Paper - The development of the adrenal gland in man: Difference between revisions

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==Definitions==
==Definitions==
The following terms are defined to clarify the presentation:
The following terms are defined to clarify the presentation:


Fetal cortex: The fetal adrenal cortex is composed of the same cell types as the adrenal cortex of the adult human, but the proportions and arrangements differ.
Fetal cortex: The fetal adrenal cortex is composed of the same cell types as the adrenal cortex of the adult human, but the proportions and arrangements differ.


Medullary cells: The cells of neural origin which take up a location in the central portion of the gland during development. The process of localization begins in the embryonic period and continues until childhood.
Medullary cells: The cells of neural origin which take up a location in the central portion of the gland during development. The process of localization begins in the embryonic period and continues until childhood.


Cell types: Cell types in the cortex are differentiated by the structure of their nuclei; they are numbered according to the order of their appearance in the primordium, as C-I (fig. 38, pl. 8), C-II (fig. 39, pl. 8), and C-III (fig. 40, pl. 8). As will be made clear, three distinct cell types may be recognized in the development of the medulla; these cells, designated M-I, M-II, and M-III, are described beginning on page 202.
Cell types: Cell types in the cortex are differentiated by the structure of their nuclei; they are numbered according to the order of their appearance in the primordium, as C-I (fig. 38, pl. 8), C-II (fig. 39, pl. 8), and C-III (fig. 40, pl. 8). As will be made clear, three distinct cell types may be recognized in the development of the medulla; these cells, designated M-I, M-II, and M-III, are described beginning on page 202.


Celomic epithelial cells: The cells of the adrenal cortex which have their origin in the celomic epithelium. They are of two types: Type C-I (fig. 38, pl. 8), the first cells to enter the primordium, are large polyhedral cells, each of which contains a large round nucleus with little chromatin, a large round nucleolus, and a spherical, refractile inclusion. In a later stage, type C-III cells (Hg. 40, pl. 8) begin to emerge from the celomic epithelium, together with type C-I. Type C-III cells are smaller than type C-1, and each contains a smaller round nucleus which stains more deeply with hematoxylin. In the nucleus. there is a large amount of chromatin, which tends to aggregate into four to five major clumps and one or two globules or vacuoles.
Celomic epithelial cells: The cells of the adrenal cortex which have their origin in the celomic epithelium. They are of two types: Type C-I (fig. 38, pl. 8), the first cells to enter the primordium, are large polyhedral cells, each of which contains a large round nucleus with little chromatin, a large round nucleolus, and a spherical, refractile inclusion. In a later stage, type C-III cells (Hg. 40, pl. 8) begin to emerge from the celomic epithelium, together with type C-I. Type C-III cells are smaller than type C-1, and each contains a smaller round nucleus which stains more deeply with hematoxylin. In the nucleus. there is a large amount of chromatin, which tends to aggregate into four to five major clumps and one or two globules or vacuoles.


Cells of mesonephric origin: These cells arise from Bowman's capsule of the mesoncphros. They are designated as type C-II cells (fig. 39, pl. 8), and are first seen streaming from the glomerular capsule across the ventral portion of the early primordium (figs. 14, 15, pl. 2). The type C-II cells and their successors form most of the adrenal capsule, and, during horizon xxiii, are diverted inward by the arteriolar capillaries. Thereafter, they are found in the cords of cells within the cortex. Their nucleus is round, and stains a little more deeply than does that of type C-I. The chromatin is distributed in fine strands in a spider-like manner just beneath the nuclear membranes, the nucleolus corresponding to the body of the spider. The center of the nucleus is occupied by one or more large globules or vacuoles.
Cells of mesonephric origin: These cells arise from Bowman's capsule of the mesoncphros. They are designated as type C-II cells (fig. 39, pl. 8), and are first seen streaming from the glomerular capsule across the ventral portion of the early primordium (figs. 14, 15, pl. 2). The type C-II cells and their successors form most of the adrenal capsule, and, during horizon xxiii, are diverted inward by the arteriolar capillaries. Thereafter, they are found in the cords of cells within the cortex. Their nucleus is round, and stains a little more deeply than does that of type C-I. The chromatin is distributed in fine strands in a spider-like manner just beneath the nuclear membranes, the nucleolus corresponding to the body of the spider. The center of the nucleus is occupied by one or more large globules or vacuoles.


