GOLGI APPARATUS 



144 



GOLGI APPARATUS 



rounded by a zone of dense phospholipid 

 material and in some cases embedded 

 in a zone of diffuse phospholipid. 

 Palade and Claude (J. Morph. 1949, 85, 

 71) claim that the Golgi apparatus is 

 composed solely of neutral red staining 

 vacuoles which possess a membrane con- 

 taining an appreciable proportion of 

 lecithin and that the classical Golgi 

 apparatus is formed by myelin figures 

 produced by the action of fixatives on 

 these vesicles. Bensley (Exp. Cell. 

 Res. 1951, 2, 1-19) contends that the 

 Golgi apparatus is composed of vesicles 

 or canals containing a watery solution 

 in which various substances, e.g., 

 polysaccharides are dissolved (Gersh, 

 I., Arch. Path. 1949, 47, 99-109). The 

 canalicular conception of the Golgi ap- 

 paratus is also claimed by Gatenby and 

 Moussa for the mammalian neurone. 



Against these views we must put the 

 fact that no canals have been seen in 

 cells by numerous workers using the 

 phase contrast microscope and as 

 Hibbard, H. (Quart. Rev. Biol. 1945, 

 20, 1-19) has pointed out Lewis has 

 never seen anything resembling the 

 classical apparatus in a life-time of 

 study of tissue culture cells. Most 

 workers on the Golgi apparatus have 

 tended to range themselves behind one 

 or other of these views. It should be 

 realized, however, that all these points 

 of view about the structure of the ap- 

 paratus simply continue the con- 

 troversy which has gone on almost since 

 its discovery. The canalicular concep- 

 tion of the apparatus originated in 1902 

 with Holmgren's Trophospongium, the 

 neutral red vacuole theory found its 

 origin in Parat's 1928 work and has 

 seen some extension in Cowdry's labora- 

 tory by the work of Cowdry, E. V. and 

 Scott, G. H. (Arch. Inst. Pasteur de 

 Tunis, 1928, 233), Covell, W. P. and 

 Scott, G. H. (Anat. Rec, 1928, 38, 377), 

 and of course the argentophil network 

 theory originated with Golgi himself 

 in 1898 (Arch. Ital. de Biol., 30, 60) and 

 in more recent years has been supported 

 bj'^ Ludford and particularly by 

 Gatenby. So, it is apparent that our 

 knowledge of the nature of the Golgi 

 apparatus has remained more or less in 

 the same state as it was 30 or 40 

 years ago. 



In view of this discrepancy of views 

 techniques wall be given for demon- 

 strating the classical Golgi net and also 

 for demonstrating the apparatus as 

 discrete bodies. 



The Classical Golgi Apparatus: Even 

 if the Golgi network is an artefact as 

 some workers believe the conventional 



Golgi preparation does give us informa- 

 tion about its position in the cell and 

 whether the Golgi material is present in 

 small or in large amount. 



Golgi's original technique involved 

 the fixation of tissue in a mixture con- 

 taining potassium bichromate and 

 osmic acid followed by impregnation 

 with silver. The apparatus with this 

 technique appears jet black against a 

 yellowish background. It is a con- 

 spicuous structure consisting of an 

 intricate network of anastomosing 

 strands. This network may closely en- 

 velop the nucleus, be concentrated to 

 one side of it or else be scattered rather 

 diffusely throughout the cytoplasm. 

 In glandular cells the apparatus grows 

 in size with the development of secre- 

 tory granules and strands from it ramify 

 between the various granules. 



Kopsch, F. (Sitzungsber. K. Akad. 

 Wissensch, 1902, 40, 929) showed that 

 the Golgi apparatus can be blackened 

 by prolonged treatment with 2% osmic 

 acid. On this affinity for both silver 

 and osmium all the conventional meth- 

 ods of demonstrating the Golgi appara- 

 tus are based. Few cytological re- 

 actions are more fickle and incon- 

 stant; but when, after many attempts, 

 the technique is successful, convincing 

 and very beautiful preparations result. 

 Mitochondria can be stained supra- 

 vitally by some vital dyes but no vital 

 dyes will show up the reticular Golgi 

 apparatus, a fact which is taken by 

 some authors to indicate the non-exist- 

 ence in vivo of such networks. How- 

 ever, neutral red will stain spheres in 

 the region of the apparatus. 



With both silver and osmium methods 

 considerable experimentation is neces- 

 sary in order to obtain the best results. 

 The factors to be varied are principally 

 the composition of the fixative and im- 

 pregnating substance and the time 

 during which they are allowed to act. 

 During impregnation it is always ad- 

 visable to keep the tissues in the dark 

 and instructions as to temperature re- 

 quirements should be carefully fol- 

 lowed. When either the silver or 

 osmium solution becomes blackened 

 it should be renewed. It is important 

 for the beginner to start with the most 

 favorable material. The spinal 

 ganglion cells of young mammals such 

 as the rabbit are perhaps the best for 

 this purpose. The acinous cells of the 

 pancreas are also recommended but are 

 somewhat more difficult to handle. All 

 the methods of impregnation outlined 

 below frequently bring to light the 

 mitochondria also. 



