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The Cytology of Amoeba Proteus 'Y' and the Effects of Large and Small Centrifugal Force S

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The Cytology of Amoeba Proteus 'Y' and the Effects of Large and Small Centrifugal Force S The Cytology of Amoeba proteus 'Y' and the effects of large and small Centrifugal Force s. By B. N. Singh, M.Sc. (Allahabad) (Department of Zoology, Trinity College, Dublin.) With Plates 50-51 and 2 Text-figures. CONTENTS. PAGE 1. INTRODUCTION AND PREVIOUS WORK ..... 601 2. MATERIAL AND METHODS 612 3. OBSERVATIONS ON AMOEBA PROTEUS'Y' 615 (i) General Morphology of Normal and Centrifuged Organisms, p. 615. (ii) Nutritive Spheres, p. 616. (iii) Crystals, p. 619. (iv) Neutral-red Staining Bodies, p. 619. (v) Fat, p. 620. (vi) Nucleus, p. 622. (vii) Contractile Vacuole, p. 622. (viii) Mitochondria, p. 622. (ix) Ectoplasmic Folds, p. 624. (x) Vitamin C, p. 624. 4. DISCUSSION .......... 625 5. SUMMARY 632 1. INTRODUCTION AND PEEVIOUS WORK. THE development of the air-driven ultra-centrifuge by J. W. Beams and his associates (1930-3) has proved a valuable means of discriminating between the pre-existing cytoplasmic bodies and those that are fixation artefacts. Ultra-centrifuging has very clearly shown that the Golgi apparatus is not an artefact produced by the action of fixatives on cytoplasmic colloids. It is also a very satisfactory means by which to distinguish between the pre-existing neutral-red bodies and those produced by the segregation of the dye. One can very easily distinguish, by the above method, the separate identity of the Golgi bodies from neutral-red staining bodies where both types of inclusions occur side by side. In the Sporozoa, where the true homologue of the Golgi apparatus of the Metazoa has been clearly shown to exist by different workers, one can very easily identify 602 B. N. SINGH the separate Golgi bodies and neutral-red staining bodies by centrifuging, and we may also ascertain whether the latter bodies are pre-existing, or artefacts produced by the neutral- red staining. As Amoeba proteus is very often used as a type of simple cell, it was thought that the detailed study of its cytology and the effect of centrifuging would be of interest, and might shed some further light on the question of the homology of the protozoan and the metazoan Golgi apparatus. It was also thought possible to ascertain whether any homologue of the Golgi apparatus exists in Amoeba proteus as claimed by Causey (1925), Hirschler (1927 a), Hall (19806), Hall and Loefer (1980), Nigrelli and Hall (1980), and V. E. Brown (1930) in the Protozoa Sarcodina. A little work has been done on this subject by V. E. Brown (1980), who claims to have demon- strated two types of Golgi bodies in Amoeba proteus. One of these is composed of granules and the other of spherules with black rims (after the Mann-Kopsch fixative). After Champy fixation and staining with acid fuchsin, thionin, and aurantia, he found that the spherules are light blue with dark blue rims. According to him, the spherules when viewed at a central focus appear as rings and crescents having clear centres. The contractile vacuole does not blacken with osmic acid, and in no case are the Golgi bodies associated with it. He found small granules grouped around the contractile vacuole which do not take osmic stain, but are revealed by acid fuchsin. In addition to these he finds small vacuoles of variable size which are attached to the supposed Golgi bodies, and are also found in the endoplasm, and they are numerous around the contractile vacuole. He believes that the small vacuoles originate from small granules which move about in the cyto- plasm and are grouped around the contractile vacuole. They flow together to form the new vacuole after systole. Brown thinks that it may be possible that the black granules, which are of variable size, and which he identifies as Golgi bodies, grow in size to form black spheres with black rims, and compares these two types of Golgi bodies with those observed by Hirschler (1914), King and Gatenby (1928), Joyet Lavergne AMOEBA PROTEUS 603 (1926), and Hall (19306). He did not use silver methods to confirm his results, and has not drawn mitochondria in his diagrams, to avoid confusion with the various types of bodies that he has described. According to Brown no network exists in Amoeba proteus, so he is led to believe that in Pro- tozoa secretion is a constant process and, therefore, it is natural to expect globules, which are hypertrophied Golgi network. He believes that the relationship existing between crescent-shaped Golgi bodies and the small vacuoles which are present throughout the endoplasm, and are also attached to the supposed Golgi crescents, is the same as pointed out by Bowen (1926) in the case of gland-cells. Sister Carmela Hayes (1924) pointed out that the nutritive spheres in Amoeba proteus, as so termed by Sister Monica Taylor (1924), are of the nature of glycogen