Immigration and migration: The passage of cells from one region to another; the addition of cells by movement from an adjacent region. Ameboid movement is not implied.
Immigration and migration: The passage of cells from one region to another; the addition of cells by movement from an adjacent region. Ameboid movement is not implied.


Invasion: The entrance of cells after destruction or disruption of pre-existing tissue, and the assumption of position without rcference to former relations.
Invasion: The entrance of cells after destruction or disruption of pre-existing tissue, and the assumption of position without rcference to former relations.


Staging of embryos: The dating of the embryos follows the horizons described and graphed by Streeter (1951). This method of seriation affords a much better concept of the progress of development than does measurement of the length. Frequently, the organogenesis of the adrenal gland was found to be in a much further advanced stage in an embryo with a shorter crown-rump length than in a longer one, particularly in the earlier stages when the dorsal curvature varies so remarkably among embryos in the same horizon. This finding speaks well for the method, since Streeter did not use the adrenal among his criteria for staging. The age in weeks is the
Staging of embryos: The dating of the embryos follows the horizons described and graphed by Streeter (1951). This method of seriation affords a much better concept of the progress of development than does measurement of the length. Frequently, the organogenesis of the adrenal gland was found to be in a much further advanced stage in an embryo with a shorter crown-rump length than in a longer one, particularly in the earlier stages when the dorsal curvature varies so remarkably among embryos in the same horizon. This finding speaks well for the method, since Streeter did not use the adrenal among his criteria for staging. The age in weeks is the estimated conception age. To obtain menstrual age, two weeks must be added. There are enough stages to pro- vicle a consecutive account of the organogenesis of the adrenal.
 
estimated conception age. To obtain menstrual age, two weeks must be added. There are enough stages to pro- vicle a consecutive account of the organogenesis of the adrenal.


==Materials and Methods==
==Materials and Methods==

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Crowder RE. The development of the adrenal gland in man, with special reference to origin and ultimate location of cell types and evidence in favor of the "cell migration" theory. (1957) Contrib. Embryol., Carnegie Inst. Wash. 36, 193-210.

Online Editor Note 
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This historic 1957 paper by Crowder is a description of the development of the human adrenal gland.



Week: 1 2 3 4 5 6 7 8
Carnegie stage: 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23


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Endocrine Links: Introduction | BGD Lecture | Science Lecture | Lecture Movie | pineal | hypothalamus‎ | pituitary | thyroid | parathyroid | thymus | pancreas | adrenal | endocrine gonad‎ | endocrine placenta | other tissues | Stage 22 | endocrine abnormalities | Hormones | Category:Endocrine
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1903 Islets of Langerhans | 1903 Pig Adrenal | 1904 interstitial Cells | 1908 Pancreas Different Species | 1908 Pituitary | 1908 Pituitary histology | 1911 Rathke's pouch | 1912 Suprarenal Bodies | 1914 Suprarenal Organs | 1915 Pharynx | 1916 Thyroid | 1918 Rabbit Hypophysis | 1920 Adrenal | 1935 Mammalian Hypophysis | 1926 Human Hypophysis | 1927 Adrenal | 1927 Hypophyseal fossa | 1930 Adrenal | 1932 Pineal Gland and Cysts | 1935 Hypophysis | 1935 Pineal | 1937 Pineal | 1935 Parathyroid | 1940 Adrenal | 1941 Thyroid | 1950 Thyroid Parathyroid Thymus | 1957 Adrenal


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Historic Embryology - Neural Crest  
1879 Olfactory Organ | 1905 Cranial and Spinal Nerves | 1908 10 mm Peripheral | 1910 Mammal Sympathetic | 1920 Human Sympathetic | 1928 Cranial ganglia | 1939 10 Somite Embryo | 1942 Origin | 1957 Adrenal
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The Development Of The Adrenal Gland In Man, With Special Reference To Origin And Ultimate Location Of Cell Types And Evidence In Favor Of The “Cell Migration” Theory

by Roy E. Crowder

With 8 plates and 7 text figures.

