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Morphological and Histochemical Studies on the Uropygial Glands of Pigeon and Domestic Fowl

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Morphological and Histochemical Studies on the Uropygial Glands of Pigeon and Domestic Fowl 124 Cytologia 26 Morphological and Histochemical Studies on the Uropygial Glands of Pigeon and Domestic Fowl Kuldip Chand Kanwar1 Department of Zoology, Panjab University, Hoshiarpur, India Received June 28, 1960 Introduction Although quite a good deal of cytological work has been done in recent years on glands secreting triglycerides (e. g., sebaceous glands, mammary glands and Harderian glands), uropygial glands or more popularly known as oil glands of birds have not received closer attention of the workers. A few papers available (Altmann 1894, Bowen 1926, Bhatia 1943, Das and Ghosh 1959) dealing with the morphological and histochemical studies of these glands supply insufficient data as to how exactly the secretion is manufactured inside these holocrine glands and more particularly, as to what is the role of the mitochondria during the secretory genesis. Altmann (1894) and Bowen (1926) employed, almost exclusively, techniques involving prolonged osmication which, as Nath (1944) and Baker (1953) have also observed, have little morphological and absolutely no histochemical validity. Similarly Bhatia (1943) also depended mainly on Kolatchev and F. W. A./iron haematoxylin preparations. Recently Das and Ghosh (1959), who worked out the histochemistry of the oil gland cells of the pigeon, have published only a brief note. As early as 1894, Altmann observed that the peripheral envelope of each secretion droplet differs chemically from the central part. He based his observations on the fact that in various osmium preparations it is only the periphery of the secretion droplet which gets impregnated whereas the se cretion droplet proper remains untinged. Bowen (1926), who worked on a variety of holocrine cells, contrary to the findings of Altmann (1894), established that there is no peripheral im pregnation of the secretion droplets and whatever is impregnated in various osmium preparations is not a part of the secretion droplets but the remnant of the 'Golgi apparatus'. Bhatia (1943) recorded that the secretion is always formed within the 'Golgi interna' surrounded with highly o smiophil cortices ('Golgi externa') . Quite recently, Baker (1954, 1957) and Nath (1957, 1959) have argued that the use of the terms like 'Golgi material', 'Golgi apparatus', 'Golgi 1 Present address: Department of Zoology, Panjab University, Chandigarh, India. 1961 Morphological and Histochemical Studies on the Uropygial Glands 125 bodies' etc. should be dropped , as these terms do not refer to structures which can be held homologous in different types of cells . These renowned cytologists further deny the existence of the classical 'Golgi apparatus' , from all types of cells, to which Bowen (1926) attributes a crucial role in the synthesis of the secretion. These observations of Baker and Nath call for fresh investigations by employing the rational morphological and various other cytochemical techniques. The purpose of this communication is to correlate the present findings with those of other authors and, if possible, to bring about unanimity in the varied descriptions available upto this date. Material and techniques Only the male pigeons and fowls were used for the present investiga tions. The animals were dissected alive and the uropygial glands were taken out. These were further cut into small pieces which in turn were transferred to the physiological solution. The material was then dropped into various fixatives and processed accordingly. The various techniques which have been employed include formaldehyde osmium (Baker 1944) followed by gelatin as well as paraffin embedding, Kolatchev, formaldehyde calcium (Baker 1946) -both with and without postchroming , Regaud, Lewitsky saline (Baker 1956a), Zenker and Carnoy. The paraffin sections of the last four fixatives were usually stained with 0.5% iron haematoxylin. Regaud fixed material was also coloured with Sudan black B (SBB) and also treated with acid fuchsine for the demonstration of the mitochondria. Carnoy fixed material was mainly used for staining with mercuric bromophenol blue (Hg-BPB) for the detection of proteins (Mazia et al. 1953). The various dyes employed for colouring the lipids include ethanolic Sudan black B (Baker 1949), nile blue sulphate (Cain 1947), Sudan III and IV (Kay and Whitehead 1941) and Fettrot (Pearse 1954). These have been mainly used on gelatin sections. Extractions with lipid solvents like acetone and ethanol were tried on formaldehyde calcium (FCa) unpostchromed gelatin sections. In addition to all this, acid haematein (AH) technique for the identification of phospholipids (Baker 1946) along with its pyridine extraction control, Performic