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| {{Header}}
| | #REDIRECT [[Paper - The development of the adrenal gland in man (1957)]] |
| {{Ref-Crowder1957}}
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| {{Historic Disclaimer}}
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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=
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| by Roy E. Crowder
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| With 8 plates and 7 text figures.
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| * The ideas herein expressed are those of the author and do not reflect those of the United States Navy.
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| ==Introduction==
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| A voluminous literature has accumulated on the development of the adrenal gland. In 1902, Iosef \Viesel
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| apologized for adding one more article. Others, including the present author, have not been so restrained.
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| In a study of the sources of certain of the adrenal
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| hormones, a review of the literature on the embryology
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| and histology of the gland was undertaken. In spite
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| of the number of articles published on the subject, including extensive reviews (Parkes, I945; Lanman, 1953),
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| there was no consecutive life history of the source and
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| ultimate distribution of the various cell types found in
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| the gland. and, therefore, it was not possible to associate
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| the synthesis of the adrenal hormones with specific cell
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| types.
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| The wealth of closely seriated, well preserved, and adequately stained material available in the [[Carnegie Collection|Carnegie Embryological Collection]], in Baltimore, Maryland, made a complete histologic and embryologic study possible.
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| The study was begun with the idea of determining the
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| types of cells in the adrenal cortex. It was soon found
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| that, to do this adequately, the whole embryologic history
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| must be reviewed. The results of this undertaking are
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| set forth herein. It is hoped not only that they will assist
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| in an understanding of the sequence of events in the
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| histogenesis of the adrenal cortex and clarify a few of
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| the points of controversy, but also that the descriptions
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| of cell types in the cortex may be of value in determining
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| the oligoplasia or hypcrplasia of certain of these cell types
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| in relation to many clinical entities.
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| ==Definitions==
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| The following terms are defined to clarify the presentation:
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| Fetal cortex: The fetal adrenal cortex is composed of
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| the same cell types as the adrenal cortex of the adult
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| human, but the proportions and arrangements differ.
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| Medullary cells: The cells of neural origin which take
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| up a location in the central portion of the gland during
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| development. The process of localization begins in the
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| embryonic period and continues until childhood.
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| Cell types: Cell types in the cortex are differentiated
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| 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
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| C-III (fig. 40, pl. 8). As will be made clear, three distinct
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| cell types may be recognized in the development of the
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| medulla; these cells, designated M-I, M-II, and M-III, are
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| described beginning on page 202.
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| Celomic epithelial cells: The cells of the adrenal cortex
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| which have their origin in the celomic epithelium. They
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| are of two types: Type C-I (fig. 38, pl. 8), the first cells
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| to enter the primordium, are large polyhedral cells, each
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| of which contains a large round nucleus with little
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| chromatin, a large round nucleolus, and a spherical, refractile inclusion. In a later stage, type C-III cells (Hg. 40,
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| pl. 8) begin to emerge from the celomic epithelium,
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| together with type C-I. Type C-III cells are smaller than
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| type C-1, and each contains a smaller round nucleus
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| which stains more deeply with hematoxylin. In the
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| 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.
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| Cells of mesonephric origin: These cells arise from
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| Bowman's capsule of the mesoncphros. They are designated as type C-II cells (fig. 39, pl. 8), and are first seen
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| streaming from the glomerular capsule across the ventral
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| portion of the early primordium (figs. 14, 15, pl. 2). The
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| type C-II cells and their successors form most of the
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| adrenal capsule, and, during horizon xxiii, are diverted
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| inward by the arteriolar capillaries. Thereafter, they are
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| found in the cords of cells within the cortex. Their
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| nucleus is round, and stains a little more deeply than
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| does that of type C-I. The chromatin is distributed in
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| fine strands in a spider-like manner just beneath the
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| nuclear membranes, the nucleolus corresponding to the
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| body of the spider. The center of the nucleus is occupied
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| by one or more large globules or vacuoles.
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| Immigration and migration: The passage of cells from
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| one region to another; the addition of cells by movement
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| from an adjacent region. Ameboid movement is not
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| implied.
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| Invasion: The entrance of cells after destruction or disruption of pre-existing tissue, and the assumption of position without rcference to former relations.