  • The ideas herein expressed are those of the author and do not reflect those of the United States Navy.

Introduction

A voluminous literature has accumulated on the development of the adrenal gland. In 1902, Iosef \Viesel apologized for adding one more article. Others, including the present author, have not been so restrained.

In a study of the sources of certain of the adrenal hormones, a review of the literature on the embryology and histology of the gland was undertaken. In spite of the number of articles published on the subject, including extensive reviews (Parkes, I945; Lanman, 1953), there was no consecutive life history of the source and ultimate distribution of the various cell types found in the gland. and, therefore, it was not possible to associate the synthesis of the adrenal hormones with specific cell types.


The wealth of closely seriated, well preserved, and adequately stained material available in the Carnegie Embryological Collection, in Baltimore, Maryland, made a complete histologic and embryologic study possible.


The study was begun with the idea of determining the types of cells in the adrenal cortex. It was soon found that, to do this adequately, the whole embryologic history must be reviewed. The results of this undertaking are set forth herein. It is hoped not only that they will assist in an understanding of the sequence of events in the histogenesis of the adrenal cortex and clarify a few of the points of controversy, but also that the descriptions of cell types in the cortex may be of value in determining the oligoplasia or hypcrplasia of certain of these cell types in relation to many clinical entities.

Definitions

The following terms are defined to clarify the presentation:


Fetal cortex: The fetal adrenal cortex is composed of the same cell types as the adrenal cortex of the adult human, but the proportions and arrangements differ.


Medullary cells: The cells of neural origin which take up a location in the central portion of the gland during development. The process of localization begins in the embryonic period and continues until childhood.


Cell types: Cell types in the cortex are differentiated by the structure of their nuclei; they are numbered according to the order of their appearance in the primordium, as C-I (fig. 38, pl. 8), C-II (fig. 39, pl. 8), and C-III (fig. 40, pl. 8). As will be made clear, three distinct cell types may be recognized in the development of the medulla; these cells, designated M-I, M-II, and M-III, are described beginning on page 202.


Celomic epithelial cells: The cells of the adrenal cortex which have their origin in the celomic epithelium. They are of two types: Type C-I (fig. 38, pl. 8), the first cells to enter the primordium, are large polyhedral cells, each of which contains a large round nucleus with little chromatin, a large round nucleolus, and a spherical, refractile inclusion. In a later stage, type C-III cells (Hg. 40, pl. 8) begin to emerge from the celomic epithelium, together with type C-I. Type C-III cells are smaller than type C-1, and each contains a smaller round nucleus which stains more deeply with hematoxylin. In the nucleus. there is a large amount of chromatin, which tends to aggregate into four to five major clumps and one or two globules or vacuoles.


Cells of mesonephric origin: These cells arise from Bowman's capsule of the mesoncphros. They are designated as type C-II cells (fig. 39, pl. 8), and are first seen streaming from the glomerular capsule across the ventral portion of the early primordium (figs. 14, 15, pl. 2). The type C-II cells and their successors form most of the adrenal capsule, and, during horizon xxiii, are diverted inward by the arteriolar capillaries. Thereafter, they are found in the cords of cells within the cortex. Their nucleus is round, and stains a little more deeply than does that of type C-I. The chromatin is distributed in fine strands in a spider-like manner just beneath the nuclear membranes, the nucleolus corresponding to the body of the spider. The center of the nucleus is occupied by one or more large globules or vacuoles.


Immigration and migration: The passage of cells from one region to another; the addition of cells by movement from an adjacent region. Ameboid movement is not implied.


Invasion: The entrance of cells after destruction or disruption of pre-existing tissue, and the assumption of position without rcference to former relations.

