primary genital cells). Their wide extraregional distribution indicates that these cells undergo a rather extensive migration before reaching the germinal epithelium. This of course is in keeping with what is known about the early development of germ cells in other vertebrates.
mass of cells clearly difierentiated from the mesenchyme. Its
 
relation to the ramus communicans is somewhat different from
 
  
that described for the 9-mm. embryo. There is no longer any
+
Embryo no. 4, 12 mm. The suprarenal glands in this specimen are distinctly marked off from the surrounding tissues (fig. 2). Each one lies with its dorsal surface against the pleuroperitoneal cushion, while in close contact with its median side are bundles of nerve fibers and -ganglionic clumps. Near the ventral border, as shown in figure 2, some nervous tissue is pushed into the substance of the gland, but only to a very slight extent, definite immigration, according to other authors, not taking place until a much later period.
evidence of a branching in the ramus cominunicans at the level
 
of the ganglion, the latter lying more directly in the path of the
 
ramus. The ventral extension of the ramus (fig. 1) now passes
 
directly ventralward from the ganglion (fig 2, 2).r.). Among the
 
nerve fibers of the ventral extension of the ramus can be seen two
 
kinds of cells: 1) those distinguished by a long narrow outline
 
with nucleus of the same shape, which are probably sheath cells
 
and, 2) large irregular cells with yellowish cytoplasm drawn out
 
into processes and possessing rounded nuclei. The latter are undoubtedly migrating nerve cells (fig. 7). They are larger and
 
difi'er otherwise in their appearance from the nerve cells found in
 
the prevertebral ganglia (fig. 6), but are practically identical
 
with the large nerve cells partially embedded in the ventral border of the suprarenal gland (figs. 2, 8). Comparing the pictures
 
presented by figures 1 and 2 -it would seem that in the latter
 
stage the nerve cells are in the actual process of migrating ventralward along the paths marked out by the nerve fibers, which
 
alone are present in the former stage. If some of these migrating
 
nerve cells are destined to enter the gland to form its chromaffin
 
tissue and others to pass on to form the ganglia of the coeliac
 
plexus, there is no way in the preparations at hand of distinguishing the two kinds. According to Souilé (’03), the penetration of the cortical portion of the suprarenal by the parasympathetic cells commences at the 19-mm. stage, which is of course
 
considerably older than the one dealt with here.
 
  
The cells of the gland itself are arranged in the form of a branching network penetrated with blood-vessels, and resembling the
+
The prevertebral sympathetic ganglion is now a very distinct mass of cells clearly difierentiated from the mesenchyme. Its relation to the ramus communicans is somewhat different from that described for the 9-mm. embryo. There is no longer any evidence of a branching in the ramus cominunicans at the level of the ganglion, the latter lying more directly in the path of the ramus. The ventral extension of the ramus (fig. 1) now passes directly ventralward from the ganglion (fig 2, 2).r.). Among the nerve fibers of the ventral extension of the ramus can be seen two kinds of cells: 1) those distinguished by a long narrow outline with nucleus of the same shape, which are probably sheath cells and, 2) large irregular cells with yellowish cytoplasm drawn out into processes and possessing rounded nuclei. The latter are undoubtedly migrating nerve cells (fig. 7). They are larger and difi'er otherwise in their appearance from the nerve cells found in the prevertebral ganglia (fig. 6), but are practically identical with the large nerve cells partially embedded in the ventral border of the suprarenal gland (figs. 2, 8). Comparing the pictures presented by figures 1 and 2 -it would seem that in the latter stage the nerve cells are in the actual process of migrating ventralward along the paths marked out by the nerve fibers, which alone are present in the former stage. If some of these migrating nerve cells are destined to enter the gland to form its chromaffin tissue and others to pass on to form the ganglia of the coeliac plexus, there is no way in the preparations at hand of distinguishing the two kinds. According to Souilé (’03), the penetration of the cortical portion of the suprarenal by the parasympathetic cells commences at the 19-mm. stage, which is of course considerably older than the one dealt with here.
zona reticularis of the adult organ. A distinct endothelium forms
 
the walls of the capillaries (fig. 9). The only other indication
 
of differentiation in the gland is the ingrowing of connective tissue trabeculae at the periphery.
 