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| Staging of embryos: The dating of the embryos follows the horizons described and graphed by Streeter
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| (1951). This method of seriation affords a much better
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| concept of the progress of development than does measurement of the length. Frequently, the organogenesis
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| of the adrenal gland was found to be in a much further | |
| advanced stage in an embryo with a shorter crown-rump
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| length than in a longer one, particularly in the earlier
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| 196 DEVELOPMENT OF THE ADRENAL GLAND IN MAN
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| stages when the dorsal curvature varies so remarkably
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| among embryos in the same horizon. This finding speaks
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| well for the method, since Streeter did not use the adrenal
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| among his criteria for staging. The age in weeks is the
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| estimated conception age. To obtain menstrual age, two
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| weeks must be added. There are enough stages to pro-
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| vicle a consecutive account of the organogenesis of the
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| adrenal.
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| ==Materials and Methods==
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| The following embryos of the Carnegie Collection were selected for the study of the embryonic and early fetal development of the adrenal gland.
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| (table to be formatted)
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| Crown-Rump Length (mm.)
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| saw
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| Horizon xnr
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| 836 . . . . . . 4.0 Trans. Exc. A1. coch.
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| 7433 . . . . . . 5.2 Frontal Exc. {{HE}}
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| 8066 . . . . . . 5.3 Trans. Exc. {{HE}}
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| 8119 . . . . . . 5.3 Trans. Exc. {{HE}}
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| Homzo.-' xtv
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| 3805 . . . . . . 5.9 Trans. Exc. {{HE}}
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| 6502 . . . . . . 6.7 Trans. Exc. {{HE}}
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| 7324 . . . . . . 6.6 Trans. Good {{HE}}
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| 7829 . . . . . . 7.0 Trans. Exc. {{HE}}
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| 7370 . . . . . . 7.2 Trans. Exc. {{HE}}
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| 8314 . . . . . . 8.2 Trans. Exc. Azan
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| 1-lomzow xv
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| 2. . . . . . 7.0 Trans. Good Al. carm.
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| 721 . . . . . . 9.0 Trans. lixc.
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| 3385 . . . . . . 8.3 Trans. Exc. {{HE}} or G.
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| 3441 . . . . .. 8.0 Sag. Exc. Al. coch.
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| 3512 . . . . . . S 5 Trans. Exc. Al. coch.
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| 6504 . . . . . . 7 5 Sag. Exc. A1. coch.
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| 6508 . . . . . . 7.3 Trans. Good Al. coch.
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| 6595. . . . , 3.0 Sag. Good I-1.-17..
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| 7199 . . . . . , 8.-'1 Frontal Good Al. coch. phloxine
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| 7364 . . . . . 9 5 Frontal Poor 1-1.-17..
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| 8929 . . . . . , 6 3 Frontal Exc. Azan
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| 8997 . . . . . . 9 0 Trans. Exc. Azan
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| 9140 . . . . . . 7 0 Trans. Exc. Azan
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| I-Ioruzox xvt
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| 792 . . . . . . 8 0 Trans. Good A1. coch.
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| 6511 . . . . . . 8 1 Sag. Good Al. coch. iron H.
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| 6512 . . . . .. 70 Trans. Exc. A1. coch.
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| 6517 . . . . . . 10 5 Trans. Exc. Al. coch.
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| 7115 . . . . .. 9 7 Frontal Exc. H. and pltloxinc
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| 7804 . . . . . . 9 5 Trans. Good {{HE}}
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| 7897 . . . . . . 12 2 Trans. Good {{HE}}
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| 8773 . . . . . . ll 0 Frontal Exc. Azan
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| l-lot-:1zo.\' xvu
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| 940 . . . . . . 14 0 Trans. Good {{HE}} or G.
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| 5893 . . . . . . 13 2 Trans. Good Al. coch.
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| 6253 . . . . . . 1-10 Trans. Good {{HE}}
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| 6519 , . . . . . 10 3 Sag. Exc. Al. coch.
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| 6520 . . . . . . 14 2 Trans. Exc. Al. coch.