Staging of embryos: The dating of the embryos follows the horizons described and graphed by Streeter (1951). This method of seriation affords a much better concept of the progress of development than does measurement of the length. Frequently, the organogenesis of the adrenal gland was found to be in a much further advanced stage in an embryo with a shorter crown-rump length than in a longer one, particularly in the earlier stages when the dorsal curvature varies so remarkably among embryos in the same horizon. This finding speaks well for the method, since Streeter did not use the adrenal among his criteria for staging. The age in weeks is the estimated conception age. To obtain menstrual age, two weeks must be added. There are enough stages to pro- vicle a consecutive account of the organogenesis of the adrenal.

Materials and Methods

The following embryos of the Carnegie Collection were selected for the study of the embryonic and early fetal development of the adrenal gland.

(table to be formatted)

Crown-Rump Length (mm.)

saw

Horizon xnr

836 . . . . . . 4.0 Trans. Exc. A1. coch. 7433 . . . . . . 5.2 Frontal Exc. (Stain - Haematoxylin Eosin) 8066 . . . . . . 5.3 Trans. Exc. (Stain - Haematoxylin Eosin) 8119 . . . . . . 5.3 Trans. Exc. (Stain - Haematoxylin Eosin) Homzo.-' xtv 3805 . . . . . . 5.9 Trans. Exc. (Stain - Haematoxylin Eosin)

6502 . . . . . . 6.7 Trans. Exc. (Stain - Haematoxylin Eosin)

7324 . . . . . . 6.6 Trans. Good (Stain - Haematoxylin Eosin)

7829 . . . . . . 7.0 Trans. Exc. (Stain - Haematoxylin Eosin)

7370 . . . . . . 7.2 Trans. Exc. (Stain - Haematoxylin Eosin)

8314 . . . . . . 8.2 Trans. Exc. Azan 1-lomzow xv 2. . . . . . 7.0 Trans. Good Al. carm. 721 . . . . . . 9.0 Trans. lixc.

3385 . . . . . . 8.3 Trans. Exc. (Stain - Haematoxylin Eosin) or G.

3441 . . . . .. 8.0 Sag. Exc. Al. coch.

3512 . . . . . . S 5 Trans. Exc. Al. coch.

6504 . . . . . . 7 5 Sag. Exc. A1. coch.

6508 . . . . . . 7.3 Trans. Good Al. coch. 6595. . . . , 3.0 Sag. Good I-1.-17.. 7199 . . . . . , 8.-'1 Frontal Good Al. coch. phloxine 7364 . . . . . 9 5 Frontal Poor 1-1.-17.. 8929 . . . . . , 6 3 Frontal Exc. Azan 8997 . . . . . . 9 0 Trans. Exc. Azan 9140 . . . . . . 7 0 Trans. Exc. Azan I-Ioruzox xvt

792 . . . . . . 8 0 Trans. Good A1. coch.

6511 . . . . . . 8 1 Sag. Good Al. coch. iron H.

6512 . . . . .. 70 Trans. Exc. A1. coch.

6517 . . . . . . 10 5 Trans. Exc. Al. coch.

7115 . . . . .. 9 7 Frontal Exc. H. and pltloxinc

7804 . . . . . . 9 5 Trans. Good (Stain - Haematoxylin Eosin)

7897 . . . . . . 12 2 Trans. Good (Stain - Haematoxylin Eosin)

8773 . . . . . . ll 0 Frontal Exc. Azan l-lot-:1zo.\' xvu 940 . . . . . . 14 0 Trans. Good (Stain - Haematoxylin Eosin) or G.

5893 . . . . . . 13 2 Trans. Good Al. coch.

6253 . . . . . . 1-10 Trans. Good (Stain - Haematoxylin Eosin)

6519 , . . . . . 10 3 Sag. Exc. Al. coch.

6520 . . . . . . 14 2 Trans. Exc. Al. coch.