  
 +
The cells of the gland itself are arranged in the form of a branching network penetrated with blood-vessels, and resembling the zona reticularis of the adult organ. A distinct endothelium forms the walls of the capillaries (fig. 9). The only other indication of differentiation in the gland is the ingrowing of connective tissue trabeculae at the periphery.
  
 
==Discussion==
 
==Discussion==
 +
His, Jun. (’91), described a preliminary nerve-fiber framework laid out in the form of rami communicantes, along which the sympathetic cells wander to form the ganglia. Streeter (Keibel and Mall, v. 2, p. 149) states that in human embryos the migrating cells can be recognized in advance of the loose strands of the tip of the growing nerve which extend through the mesenchyme toward the aorta, and that by the time a Well-defined nerve trunk is established, the sympathetic cells have already completed that part of their migration, and the cells then found on the nerve trunk are sheath cells only.
  
His, Jun. (’91), described a preliminary nerve-fiber framework
+
In the 9-mm. embryo under discussion the nerve fibers forming the ventral extension of the ramus communicans beyond the sympathetic ganglia (fig. 1) may have been preceded by amigration of ganglion cells through the mesenchyme, but my preparations do not show nerve cells either among the fibers or at their distal ends (fig. 3). On the other hand, in the 12-mm. embryo the cells found scattered along the course of the fibers are undoubtedly nerve cells rather than sheath cells. I am therefore inclined to believe that some at least of the ganglia migrate to their final location along paths formed of nerve fibers. The well-known work of Harrison (’06) which shows very conclusively that ganglion cells of amphibian larvae develop axis cylinders as outgrowths of the cell body, strengthens my conviction that the nerve fibers forming the pathway develop originally as outgrowths from cells located in the cord. Whether the nerve cells subsequently found among the nerve fibers come directly from the spinal ganglia or from the prevertebral ganglia is another question. I can only say that the migrating nerve cells are larger and differ in outline from those found in the prevertebral ganglia.
laid out in the form of rami communicantes, along which the sympathetic cells wander to form the ganglia. Streeter (Keibel and
 
Mall, v. 2, p. 149) states that in human embryos the migrating
 
cells can be recognized in advance of the loose strands of the tip
 
of the growing nerve which extend through the mesenchyme
 
toward the aorta, and that by the time a Well-defined nerve trunk
 
is established, the sympathetic cells have already completed that
 
part of their migration, and the cells then found on the nerve
 
trunk are sheath cells only.
 
  
In the 9-mm. embryo under discussion the nerve fibers forming
+
As has been noted above, Soulié (’03) states that the penetration of the parasympathetic cells into the cortical portion of the suprarenal gland commences at the 19-mm. stage. Zuckerkandl (Keibeland Mall, v. 2, p. 173) states that the elements of the migrating cell masses, which are entirely or for the most part chromaffin-forming cells, are sharply distinguished from the neighboring cortical cells by their smallness and intense stain. In my preparations one cannot say with certainty Whether such cells are present or not. Deeply staining cells occur bordering the nerve strands and the ganglionic masses, and these may represent the chromaflin cells, but if so they are sharply defined from the nerve cells and are more distinctly epithelial in character.
the ventral extension of the ramus communicans beyond the
 
sympathetic ganglia (fig. 1) may have been preceded by amigration of ganglion cells through the mesenchyme, but my preparations do not show nerve cells either among the fibers or at their
 
distal ends (fig. 3). On the other hand, in the 12-mm. embryo
 
the cells found scattered along the course of the fibers are undoubtedly nerve cells rather than sheath cells. I am therefore
 
inclined to believe that some at least of the ganglia migrate to
 
their final location along paths formed of nerve fibers. The
 
well-known work of Harrison (’06) which shows very conclusively
 
that ganglion cells of amphibian larvae develop axis cylinders as
 
outgrowths of the cell body, strengthens my conviction that the
 
nerve fibers forming the pathway develop originally as outgrowths from cells located in the cord. Whether the nerve cells
 
subsequently found among the nerve fibers come directly from
 
the spinal ganglia or from the prevertebral ganglia is another
 
question. I can only say that the migrating nerve cells are
 
larger and differ in outline from those found in the prevertebral
 
ganglia.
 