acid-Schiff (Pearse 1951) for unsaturated bonds and Fischler's test for fatty acids (Pearse 1954) have also been employed. Schultz's test for cholesterols (Gomori 1952) was also performed. Periodic acid-Schiff (PAS) technique (Hotchkiss 1948) for the detection of general carbohydrates (1•F2 glycol group) coupled with its acetylation control and KOH reversal treatment (Gersh 1949) was used both on paraffin and gelatin sections. For the cytochemical detection of RNA (cytoplasmic basiphilia), paraffin sections of Zenker (chilled) fixed material were stained with pyronin G/methyl 126 K. C. Kanwar Cytologia 26 green (Jordan and Baker 1955). Control sections, prior to their staining with pyronin G/methyl green, were treated with trichloroacetic acid (Schneider 1945) and perchloric acid (Erickson et al. 1949), both of which dissolve out the RNA completely. Observations The uropygial glands of the pigeon and the fowl are multiple acinar holocrine glands. The whole gland, in both the animals, consists essentially of a sac with a central lumen. Since the new cells are regularly being added to replace those which are constantly being destroyed along with the dis charge of the secretion products (holocrine), all stages of secretory cycle can be observed even in a small portion of the gland. The glandular acini consist of small undifferentiated peripheral cells and large 'vacuolated' cells. Progressive maturity of the cells from the periphery towards the centre is indicated by the increasing quantity of lipid (sudanophil) material in the cytoplasm-the cells just near the lumen being fully packed with lipid droplets (Fig. 22). The gland cells form a stratified epithelium arranged in the form of long tubes. The centre of each tube is pierced by a narrow lumen through which the secretion products are discharged into the common duct. The earliest undifferentiated cells or 'indifferent cells' of Das and Ghosh (1959) are small, peripheral in position and these rest on an acinar basement membrane. There is a gradual increase in size of these cells from the peri phery towards the lumen. These earliest cells-farthest from the lumen, contain comparatively large, more or less centrally placed nuclei (with one or sometimes more nucleoli each). The cytoplasm of these cells is more or less homogeneously basiphil. Since the basiphilia is labile in salivary RNase, perchloric acid and trich loroacetic acid, it is evidently due to the presence of RNA. The ground cytoplasm of these cells is stained appreciably with mercuric bromophenol blue and also gives homogeneous diffuse pink coloration after periodic acid -Schiff . There is observed a gradual decrease in the cytoplasmic basiphilia and cytoplasmic affinity for mercuric bromophenol blue and periodic acid -Schiff from periphery towards lumen, so much so that the cells adjoining the lumen are almost completely devoid of these various colorations. Mitochondria. The cytoplasm of the early undifferentiated cells reveals uniformly distributed mitochondria which are in the form of rods or small stout filaments, each usually with one granule (Figs. 1-10). These mitochondria appear sufficiently impregnated in Kolatchev and formaldehyde osmium preparations (Fig. 1). The surface or end granules in such preparations appear jet black. The mitochondria are very well preserved in all the fixatives employed. In FCa/SBB (Fig. 4) and Lewitsky saline/SBB (Fig. 6) preparations, the mitochondria are stained distinctly with intense 1961 Morphological and Histochemical St udies on the Uropygial Glands 127 Figs. 1-22. Abbreviations. LG, lipid granule. M, mitochondria. N, nucleus. N, nucleolus . P, peripheral investment. PR, peripheral remnants. S, secretion granule or droplet . SG, surface granule. SV, secretion vacuoles. All the figures have been drawn with the help of camera lucida. 1, fowl. Kolatchev. 2, 3, fowl. Formaldehyde calcium/acid haematein- Sudan III and IV. 4, pigeon. Formaldehyde calcium/Sudan black B. 5, pigeon. Formaldehyde calcium/nile blue. 6, pigeon. Lewitsky saline/Sudan black B. 7, fowl. Carnoy/mercuric bromophenol blue. 8, pigeon. Formaldehyde calcium/acid haematein- Sudan III and IV . 9, fowl. Formaldehyde calcium (unpostchromed)-acetone extraction/Sudan black B . 10, fowl. Formaldehyde calcium (unpostchromed) ethanol extraction Sudan black B. 11, pigeon. Formaldehyde calcium/nile blue. 12, pigeon. Formaldehyde osmium. 13, fowl. Kolatchev . 14, fowl. Formaldehyde calcium/acid haematein-Sudan III and IV. 15, pigeon. Formaldehyde calcium/acid haematein-Sudan III and IV. 16, fowl. Formaldehyde calcium/nile blue . 17, fowl. Lewitsky saline,/Sudan black B. 18, pigeon. Lewitsky saline/Sudan black B. 19, fowl. Carnoy/mercuric bromophenol blue. 20, fowl. Formaldehyde calcium (unpostchromed) -acetone extraction/Sudan black B. 21, pigeon. Formaldehyde calcium (unpostchromed)- ethanol extraction/Sudan black B. 22,
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