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| 6742 . . . . . . 11 0 Trans. Good I-1.-pltloxine
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| 6758 . , . . . . 12 8 Trans. Good 1-1.-phloxine
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| 8101 . . . . .. 13.0 Trans. Exc. {{HE}}
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| 811$ . . . . .. 12.6 Frontal Exc. {{HE}}
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| 8789 . . . . . . ll 7 Sag. Exc. Azan
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| 3998 . . . . . . 11 0 Frontal Exc. Azan
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| Crown- _
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|
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| (mm.) | |
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| Horizon xvrn
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| -192 . . . , . . 16.3 Frontal Exc. Al. coch.
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| 1909 . . . . . . 14.6 Frontal Good .'-\l. coch. or G.
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| 4430 . . . . .. 14.0 Trans. Exc. A1. coch. or G.
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| 6524 . . . . . . 11.7 Trans. Exc. A1. coch.
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| 6525 . . . . .. 13.8 Sag. Exc. Al. coch.
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| 6527 . . . . .. 14.4 Trans. Exc. Al. coch.
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| 6528 . . . . .. 13.4 Frontal Exc. A1. coch.
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| 7707 . . . . .. 14 5 Trans. F.xc. 1-1.—F.. phloxine
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| 8172 . . . . . . 16 5 Trans. Exc. {{HE}}
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| S312 . . . . .. 12.2 Trans. F..\:c. I-1.717..
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| 9247 . . . . .. 15.0 Sag. Exc. Azan
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| Horizon xxx
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| 4-105 . . . . . . 15 5 Trans. Good Coch. l\1allor_\-'
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| 4501 . . . . .. 180 Trans. F..\'c. Coclt. or G.
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| 5609. . 18 0 Frontal F..\:c. Al. coch.
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| 6150 . . . . . . 17.0 Trans. Good
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| 632-l . . . . .. 18.5 Sag. Good {{HE}}
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| S092. . . . . . 16 3 Trans. lixc. I-1.-E. phlo.\;ine
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| 9113 . . . . . . 18 5 Trans. Exc. Azan
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| Horizon xx
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| 966.. . 23.0 Frontal F.xc. A1. coch.
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| 5537. . . . .. 22.0 Trans. Fxc. Al. coch.
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| 6202... . .. 21.0 Sag. F.xc. {{HE}}
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| 7274 . . . . .. 18 3 Trans. Exc. {{HE}} phloxinc
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| 7906 . , . . .. 19.5 Frontal F..\:c. {{HE}}
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| S226 . . . . .. 18.0 Sag. Exc. Azan
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| Horizon xxx
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| 5.596. . 21.5 Sag. Good I-1.-Ii.
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| 725-1.. . 22.5 Trans. Exc.
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| 7392.. . 22.7 Trans. Exc. {{HE}}
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| Z864. . 24 0 Frontal F.:~:c. {{HE}}
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| S553 . _ . _ . , 22.0 Trans. Exc. {{HE}}
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| Horizon xxn
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| 1458. . . . . , 27.5 Sag. Exc. or G
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| 430-1. . 25 0 Trans. Good {{HE}}
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| 4638. , . 23.4 Trans. Exc. Al. coch
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| 6701 . . . 24.0 Frontal Poor I-I.-E.
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| 6832. . 25.8 Frontal Exc. I-1..F_.
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| 339-1. . . 25 3 Trans. lixc. I1.-I’.
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| Horizon mm
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| 4205. . . . . . 29.5 Trans. Good Al coch
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| [[:Category:Carnegie Embryo 4289|'''No. 4289''']]. . . . . . 32.2 Trans. Good Al coch
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| 4570 . . . . .. 30.7 Trans. Exc. {{HE}}
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| 5-122. . . . . . 27.0 Sag. Good {{HE}}
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| 5621.-\ . . . . 27.5 Trans. Good
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| 6573. . . . . . 31.5 Trans. Good {{HE}}
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| 6719 . . . . .. 30.1 Trans. Good Chrom.alum hem.
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| 7425 . . . . .. 27.0 Frontal Exc. Cl1I‘0111.:1lUI11l1CI]'l.
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| 9226. . . . . . 30.1 Trans. Exc. Azan
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| {{Footer}}
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| [[Category:Adrenal]]
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| [[Category:Draft]]
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| [[Category:Carnegie Embryo 4289]]
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