  
As has been noted above, Soulié (’03) states that the penetration of the parasympathetic cells into the cortical portion of the
+
Zuckerkandl (Keibel and Mall, V. 2, p. 171) found that the suprarenal glands are already vascularized in a 9-mm. embryo, whereas in my specimen of this age the only indication of vascularization is the occasional appearance in the suprarenal ridges of a cross-section of a blood capillary, which simply means that my embryo is younger than his, though both are the same length. In the 12-mm. embryo of my collection delicate endothelial capillaries form a very rich vascular network involving practically all of the gland except the cortical region. The central vein is not visible.
suprarenal gland commences at the 19-mm. stage. Zuckerkandl
 
(Keibeland Mall, v. 2, p. 173) states that the elements of the
 
migrating cell masses, which are entirely or for the most part
 
chromaffin-forming cells, are sharply distinguished from the neighboring cortical cells by their smallness and intense stain.
 
In my preparations one cannot say with certainty Whether such
 
cells are present or not. Deeply staining cells occur bordering
 
the nerve strands and the ganglionic masses, and these may
 
represent the chromaflin cells, but if so they are sharply
 
defined from the nerve cells and are more distinctly epithelial in
 
character.
 
  
Zuckerkandl (Keibel and Mall, V. 2, p. 171) found that the
+
Hoffmann (93) and others have shown that the primordial germ cells are distinct from the elements making up the germinal epithelium of the gonad and that they exist a long time before the appearance of the latter. More recently, Swift (’14) has traced the history of the primordial germ cells of the chick from their origin in a specialized region of the germ-wall entoderm just at the margin of the area pellucida. These cells are carried by their own movement and later by that of the blood to all parts of the embryo and Vascular area until in embryos of twentysix to twenty-nine somites they are found in the splanchnic mesoderm near the radix mesenterii. With the formation of the gonad they gradually pass to that organ. Fuss (’11) describes extraregional germ cells in a human embryo aged four Weeks. He finds them in the mesentery directly under the peritoneum, but not in the germinal epithelium. According to his description, they are large cells of rounded outline, with clear cytoplasm and distinct nucleus. The cells measure 19 to 20 p in diameter and the nuclei 12.75 p.
suprarenal glands are already vascularized in a 9-mm. embryo,
 
whereas in my specimen of this age the only indication of vascularization is the occasional appearance in the suprarenal ridges
 
of a cross-section of a blood capillary, which simply means that
 
my embryo is younger than his, though both are the same length.
 
In the 12-mm. embryo of my collection delicate endothelial
 
capillaries form a very rich vascular network involving practically all of the gland except the cortical region. The central
 
vein is not visible.
 
  
Hoffmann (93) and others have shown that the primordial
+
In my 9-mm. embryo, which is somewhat older than the one studied by Fuss, I have found cells resembling the one pictured by Fuss in his figure and corresponding in every way to those described in his text, except that the measurements I have made are somewhat less than his. Likewise I find the distribution of these primitive germ cells to be somewhat wider than he found, and in my preparations some of the cells have approached very close to the germinal epithelium (fig. 4). The 12—mm. embryo did not prove favorable for the study of these cells, so that I have no data on the range of distribution at this later stage. My observations on the 9-mm. embryo corroborate the statements of Fuss, which indicates that the germ cells of man like those of other vertebrates are characterized by period of migration in the early part of their history.
germ cells are distinct from the elements making up the germinal
 
epithelium of the gonad and that they exist a long time before
 
the appearance of the latter. More recently, Swift (’14) has
 
traced the history of the primordial germ cells of the chick from
 
their origin in a specialized region of the germ-wall entoderm
 
just at the margin of the area pellucida. These cells are carried
 
by their own movement and later by that of the blood to all
 
parts of the embryo and Vascular area until in embryos of twentysix to twenty-nine somites they are found in the splanchnic mesoderm near the radix mesenterii. With the formation of the gonad
 
they gradually pass to that organ. Fuss (’11) describes extraregional germ cells in a human embryo aged four Weeks. He
 
finds them in the mesentery directly under the peritoneum, but
 
not in the germinal epithelium. According to his description,
 
they are large cells of rounded outline, with clear cytoplasm and
 
distinct nucleus. The cells measure 19 to 20 p in diameter and
 
the nuclei 12.75 p.
 
