The different suprarenal bodies exhibit variations in structure mainly dependent on the ganglion cells and nerves in them, and their typical structure is best exhibited in a posterior one, in which there is a comparatively small development of nervous elements.
A portion of a section through one of these is represented on Pl. 19, fig. 6, and presents the following features. Externally there is present a fibrous capsule, which sends in the septa, imperfectly dividing up the body into a series of alveoli or lobes. Penetrating and following the septa there is a rich capillary network. The parenchyma of the body itself exhibits a well-marked distinction in the majority of instances into a cortical and medullary substance. The cortical substance is formed of rather irregular columnar cells, for the most part one row deep, arranged round the periphery of the body. Its cells measure on about an average .03 Mm. in their longest diameter. The medullary substance is more or less distinctly divided into alveoli, and is formed of irregularly polygonal cells; and though it is difficult to give an estimate of their size on account of their irregularity, .021 Mm. may be taken as probably about the diameter of an average cell. The character of the cortical and medullary cells is nearly the same, and the cells of the two strata appear rather to differ in shape than in any other essential point. The protoplasm of both has a markedly yellow tinge, giving to the suprarenal bodies a yellowish brown colour. The nuclei are small compared to the size of the cells, being about .009 Mm. in both cortical and medullary cells. In the anterior suprarenal body there is a less marked distinction between the cortical and the medullary layers, and a less pronounced yellow coloration of the whole, than in the posterior bodies. The suprarenal bodies are often partially or completely surrounded by a lymphoid tissue, which is alluded to in the account of their development.
The most interesting features of my sections of the anterior bodies are the relations they bring to light between the sympathetic ganglia and the suprarenal bodies. In the case of one of the posterior suprarenal bodies, a small ganglion is generally found attached to both ends of the body, and invested in the same sheath; in addition to this a certain number of ganglion cells (very conspicuous by their size and other characters) are to be found scattered through the body. In the anterior suprarenal bodies the development of ganglion cells is very much greater. If a section is taken through the region where the large sympathetic ganglion (already mentioned) is attached to the body, one half of the section is composed mainly of sympathetic ganglion cells and nerve fibres, and the other of suprarenal tissue, but the former spread in considerable numbers into the latter. A transverse section through the suprarenal body in front of, or behind this point, is still more instructive. One of these is represented in Pl. 19, fig. 7. The suprarenal tissue is not inserted, but fills up the whole space within the outline of the body. At one point a nerve (n) is seen to enter. In connection with this are a number of ganglion cells, the exact distribution of which has been reproduced. They are scattered irregularly throughout the suprarenal body, but are more concentrated at the smaller than at the large end. It is this small end which, in succeeding sections, is entirely replaced by a sympathetic ganglion. Wavy fibres (which I take to be nervous) are distributed through the suprarenal body in a manner which, roughly speaking, is proportional to the number of ganglion cells. At the large end of the body, where there are few nerve cells, the typical suprarenal structure is more or less retained. Where the nerve fibres are more numerous at the small end of the section, they give to the tissue a somewhat peculiar appearance, though the individual suprarenal cells retain their normal structure. In a section of this kind the ganglion and nerves are clearly so intimately united with the suprarenal body as not to be separable from it.
The question naturally arises as to whether there are cells of an intermediate character between the ganglion cells and the cells of the suprarenal body. I have not clearly detected any such, but my observations are of too limited a character to settle the point in an adverse sense.
The embryological part of my researches on these bodies is in reality an investigation of later development of the sympathetic ganglia. The earliest stages in the development of these have already been given[332], and I take them up here as they appear during stage L, and shall confine my description to the changes they undergo in the anterior part of the trunk. They form during stage L irregular masses of cells with very conspicuous branches connecting them with the spinal nerves (Pl. 18, fig. 3). There may be noticed at intervals solid rods of cells passing from the bodies to the aorta, Pl. 18, fig. 2. These rods are the rudiments of the aortic branches to which the suprarenal bodies are eventually attached.
In a stage between M and N the trunks connecting these bodies with the spinal nerves are much smaller and less easy to see than during stage L. In some cases moreover the nerves appear to attach themselves more definitely to a central and inner part of the ganglia than to the whole of them. This is shewn in Pl. 19, fig. 8, and I regard it as the first trace of a division of the primitive ganglia into a suprarenal part and a ganglionic part. The branches from the aorta have now a definite lumen, and take a course through the centre of these bodies, as do the aortic branches in the adult.
By stage O these bodies have acquired a distinct mesoblastic investment, which penetrates into their interior, and divides it, especially in the case of the anterior bodies, into a number of distinct alveoli. These alveoli are far more distinct in some parts of the bodies than in others. The nerve-trunks uniting the bodies with the spinal nerves are (at least in specimens hardened in picric and chromic acids) very difficult to see, and I have failed to detect that they are connected with special parts of the bodies, or that the separate alveoli differ much as to the nature of their constituent cells. The aortic branches to the bodies are larger than in the previous stage, and the bodies themselves fairly vascular.
By stage Q (Pl. 19, fig. 9) two distinct varieties of cells are present in these bodies. One of these is large, angular, and strikingly resembles the ganglion cells of the spinal nerves at the same period. This variety is found in separate lobules or alveoli on the inner border of the bodies. I take them to be true ganglion cells, though I have not seen them in my sections especially connected with the nerves. The cells of the second variety are also aggregated in special lobules, and are very markedly smaller than the ganglionic cells. They form, I imagine, the cells of the true suprarenal tissue. At this and the earlier stage lymphoid tissue, like that surrounding the suprarenal bodies in the adult, is found adjacent to these bodies.
Stage Q forms my last embryonic stage, and it may perhaps be asked on what grounds I regard these bodies as suprarenal bodies at all and not as simple sympathetic ganglia.
My determination mainly rests on three grounds: (1) That a branch from the aorta penetrates these bodies and maintains exactly the same relations to them that the same branches of the aorta do in the adult to the true suprarenal bodies. (2) That the bodies are highly vascular. (3) That in my last stage they become divided into a ganglionic and a non-ganglionic part, with the same relations as the ganglia and suprarenal tissue in the adult. These grounds appear to me to afford ample justification for my determinations, and the evidence adduced above appears to me to render it almost certain that the suprarenal tissue is a product of the primitive ganglion and not introduced from the mesoblast without, though it is not to be denied that a more complete investigation of this point than it has been possible for me to make would be very desirable.