6742 . . . . . . 11 0 Trans. Good I-1.-pltloxine

6758 . , . . . . 12 8 Trans. Good 1-1.-phloxine

8101 . . . . .. 13.0 Trans. Exc. (Stain - Haematoxylin Eosin)

811$ . . . . .. 12.6 Frontal Exc. (Stain - Haematoxylin Eosin)

8789 . . . . . . ll 7 Sag. Exc. Azan

3998 . . . . . . 11 0 Frontal Exc. Azan

Crown- _

(mm.)

Horizon xvrn

-192 . . . , . . 16.3 Frontal Exc. Al. coch.

1909 . . . . . . 14.6 Frontal Good .'-\l. coch. or G.

4430 . . . . .. 14.0 Trans. Exc. A1. coch. or G.

6524 . . . . . . 11.7 Trans. Exc. A1. coch.

6525 . . . . .. 13.8 Sag. Exc. Al. coch.

6527 . . . . .. 14.4 Trans. Exc. Al. coch.

6528 . . . . .. 13.4 Frontal Exc. A1. coch.

7707 . . . . .. 14 5 Trans. F.xc. 1-1.—F.. phloxine

8172 . . . . . . 16 5 Trans. Exc. (Stain - Haematoxylin Eosin)

S312 . . . . .. 12.2 Trans. F..\:c. I-1.717..

9247 . . . . .. 15.0 Sag. Exc. Azan

Horizon xxx

4-105 . . . . . . 15 5 Trans. Good Coch. l\1allor_\-' 4501 . . . . .. 180 Trans. F..\'c. Coclt. or G. 5609. . 18 0 Frontal F..\:c. Al. coch.

6150 . . . . . . 17.0 Trans. Good

632-l . . . . .. 18.5 Sag. Good (Stain - Haematoxylin Eosin)

S092. . . . . . 16 3 Trans. lixc. I-1.-E. phlo.\;ine 9113 . . . . . . 18 5 Trans. Exc. Azan

Horizon xx 966.. . 23.0 Frontal F.xc. A1. coch.

5537. . . . .. 22.0 Trans. Fxc. Al. coch. 6202... . .. 21.0 Sag. F.xc. (Stain - Haematoxylin Eosin)

7274 . . . . .. 18 3 Trans. Exc. (Stain - Haematoxylin Eosin) phloxinc 7906 . , . . .. 19.5 Frontal F..\:c. (Stain - Haematoxylin Eosin)

S226 . . . . .. 18.0 Sag. Exc. Azan

Horizon xxx

5.596. . 21.5 Sag. Good I-1.-Ii.

725-1.. . 22.5 Trans. Exc.

7392.. . 22.7 Trans. Exc. (Stain - Haematoxylin Eosin)

Z864. . 24 0 Frontal F.:~:c. (Stain - Haematoxylin Eosin)

S553 . _ . _ . , 22.0 Trans. Exc. (Stain - Haematoxylin Eosin)

Horizon xxn

1458. . . . . , 27.5 Sag. Exc. or G 430-1. . 25 0 Trans. Good (Stain - Haematoxylin Eosin)

4638. , . 23.4 Trans. Exc. Al. coch

6701 . . . 24.0 Frontal Poor I-I.-E.

6832. . 25.8 Frontal Exc. I-1..F_.

339-1. . . 25 3 Trans. lixc. I1.-I’.

Horizon mm

4205. . . . . . 29.5 Trans. Good Al coch

No. 4289. . . . . . 32.2 Trans. Good Al coch

4570 . . . . .. 30.7 Trans. Exc. (Stain - Haematoxylin Eosin)

5-122. . . . . . 27.0 Sag. Good (Stain - Haematoxylin Eosin)

5621.-\ . . . . 27.5 Trans. Good

6573. . . . . . 31.5 Trans. Good (Stain - Haematoxylin Eosin)

6719 . . . . .. 30.1 Trans. Good Chrom.alum hem. 7425 . . . . .. 27.0 Frontal Exc. Cl1I‘0111.:1lUI11l1CI]'l. 9226. . . . . . 30.1 Trans. Exc. Azan




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