 
 
In my 9-mm. embryo, which is somewhat older than the one
 
studied by Fuss, I have found cells resembling the one pictured by Fuss in his figure and corresponding in every way to those
 
described in his text, except that the measurements I have made
 
are somewhat less than his. Likewise I find the distribution of
 
these primitive germ cells to be somewhat wider than he found,
 
and in my preparations some of the cells have approached very
 
close to the germinal epithelium (fig. 4). The 12—mm. embryo
 
did not prove favorable for the study of these cells, so that I
 
have no data on the range of distribution at this later stage. My
 
observations on the 9-mm. embryo corroborate the statements of
 
Fuss, which indicates that the germ cells of man like those of
 
other vertebrates are characterized by period of migration in the
 
early part of their history.
 
  
 
==Summary==
 
==Summary==
Line 192: Line 55:
 
9-mm. embryo
 
9-mm. embryo
  
1. The suprarenal ridges extend from the pleuroperitoneal
+
# The suprarenal ridges extend from the pleuroperitoneal membranes posteriorly to the genital ridges.
membranes posteriorly to the genital ridges.
+
# The only indication of vascularization in the suprarenal tissue consists in an occasional cross-section of a capillary.
 
+
# In the suprarenal region the rami communicantes extend ventrally beyond the prevertebral sympathetic ganglia as nerve fibers apparently free of nerve cells.
2. The only indication of vascularization in the suprarenal
+
# Extraregional germ cells are found widely scattered in the neighborhood of the genital ridges.
tissue consists in an occasional cross-section of a capillary.
 
 
 
3. In the suprarenal region the rami communicantes extend
 
ventrally beyond the prevertebral sympathetic ganglia as nerve
 
fibers apparently free of nerve cells.
 
 
 
4. Extraregional germ cells are found widely scattered in the
 
neighborhood of the genital ridges.
 
  
 
12-mm. embryo
 
12-mm. embryo
  
1. The suprarenal glands are distinctly marked off from surrounding tissues.
+
# The suprarenal glands are distinctly marked off from surrounding tissues.
 
+
# The central region of the gland is in the form of a network, and is highly vascular, the blood-vessels having distinct endothelial walls.
2. The central region of the gland is in the form of a network,
+
# The median side of each gland is in close contact with the ventral prolongation of the ramus communicans and masses of ganglia cells. Some of the latter are found among the fibers of the distal part of the ramus along which they seem to be migrating to form the coeliac and visceral plexuses.
and is highly vascular, the blood-vessels having distinct endothelial walls.
 
  
3. The median side of each gland is in close contact with the
+
==Literature Cited==
ventral prolongation of the ramus communicans and masses of
 
ganglia cells. Some of the latter are found among the fibers of
 
the distal part of the ramus along which they seem to be migrating
 
to form the coeliac and visceral plexuses.
 
  
 +
FUSS, A. 1911 Ueber extraregionare Geschlechtszellen bei einen menschlichen Embryo Von vier Wochen. Anat. Anz., Bd. 39.
  
LITERATURE CITED
+
{{Ref-Harrison1906}}
  
FUSS, A. 1911 Ueber extraregionare Geschlechtszellen bei einen menschlichen
+
HIS, W., JUN. 1891 Die Entwicklung des Herznervensystems bei Wirbeltiere. Abh. d. math.-phys. Kl. d. Kgl. Sachs. Ges. (l. Wiss., Bd. 18.
Embryo Von vier Wochen. Anat. Anz., Bd. 39.
 
  
HARRISON, R. G. 1906 Further experiments on the development of peripheral
+
HOFFMANN, C. K. 1893 Etude sur le développement de l’appareil urogenital des oiseaux. Verh. d. Kgl. Akad. V. Wetenschoppen. Amsterdam, vol. 1.
nerves. Am. Jour. Anat., vol. 5.
 
  
HIS, W., JUN. 1891 Die Entwicklung des Herznervensystems bei Wirbeltiere.
+
{{Ref-KeibelMall1912}}
Abh. d. math.-phys. Kl. d. Kgl. Sachs. Ges. (l. Wiss., Bd. 18.
 
  
HOFFMANN, C. K. 1893 Etude sur le développement de l’appareil urogenital
+
MORRIS, J. T. 1909 A note on orange—G counterstaining suggesting a useful method in the management of embryonic tissue. Anat. Rec., vol. 3.
des oiseaux. Verh. d. Kgl. Akad. V. Wetenschoppen. Amsterdam,
 
vol. 1.
 
