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Studies on the Origin of Yolk. II. Oogenesis of the Scolopendra, Otostigmus Feae (Pocock)

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Studies on the Origin of Yolk. II. Oogenesis of the Scolopendra, Otostigmus Feae (Pocock) Studies on the Origin of Yolk. II. Oogenesis of the Scolopendra, Otostigmus Feae (Pocock). By Yishwa Nath, M.Sc, Ph.D. (Cantab.) and Mian Tasdique Husain, M.Sc. Department of Zoology, Government College, University of the Punjab, Lahore. With 11 Text-figures. IN a previous paper (1924) on the egg of the centipede, Lithobius f orficatus, one of us (V. N.) described two kinds of yolk, albuminous and fatty. The albuminous yolk is pre- ceded by nucleolar extrusions of a remarkable type, and its origin seems to be associated with them, although no evidence could be adduced that they are directly transformed into the yolk. It was further shown that the juxta-nuclear Golgi apparatus fragments into small granules and small crescent- shaped Golgi elements. The former grow in size and give rise to the fatty yolk. Miss S. D. King (1924) confirms the above account of the association of the albuminous yolk with the nucleolar extrusions, but in her opinion this type of yolk arises directly from these extrusions. With regard to the Golgi apparatus she admits that it fragments into small granules. Furthermore she describes fatty yolk :—' the origin of this fatty yolk is doubtful, but it may possibly be connected with the Golgi apparatus, although no evidence in support of this theory has been discovered '. In our opinion Miss King is doubtful of the origin of fatty yolk from the Golgi apparatus, because ' both Mann-Kopsch and Da Fano material was studied, but the latter gave such favourable results that it was used almost exclusively ' (spaced words ours). Now it is well NO. 287 D d 404 VISHWA NATH AND MIAN TASDIQUE HUSAIN known that the Da Fano method fails to show fat in the finished slides, while the Golgi apparatus is preserved. On the other hand the Mann-Kopsch method preserves both Golgi apparatus and fat, and thus enables one to study the trans- formation of the former into the latter. Furthermore, Flem- ming-without-acetic, which Miss King has used, shows fat, but not the Golgi apparatus, at least in the egg ofLithobius. In a later paper (1926), however, Miss King has shown that ' the formation of fatty yolk from the Golgi elements in Oniscus is strictly comparable to the process described by Nath in Lithobius. .' In view of the above conflicting evidence it was thought desir- able to work out the origin of yolk in the eggs of the centipede Scolopendra. Qur observations, both on fixed and fresh material, unmistakably show the origin of the fatty yolk from the Golgi elements and furnish a strong confirmation of the same process in Lithobius. The nucleolar extrusions of Scolopendra are few and they disappear at a particular stage ; and between their disappearance and the origin of albuminous yolk there is a long pause which eliminates all possibility of the direct origin of this type of yolk from these extrusions. The routine laboratory technique employed is mentioned in the text and in the explanatory notes on the figures. One of us (V. N.) has also studied fresh material with the help of neutral red and janus B. All diagrams have been drawn by Vishwa Nath. It is a very great pleasure to us to thank Colonel S. E. Christophers, F.E.S., Director of the Central Eesearch Institute, Kasauli, who gave us all facilities for this work in his institute in the summer of 1926. Our thanks are also due to the authorities of the Indian Museum for identifying the Scolopendra and for supplying us with the necessary literature. OOGENESIS OF SCOLOPENDKA 405 OBSERVATIONS. Staining with vital dyes gave good results. The ovary is kept in neutral red or janus green B for about fifteen minutes. It is then mounted on a slide in a drop of the stain, covered with a cover-slip, and studied under the high power of the micro- scope. Text-fig. 9 represents a fairly young oocyte. The whole oocyte is full of very clear colourless vacuoles of similar size distributed in a uniform manner throughout the cytoplasm ; these are the Golgi vacuoles. In the spaces between the vacuoles lie the granular mitochondria also distributed uniformly. Both the cytoplasm and the mitochondria are stained red or green according to the dye used. The mitochondria, however, stain deeper than the cytoplasm and appear more distinctly with janus green B than with neutral red. With the growth of the oocyte both the Golgi vacuoles and the mitochondria increase in number (Text-fig. 10). The mitochondria remain granular, but some of the Golgi vacuoles increase in size and give rise to the fatty yolk. Both the Golgi vacuoles and the fatty yolk- vacuoles appear as colourless vacuoles, but their contents are