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A Peculiar Structure in the Electroplax of the Stargazer, Astroscopus Guttatus James G

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A Peculiar Structure in the Electroplax of the Stargazer, Astroscopus Guttatus James G A PECULIAR STRUCTURE IN THE ELECTROPLAX OF THE STARGAZER, ASTROSCOPUS GUTTATUS JAMES G. HUGHES, JR. From the Histological Laboratory of Princeton University, U. S. A. THREE FIGURES The purpose of this paper is to determine the function and composition of the peculiar pointed fibers and long pointed rods lying in the electric layer of the electroplaxes of the stargazer, Astroscopus guttatus. Before proceeding with a discussion of these rods, a brief description of the electric organ of this fish (according to Dahl- gren)' will be given. The electric apparatus is composed of two organs, which form two vertical columns roughly oval in horizontal section, and placed behind and somewhat under each eye. Each organ extends from the peculiar bare spot on the top of the head down to the tissues which form the roof of the oval cavity; and is composed of about 200 thin layers of electric tissue, which extend horizontally all the way across the organ. These layers of tissue are flat, and always at the same distance from one another. Each layer contains about 20 electroplaxes, the outlines of which present a very irregular or scalloped appear- ance. The electric tissue in which the electroplaxes are im- bedded is in appearance a jelly-like or mucous-like tissue, usu- ally known as electric connective tissue, and which I have shown in the course of my work to be of the same composition as white fibrous connective tissue. The nerve and blood supply runs in the above tissue. The general form of a vertical section of an electroplax is shown by figure 1, whioh is a drawing of part of a section of a single electroplax. Each electroplax is composed of three principal layers, a nervous or electric layer which forms the upper surface, a 1 Anat. Am., Bd. 29, S. 387, 1906. 97 98 JAMES G. HUGHES, JR. middle layer, and a lower or nutritive layer which along with the middle layer is evaginated into a large number of long papillae. All three layers are deeply marked with a dense series of fine striations, which are peculiar to the electroplaxes of several other fishes. The upper or electric surface is flat and smooth and receives the nerve endings. The current of electricity runs downward through the organ which produces it, and thus the nerve endings in accordance with Pacini’s law are found on the negative pole of the electroplax. Proceeding directly to the subject of this paper, we may say that one of the most interesting of the points noted in the elec- troplaxes, when properly fixed and stained with iron hematoxy- lin, is a series of rod-like or thread-like objects running hori- zontally in the electric layer, among, above and below the nuclei and without any apparent connection with them (figs. 1 and 2).2 These rods are of various sizes and shapes, and in form are said to resemble the classic thunderbolts seen in the hand of rep- resentations of Jove. They usually taper slowly end branch extensively at one or both ends. Some of these brznches some- times seem to be mere lines, while others are wide and heavily pointed; at their other ends the rods are usually rounded; this latter appearance may be due, however, to the cut ends of the rods, for as noted above they sometimes branch at both ends. Some are short and heavy in appearance while others me long and thread-like. Peculiar looping, twisting, or knot-like bends are sometimes found at points on the longer rods. The out- lines and contour of these rods are always smooth. Their size may vary from thick or thin rods of over 300 1.1 in length down to small ones that do not exceed 1 p. In those electroplaxes where the rods are few they sometimes lie parallel and point in a definite parallel direction, while in others where the rods are very numerous they do not seem to have any definite arrangement. In this latter condition the rods present a very wavy appearance. Their form may be seen in figures 1 and 2, which are drawings of the electric layer of an electroplax when All the figures are drawings of sections of electroplaxes of Astroscopus guttatus. 99 Fig. 1 Vertical section of part of an electroplax. The whole electroplax as well as the electric connective tissue is shown; rods arc sccn lying in the electric layer. X 1200. 