  
KEIBEL AND MALL 1912 Manual of human embryology. Philadelphia.
+
SOULIE, A. H. 1903 Recherches sur le développement des capsules surrénales. Journ. de l’Anat. et de la Physiol., vol. 39.
  
l\/IORRIS, J. T. 1909 A note on orange—G counterstaining suggesting a. useful
+
SWIFT, C. H. 1914 Origin and early history of the primordial germ cells in the chick. Am. Jour. Anat., vol. 15.
method in the management of embryonic tissue. Anat. Rec., vol. 3.
 
  
SOULIE, A. H. 1903 Recherches sur le développement des capsules surrénales.
+
==Description of Figures==
Journ. de l’Anat. et de la Physiol., vol. 39.
 
 
 
SVVIFT, C. H. 1914 Origin and early history of the primordial germ cells in the
 
chick. Am. Jour. Anat., vol. 15.
 
DESCRIPTION OF fiGURES
 
  
 
figs. 1 and 2 are camera drawings made at table level with Zeiss ocular 2- and
 
figs. 1 and 2 are camera drawings made at table level with Zeiss ocular 2- and
Line 250: Line 90:
 
2—mm. objective. All figures have been somewhat reduced in reproduction.
 
2—mm. objective. All figures have been somewhat reduced in reproduction.
  
ABBREVIATIONS USED IN fiGURES
+
===Abbreviations used in Figures===
  
 
b.v., blood-vessel p.c., pleuroperitoneal cushion
 
b.v., blood-vessel p.c., pleuroperitoneal cushion
Line 269: Line 109:
 
2 Transverse section through suprarenal region of 12—rnm. embryo.
 
2 Transverse section through suprarenal region of 12—rnm. embryo.
  
3 Enlargement of portion of ventral continuation of the ramus communicans
+
3 Enlargement of portion of ventral continuation of the ramus communicans of figure 1.
of figure 1.
 
  
4 Enlargement of portion of germinal ridge of figure 1, showing a primordial
+
4 Enlargement of portion of germinal ridge of figure 1, showing a primordial germ cell near the germinal epithelium.
germ cell near the germinal epithelium.
 
 
 
5 Enlargement of portion of subepithelial portion of the suprarenal ridge of
 
figure 1 also showing a primordial germ cell.
 
  
 +
5 Enlargement of portion of subepithelial portion of the suprarenal ridge of figure 1 also showing a primordial germ cell.
  
 
6 Enlargement of portion of the prevertebral ganglion of figure 2.
 
6 Enlargement of portion of the prevertebral ganglion of figure 2.
  
7 Enlargement of the ventral continuation of the ramus communicans of
+
7 Enlargement of the ventral continuation of the ramus communicans of figure 2.
figure 2.
 
  
8 Enlargement of the ganglionic mass embedded in the ventral border of the
+
8 Enlargement of the ganglionic mass embedded in the ventral border of the suprarenal gland of figure 2.
suprarenal gland of figure 2.
 
  
9 Enlargement of the central portion of the suprarenal gland of figure ‘2,
+
9 Enlargement of the central portion of the suprarenal gland of figure ‘2, showing the capillary structure.
showing the capillary structure.
 
  
  
 +
PLATE 2
  
PLATE 2
 
  
 +
{{Footer}}
  
 
[[Category:Adrenal]]
 
[[Category:Adrenal]]
 
[[Category:1920's]][[Category:Draft]]
 
[[Category:1920's]][[Category:Draft]]

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Wieman HL. Observations in connection with the early development of the human suprarenal gland. (1920) Anat. Rec. 19: 269-280.

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This historic 1920 paper by Wieman described human adrenal or suprarenal gland development.