chemically different, as will soon appear. Excellent results are obtained when a most highly developed egg is stained with neutral red. Such an egg can be studied only under the lower power as the big size of the egg does not allow the use of the cover-slip. In Text-fig, lla portion of such an egg stained with neutral red is represented. It is studied under the low power and is magnified only forty times. Nevertheless appearances are brilliant. At A.Y. is the albuminous yolk which appears as colourless, solid, and homogeneous spheres. At F.Y. are the fatty yolk-vacuoles which appear brilliantly red as the stain has penetrated inside them. The smaller Golgi vacuoles, how- ever, remain colourless. The granular mitochondria cannot be seen with this magnification. If now the egg is broken with a needle the solid albuminous yolk-spheres are seen scattered on the slide, but the vacuoles, both Golgi and fatty yolk, being delicate are ruptured and do not appear. In some cases, how- ever, a small portion of the cytoplasm of the ruptured egg D d 2 406 VISHWA NATH AND MIAN TASDIQUB HUSAIN remains intact and the vacuoles can be seen, as if held in position by the solid albuminous yolk-spheres. On account of the paucity of material oocytes younger than that represented in Text-fig. 9 could not be studied with the vital dyes. Text-fig. 1 represents an unstained Mann-Kopsch preparation G V Youngest oocyte showing Golgi vacuoles and mitochondrial ring. Mann-Kopsch unstained, x 2,100. EXPLANATION or LETTERING OF TEXT-FIGURES. AY, albuminous yolk; 0, cytoplasm; FY, fatty yolk; OV, Golgi vacuole; OV, Golgi vacuole looking solid on account of excessive impregnation; G V", Golgi' crescent' with ' idiosome'; 0 V", Golgi ' crescent' without ' idiosome'; M, mitochondria; N, nucleus; N', nucleolus ; N", secondary nucleoli; NE, nucleolar extrusions. of the youngest ooeyte that we have been able to obtain. The mitochondria appear granular and brownish and form a com- plete ring round the nucleus. At G.V. are the Golgi vacuoles, each with a sharp black chromophilic rim and a central chromo- phobic substance. The contents of each vacuole are watery. At G.V.' the Golgi vacuoles appear solid on account of the pre- cipitation of the metallic osmium inside their interior. With the growth of the oocyte the mitochondrial ring spreads out at its OOGENBSIS OF SCOLOPENDRA 407 periphery and the Golgi vacuoles increase in number, as shown in Text-fig. 2 which represents a Mann-Kopsch preparation stained with acid fuchsin. In Text-fig. 3, which is a Mann- Kopsch unstained preparation, the rnitochondrial ring has GV Slightly older oocyte showing proliferation of the Golgi vacuoles and the spreading out of the initochondrial ring. Mann-Kopsch stained. X 1,800. spread out and is touching the egg-membrane at the two poles. The Golgi vacuoles have further increased in number and most of them look solid for the reason given above. With the further growth of the oocyte a change takes place in the contents of the Golgi vacuoles. Text-fig. 4 represents a Champy-Kull prepara- tion stained with acid fuchsin. At G.V. are the Golgi vacuoles which are stained with acid fuchsin. Each vacuole shows a red 408 VISHWA NATH AND MIAN TASDIQUB HUSAIN rim and a clear central substance. The contents of these vacuoles are undoubtedly watery and non-fatty. At F.Y. are shown vacuoles whose contents have become fatty. In unstained Champy-Kull preparations these vacuoles appear solid and dull black on account of the coagulation and osmication of the fatty contents by osmic acid. But when the slide is heated with the TEXT-FIG. 3. GV Older oocyte showing the same phenomenon. Mann-Kopsch un- stained. X 1,680. acid fuchsin stain floating on it, these solid and dull black bodies appear as clear vacuoles on account of their decolorization by acid fuchsin aided probably by heat. How acid fuchsin which is dissolved in anilin-oil water decolorizes osmicated bodies we cannot explain. It is, however, certain that acid fuchsin tends to decolorize osmicated bodies in much the same way as turpen- tine and xylol. It will be noticed that in Text-fig. 4 the number OOGENESIS OF SCOLOPENDRA 409 of Golgi vacuoles is much smaller than in Text-fig. 3, although the latter represents a younger oocyte. This, however, is due to the Champy-Kull method which is certainly inferior to the Mann-Kopsch method so far as the fixation of the Golgi vacuoles with Avatery contents is concerned. Text-fig. 5 represents an oocyte of a later stage fixed with Mann-Kopsch and stained with acid fuchsin. This is one of the best preparations in our posses- sion as it comes very near to fresh eggs studied with vital dyes. The whole oocyte is full of Golgi vacuoles, each having a sharp V A young oooyte showing Golgi vacuoles, fatty yolk-vacuoles, mito- chondria, and nucleolar budding.
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