and not of fat, for which she has given various reasons. She found abundance of minute starch grains by staining amoebae with a solution of iodine in potassium iodide. When amoebae containing nutritive spheres were crushed under a coverslip and treated with iodine solution, the small nutritive spheres stained dark brown and the large ones stained less deeply. The big spheres seem to form centres around which starch granules collect. The nutri- tive spheres become yellow to reddish brown after treatment with iodine, and red after Best's carmine test for glycogen. The Delafield's haematoxylin and light green method stains nutritive spheres purple. Sister Carmela Hayes noted that in smear preparations the spheres lost their spherical form and became irregular patches which stained deeply in thionin and Delafield's haematoxylin. Eecently a detailed account of the cytology and the effects of centrifuging has been given by Mast and Doyle (1935 a and 19356). They used Amoeba proteus 'Y' and Amoeba proteus 'X' (Amoeba dubia) in their work. According to them the cytoplasmic bodies are: alpha granules, beta granules (mitochondria), crystals (lying inside crystal vacuoles), fat, and a single contractile vacuole. By centrifuging, the cytoplasmic bodies are stratified in different layers. Beginning with the aggregation of fat-globules 604 B. N. SINGH at the extreme centripetal end in the cell, the other cytoplasmic bodies, except the alpha granules, are stratified as follows: (1) crystal vacuoles without crystals; (2) crystal vacuoles with very small crystals and the contractile vacuole; (3) hyaline and very finely granular substance; (4) the nucleus and the food vacuoles containing much fat and little starch; (5) crystal vacuoles with crystals of moderate size; (6) crystal vacuoles with large crystals, mitochondria, and food vacuoles containing little if any fat; (7) refractive bodies. The alpha granules were more abundant in the centrifugal half than in the centripetal half of the organism. They have shown that the crystals lie in the centrifugal side of the crystal vacuoles, and the chromatin, being heavier than the karyo- lymph, collects at the centrifugal side of the nucleus. There is a thick layer of substance, lying at the centrifugal side of the contractile vacuole, with beta granules adhering to it, and only a thin layer at the opposite side, with no beta granules. This shows that both beta granules and the substance surrounding the contractile vacuole collect at the centrifugal side of the vacuole. By the use of the microscope centrifuge they observed that some of the amoebae had no pseudopodia and were elongated, while others had various numbers or sizes of pseudopodia. The fat-globules formed permanent aggregations in the pseudo- podia and sometimes the tips of these pseudopodia were separated from the rest of the body. In support of this, they point out that many amoebae, which were fixed after centri- fuging, had no fat. After the stratification of the cytoplasmic components, the amoebae continued to elongate, and in doing so, the central portion became smaller and smaller till the two ends rapidly separated. The alpha granules are approximately 0-25 of a micron in diameter, and are so near the limit of microscopic vision that they were neither able to make out the structure of these granules in the living condition, nor were they able to stain or identify them in fixed preparations. The beta granules (mitochondria) are about 1 micron in diameter, distributed uniformly throughout the cytoplasm, and AMOEBA PROTEUS 605 form a layer on the surface of the contractile vacuole. These granules are nearly spherical or ellipsoidal in the living condi- tion. They change in form, and protuberances like pseudopodia appear at intervals. Sometimes the spherical ones become ellipsoidal or rod-like in form and vice versa. With Janus green solution, both rod-shaped bacteria and mitochondria are stained a green colour. Due to the action of the fixatives, mitochondria become more elongated. Mast and Doyle found that the organisms, from which all the mitochondrial granules are removed, die after 12 hours, showing that these bodies are essential for life. The increase in number of the mitochondrial granules was seen in those organisms from which only 15 per cent, of them were removed. They were not able to make out whether the beta granules arise d e n o v o or by the division of the pre-existing ones. By changing the number of beta granules by cutting off different portions of the amoebae, they noted the rate of elimination of the fluid by the contractile vacuole, and thus came to the conclusion that the frequency of the contraction of the contrac- tile vacuole varied directly with the number of beta granules per unit volume of cytoplasm. Thus, according to the above conclusion, they point out that the beta granules function in the accumulation of the fluid eliminated by the contractile vacuole, i.e.
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