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Observations in Connection with the Early Development of the Human Suprarenal Gland

H. L. Wieman

Zoological Laboratory, University of Cincinnati

Two Plates (Nine figures)

Introduction

The purpose of this contribution is to call attention to certain details in connection with the development of the human suprarenal gland, because they happen to be clearly illustrated in the material at hand. This material consists of two human embryos, nos. 1 and 4 of the author’s collection, which were obtained some years ago, freshly killed and fixed, from Drs. H. L. VVoodward and Charles Goosmann, of Cincinnati. No. 1 had been killed in a bichromate-acetic mixture, and no. 4 in Bouin’s solution. The former measured 9 mm., crown—breech, and the latter 12 mm. Both measurements were made in 85 per cent alcohol after fixation, so that the sizes of the living embryos were somewhat larger than these figures. The embryos were embedded in paraffin, cut into serial sections of 10 [1, thickness, and stained on the slide with Delafield haematoxylin and orange G according to the method of Morris (’09). Mitotic figures abound in both embryos, but the fixation is somewhat better in no. 1.


Observations

Embryo no. 1, .9 mm. The suprarenal glands are not recognizable as distinct organs, but consist of a thickening in the mesenchyme on either side of the root of the mesentery, forming a pair of broad ridges projecting into the body cavity from the dorsal body wall. Anteriorly these suprarenal ridges are continuous with the dorsal portions of the pleuroperitoneal membranes, while posteriorly they blend with the genital ridges. Laterally each is separated from the mesonephros by a distinct groove (fig. 1). Occasionally a transverse section of a blood capillary can be seen in the center of each ridge, the beginning no doubt of the central vein of the adult organ.

The ridges are made up of mesenchyme which shows no evidence of differentiation. The ramus communicans divides into two at the level of the sympathetic ganglion where one of the branches terminates, the other passing ventralward into the mesenchyme. The cells composing the sympathetic ganglia stain more deeply than the surrounding cells, but beyond this they show no appreciable differentiation. The ganglia lie on either side of the aorta somewhat dorsal to it. The nerve strands (axis cylinders?) are easily distinguished and followed, owing to the fact that they stain readily with orange G. The ventral branch of the ramus communicans, which is really the direct ventral continuation of the latter, loses itself in the mesenchyme of the suprarenal region. It seems to be unaccompanied by nerve cells (fig. 3). From the picture one gets the impression that the nerve fibers are pushing their way through the mesenchyme, blazing, as it were, a track along which the ganglion cells are to follow later.

In the posterior part of the suprarenal ridge, the genital ridge, the subepithelial region of the mesonephros, and in the mesentery, one finds large cells with clear cytoplasm and standing out distinctly from all cells (fig. 3, 4, 5). These 1 take to be germ cells (primary genital cells). Their wide extraregional distribution indicates that these cells undergo a rather extensive migration before reaching the germinal epithelium. This of course is in keeping with what is known about the early development of germ cells in other vertebrates.

Embryo no. 4, 12 mm. The suprarenal glands in this specimen are distinctly marked off from the surrounding tissues (fig. 2). Each one lies with its dorsal surface against the pleuroperitoneal cushion, while in close contact with its median side are bundles of nerve fibers and -ganglionic clumps. Near the ventral border, as shown in figure 2, some nervous tissue is pushed into the substance of the gland, but only to a very slight extent, definite immigration, according to other authors, not taking place until a much later period.

The prevertebral sympathetic ganglion is now a very distinct mass of cells clearly difierentiated from the mesenchyme. Its relation to the ramus communicans is somewhat different from that described for the 9-mm. embryo. There is no longer any evidence of a branching in the ramus cominunicans at the level of the ganglion, the latter lying more directly in the path of the ramus. The ventral extension of the ramus (fig. 1) now passes directly ventralward from the ganglion (fig 2, 2).r.). Among the nerve fibers of the ventral extension of the ramus can be seen two kinds of cells: 1) those distinguished by a long narrow outline with nucleus of the same shape, which are probably sheath cells and, 2) large irregular cells with yellowish cytoplasm drawn out into processes and possessing rounded nuclei. The latter are undoubtedly migrating nerve cells (fig. 7). They are larger and difi'er otherwise in their appearance from the nerve cells found in the prevertebral ganglia (fig. 6), but are practically identical with the large nerve cells partially embedded in the ventral border of the suprarenal gland (figs. 2, 8). Comparing the pictures presented by figures 1 and 2 -it would seem that in the latter stage the nerve cells are in the actual process of migrating ventralward along the paths marked out by the nerve fibers, which alone are present in the former stage. If some of these migrating nerve cells are destined to enter the gland to form its chromaffin tissue and others to pass on to form the ganglia of the coeliac plexus, there is no way in the preparations at hand of distinguishing the two kinds. According to Souilé (’03), the penetration of the cortical portion of the suprarenal by the parasympathetic cells commences at the 19-mm. stage, which is of course considerably older than the one dealt with here.