100 JAMES G. HUGHES, JR. Fig. 2 Horizontal section through an electroplax. Only the electric layer is seen; a, an extensively branching rod; b, fine branches of the above rod; c, x large characteristic loop io a rod;d, a cut end of a rod; e, nuclei of the electric layer. X 1200. stained with iron hematoxylin. These drawings show the electric layer in which the rods are found in horizontal section (fig. 2) and in vertical section (fig. 1). The purpose, function, and chemical composition of these rods have been previously unknown to histologists. In order to determine anything in respect to their function or purpose, a knowledge as to the class of organic substance to which they belong, whether muscle connective tissue nervous or chitinous ELECTROPLAX OF ASTROSCOPUS 101 and also a rough knowledge of their chemical composition is imperative. The contour and form of these rods as they appear under the microscope resemble both smooth muscle fibers and fibers of elastic connective tissue. The belief that the function of these rods was somewhat of the nature of support for the delicate substance of the electroplax, and the fact that their form resembled connective tissue fibers led the writer to take for one of his first hypotheses, that they were of some form of connective tissue, and to perform accordingly the following series of experiments. As the most logical and best way for determining the kind of connective tissue, if any, of which the rods might be composed, a number of stains used by other investigators to identify similar substances were applied and the results noted. Controls were used on known tissues. Before taking up the connective tissue stains, however, a description of the results from the iron hematoxylin staining is now noteworthy; the material used being fixed in pure corrosive sublimate. The jelly connective tissue stained a very light gray. The nutritive, striated, and electric layers stained a much darker gray. The nuclei in all the layers stained somewhat black. The pointed fibers, and rods found in the electric layer stained a deep black, thus being clearly differentiated from the surround- ing cytoplasm. In some electroplaxes they were very numer- ous, while in others the number was rather small. The connective tissue stains applied as follows: (1) Mallory’s connective tissue stain, using the modifica- tion given in Lewis’ “Text-book of histology.” White fibrous connective tissue should stain blue in this medium. Paraffin sections of the electroplaxes fixed in corrosive subli- mate were stained for 10 to 12 minutes in a 1 per cent aqueous solution of acid fuchsin. They were then transferred directly to a stain consisting of 0.05 grain of aniline blue (soluble in water) and 0.2 grain of orange G dissolved in 100 cc. of a 110 per cent aqueous solution of phospho-molybdic acid. In this they remained from 2 to 3 minutes. They were then rinsed in distilled water, dehydrated rapidly, cleared and mounted. 102 JAMES G. HUGHES, JR. A description of an electroplax as seen under the 2 mm. oil immersion lens is as follows: The white fibrous or electric con- nective tissue stained a light blue or purple. The electric nutri- tive and middle layers or the electroplax proper stained a red- dish purple, and the nuclei as a whole in all of the layers, stained somewhat lighter than their surrounding cytoplasm. The peculiar rods and fibers stained a deep red, and were thus clearly differentiated from the other elements of the elec- troplax. They were very numerous and as noted above their outlines were always smooth. Blood corpuscles lying in the jelly connective tissue, of which there were only a few, stained a brilliant red, much the same as the rods. The white fibrous or jelly electric connective tissue (be- tween the electroplaxes), as noted above, stained a light blue. The pointed fibers and rods stained a deep red. This would indicate therefore, that these rods are not composed of white fibrous connective tissue. It would seem also that they are not muscle for they stained a different shade of color from the rest of the electroplaxes, which I have found in the course of my work to stain much the same as muscle. (2) Van Gieson’s connective tissue stain, in which white fibrous connective tissue should stain red: Pzraffin sections of the electroplaxes fixed in corrosive sub- limate were stained for 4 to 5 minutes in a 1 per cent aqueous solution of hematoxylin. They were rinsed in distilled water and transferred to a stain consisting of a saturated aqueous solution of picric acid containing 20 per cent acid-fuchsin. They remained in here 15 to 20 minutes and were then rinsed, cleared and mounted. The white fibrous connective tissue layer stained pink. The three layers of the electroplax stained brown. The nuclei and rods stained the same color, that is, brown; and, while the nuclei could be seen with difficulty, the rods were scarcely visible owing probably to their similar refractive in- dex. The fact therefore that the rods did not stain the same color as the white fibrous connective tissue, which stained pink, indicates again that they are not composed of white fibrous connective tissue.
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