The cells of the gland itself are arranged in the form of a branching network penetrated with blood-vessels, and resembling the zona reticularis of the adult organ. A distinct endothelium forms the walls of the capillaries (fig. 9). The only other indication of differentiation in the gland is the ingrowing of connective tissue trabeculae at the periphery.

Discussion

His, Jun. (’91), described a preliminary nerve-fiber framework laid out in the form of rami communicantes, along which the sympathetic cells wander to form the ganglia. Streeter (Keibel and Mall, v. 2, p. 149) states that in human embryos the migrating cells can be recognized in advance of the loose strands of the tip of the growing nerve which extend through the mesenchyme toward the aorta, and that by the time a Well-defined nerve trunk is established, the sympathetic cells have already completed that part of their migration, and the cells then found on the nerve trunk are sheath cells only.

In the 9-mm. embryo under discussion the nerve fibers forming the ventral extension of the ramus communicans beyond the sympathetic ganglia (fig. 1) may have been preceded by amigration of ganglion cells through the mesenchyme, but my preparations do not show nerve cells either among the fibers or at their distal ends (fig. 3). On the other hand, in the 12-mm. embryo the cells found scattered along the course of the fibers are undoubtedly nerve cells rather than sheath cells. I am therefore inclined to believe that some at least of the ganglia migrate to their final location along paths formed of nerve fibers. The well-known work of Harrison (’06) which shows very conclusively that ganglion cells of amphibian larvae develop axis cylinders as outgrowths of the cell body, strengthens my conviction that the nerve fibers forming the pathway develop originally as outgrowths from cells located in the cord. Whether the nerve cells subsequently found among the nerve fibers come directly from the spinal ganglia or from the prevertebral ganglia is another question. I can only say that the migrating nerve cells are larger and differ in outline from those found in the prevertebral ganglia.

As has been noted above, Soulié (’03) states that the penetration of the parasympathetic cells into the cortical portion of the suprarenal gland commences at the 19-mm. stage. Zuckerkandl (Keibeland Mall, v. 2, p. 173) states that the elements of the migrating cell masses, which are entirely or for the most part chromaffin-forming cells, are sharply distinguished from the neighboring cortical cells by their smallness and intense stain. In my preparations one cannot say with certainty Whether such cells are present or not. Deeply staining cells occur bordering the nerve strands and the ganglionic masses, and these may represent the chromaflin cells, but if so they are sharply defined from the nerve cells and are more distinctly epithelial in character.

Zuckerkandl (Keibel and Mall, V. 2, p. 171) found that the suprarenal glands are already vascularized in a 9-mm. embryo, whereas in my specimen of this age the only indication of vascularization is the occasional appearance in the suprarenal ridges of a cross-section of a blood capillary, which simply means that my embryo is younger than his, though both are the same length. In the 12-mm. embryo of my collection delicate endothelial capillaries form a very rich vascular network involving practically all of the gland except the cortical region. The central vein is not visible.

Hoffmann (93) and others have shown that the primordial germ cells are distinct from the elements making up the germinal epithelium of the gonad and that they exist a long time before the appearance of the latter. More recently, Swift (’14) has traced the history of the primordial germ cells of the chick from their origin in a specialized region of the germ-wall entoderm just at the margin of the area pellucida. These cells are carried by their own movement and later by that of the blood to all parts of the embryo and Vascular area until in embryos of twentysix to twenty-nine somites they are found in the splanchnic mesoderm near the radix mesenterii. With the formation of the gonad they gradually pass to that organ. Fuss (’11) describes extraregional germ cells in a human embryo aged four Weeks. He finds them in the mesentery directly under the peritoneum, but not in the germinal epithelium. According to his description, they are large cells of rounded outline, with clear cytoplasm and distinct nucleus. The cells measure 19 to 20 p in diameter and the nuclei 12.75 p.

In my 9-mm. embryo, which is somewhat older than the one studied by Fuss, I have found cells resembling the one pictured by Fuss in his figure and corresponding in every way to those described in his text, except that the measurements I have made are somewhat less than his. Likewise I find the distribution of these primitive germ cells to be somewhat wider than he found, and in my preparations some of the cells have approached very close to the germinal epithelium (fig. 4). The 12—mm. embryo did not prove favorable for the study of these cells, so that I have no data on the range of distribution at this later stage. My observations on the 9-mm. embryo corroborate the statements of Fuss, which indicates that the germ cells of man like those of other vertebrates are characterized by period of migration in the early part of their history.

Summary

9-mm. embryo

  1. The suprarenal ridges extend from the pleuroperitoneal membranes posteriorly to the genital ridges.
  2. The only indication of vascularization in the suprarenal tissue consists in an occasional cross-section of a capillary.
  3. In the suprarenal region the rami communicantes extend ventrally beyond the prevertebral sympathetic ganglia as nerve fibers apparently free of nerve cells.
  4. Extraregional germ cells are found widely scattered in the neighborhood of the genital ridges.

12-mm. embryo

  1. The suprarenal glands are distinctly marked off from surrounding tissues.
  2. The central region of the gland is in the form of a network, and is highly vascular, the blood-vessels having distinct endothelial walls.
  3. The median side of each gland is in close contact with the ventral prolongation of the ramus communicans and masses of ganglia cells. Some of the latter are found among the fibers of the distal part of the ramus along which they seem to be migrating to form the coeliac and visceral plexuses.

Literature Cited

FUSS, A. 1911 Ueber extraregionare Geschlechtszellen bei einen menschlichen Embryo Von vier Wochen. Anat. Anz., Bd. 39.

Harrison RG. Further experiments on the development of peripheral nerves. (1906) Amer. J Anat. 5:

HIS, W., JUN. 1891 Die Entwicklung des Herznervensystems bei Wirbeltiere. Abh. d. math.-phys. Kl. d. Kgl. Sachs. Ges. (l. Wiss., Bd. 18.

HOFFMANN, C. K. 1893 Etude sur le développement de l’appareil urogenital des oiseaux. Verh. d. Kgl. Akad. V. Wetenschoppen. Amsterdam, vol. 1.

Keibel F. and Mall FP. Manual of Human Embryology II. (1912) J. B. Lippincott Company, Philadelphia.

MORRIS, J. T. 1909 A note on orange—G counterstaining suggesting a useful method in the management of embryonic tissue. Anat. Rec., vol. 3.

SOULIE, A. H. 1903 Recherches sur le développement des capsules surrénales. Journ. de l’Anat. et de la Physiol., vol. 39.

SWIFT, C. H. 1914 Origin and early history of the primordial germ cells in the chick. Am. Jour. Anat., vol. 15.

Description of Figures

figs. 1 and 2 are camera drawings made at table level with Zeiss ocular 2- and 16-mm. objective. The remaining figures were made with Zeiss ocular 4- and 2—mm. objective. All figures have been somewhat reduced in reproduction.

Abbreviations used in Figures

b.v., blood-vessel p.c., pleuroperitoneal cushion

e.c., endothelial cell s.g., prevertebral sympathetic gang.c., primordial germ cell glion

g.r., germinal ridge s.r., suprarenal ridge

n.c., nerve cell v.r., ventral continuation of the ramus n. f ., nerve fiber communicans

PLATE 1

EXPLANATION OF fiGURES

1 Transverse section through suprarenal region of 9-mm. embryo.

2 Transverse section through suprarenal region of 12—rnm. embryo.

3 Enlargement of portion of ventral continuation of the ramus communicans of figure 1.

4 Enlargement of portion of germinal ridge of figure 1, showing a primordial germ cell near the germinal epithelium.

5 Enlargement of portion of subepithelial portion of the suprarenal ridge of figure 1 also showing a primordial germ cell.

6 Enlargement of portion of the prevertebral ganglion of figure 2.

7 Enlargement of the ventral continuation of the ramus communicans of figure 2.

8 Enlargement of the ganglionic mass embedded in the ventral border of the suprarenal gland of figure 2.

9 Enlargement of the central portion of the suprarenal gland of figure ‘2, showing the capillary structure.


PLATE 2



Cite this page: Hill, M.A. (2019, October 22) Embryology Paper - Observations in connection with the early development of the human suprarenal gland. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Paper_-_Observations_in_connection_with_the_early_development_of_the_human_suprarenal_gland

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