The Golgi Apparatus with the Historical Point of View


  • Mukaddes EŞREFOĞLU

Received Date: 23.07.2020 Accepted Date: 30.10.2020 Bezmialem Science 2021;9(3):369-372

Scientists were introduced to the Golgi apparatus (GA) in 1898, when it was discovered by Camillo Golgi in 1898 as a “cytoplasmic reticular network”. Researchers heard Camillo Golgi’s name not only because of the GA, but also because of many definitions such as Golgi silver impregnation techniques, Golgi type I and II cells, Golgi cells of the cerebellum, and Golgi tendon organ. In fact, although the GA beared the name of this scientist, many scientists did numerous studies on the morphological and functional properties of this unique organelle before him, simultaneously with him or after him. Despite the simple technical possibilities of the old times, the scientists, whom we gratefully commemorated, obtained magnificent findings about the GA and presented them to the world of science. In this short article, which was a review, the following historical developments, starting from the discovery of the GA, were summarized.

Keywords: Camillo Golgi, Golgi apparatus, discovery, historical narrative

Discovery of Golgi Apparatus and Historical Developments

The Golgi apparatus (GA), defined as the “post office of the cell”, was first described in history about 100 years ago by the Italian doctor and pathologist Camillo Golgi, who was known for his studies on the nervous system (1844-1926). Professor Golgi was working at the University of Pavia, the oldest and most respected university in Italy, founded in 1361. Camillo Golgi and Spanish Anatomist Ramón y Cajal (1852-1934) shared the Nobel Prize in Physiology and Medicine in 1906 for their separate studies on the anatomy of the nervous system (1). The first award in this field was given to the German physiologist Emil Adolf von Behring, who discovered serum therapy in the development of diphtheria and tetanus vaccines in 1901. Serum therapy was interpreted as opening a new path in the field of Medical Sciences. Emil Adolf von Behring went down in history with the sentence that “a victorious weapon was placed in the hands of the doctor against sickness and death” (2).

Camillo Golgi developed a new technique in 1873 that stained neurons and special cells in the nervous system, thus allowing them to be marked. This technique was called the ‘black reaction’ because it stained the cell bodies and extensions of the nervous system black. With his technique, Golgi determined that the axons he clearly saw formed an uninterrupted network carrying nerve impulses (1). Today, the scientific writings of Camillo Golgi, including original drawings, are exhibited in the History Museum of the University of Pavia. Golgi has studies on various parts of the brain. However, his studies on the cerebellum, olfactory bulb, hippocampus and cerebral cortex are particularly important. For example, Golgi cells, which were named after him, were identified in the cerebellum (3). While Camillo Golgi’s work on the nervous system continued, his rival, Santiago Ramo’n y Cajal, was also on the rise in the scientific world. Cajal developed a theory that challenged Golgi. While this theory, known as the “neuron theory”, was generally supported by scientists, Golgi’s work was criticized (4). These criticisms did not deter Camillo Golgi. While examining the spinal ganglia in 1897, he noticed that there was a cytoplasmic network in their cell bodies, although not in every cell. Later, he detected the same reticulated structure in the cytoplasm of Purkinje cells of the genus Tyto alba owl (barn owl) (3). Yet this new and strange structure was not stained in every cell. Therefore, he shared his observation with his assistant Emilio Veratti (1872-1967). Emilio Veratti (5), who described the sarcoplasmic reticulum in 1902, confirmed its existence by showing this cytoplasmic network in the 4th cranial nerve. He made the first official presentation of this structure, which he described as the “internal reticulated structure”, on 19 April 1898 at the Pavia Medical-Surgical Society (6,7). He stated that this newly discovered reticulated  structure consisted of anastomosing ribbon-shaped filamentous elements, small plates with a clear center that served as the nodal points of the reticulum, and rounded discs (8). Antonio Pensa (1874-1970), working in the General Pathology and Histology laboratory of Golgi in 1899, detected this organelle in the cells of the adrenal medulla (9). A short time later, 5th year medical student Adelchi Negri (1876-1912) demonstrated the presence of a similar structure in thyroid, epididymis, salivary glands, and ovarian cells besides nerve cells (10). Negri incidentally detected intraneuronal inclusions while examining rabies-infected brains. These inclusions are known as "Negri bodies". (8). Meanwhile, Edoardo Gemelli (1878-1959), one of Golgi’s students, showed that a similar structure was found in the cells of the pituitary gland (11). From these observations it became clear that this structure was probably ubiquitous in eukaryotic cell types. Camillo Golgi hypothesized that this cytoplasmic network might be related to secretory function, more broadly to cell nutrition (8). Camillo Golgi changed the technique defined by Ramón y Cajal in 1903 and developed a new technique (12). With the advantages of this new technique, he was able to observe the morphology and localization of the GA during the secretion process in the mucous glands of the frog stomach. Thus, he found that the GA was located in the apical cytoplasm above the nucleus (13). Camillo Golgi tried to explain the physiological role of the GA in gastric and intestinal mucous cells (14). Meanwhile, Giuseppe D’Agata (1927-2011) was investigating this reticular network in the gastric epithelium (15).

In fact, although the GA was defined by being inspired by the work of Camillo Golgi, between 1867 and 1887, various scientists talked about the reticular structures existing in the cell from time to time before or after Camillo Golgi introduced this organelle (3,4,16,17). Perhaps these researchers also observed the GA. Yet all these years the GA was considered almost entirely a specific subject of the University of Pavia. Camillo Golgi and his students published more than 70 articles on this organelle. In these articles, they reported the changes observed in the GA in various developmental, physiological and pathological conditions, as well as the wide variety of cell types they observed (18). Thus, the organelle was found to be highly unstable and variable. Despite all these studies, the authenticity of this organelle was questioned by many researchers in the following years. Scientists defined this structure as an unreal structure that occured due to fixation or metallic impregnation technique (3). For example, George Palade and Albert Claude, scientists of the Rockefeller Institute, who showed similar cytoplasmic structures 20 years later in various cells without applying special staining methods using 40-55% ethanol, suggested based on these observations that GA represented one or more myelin figures that emerged artificially during the preparation of cytological samples (19). These discussions continued even when the GA could not be demonstrated with the first electron microscopic examinations (20). Finally, the GA, which was observed electron microscopically in the mid-1950s, was accepted as a real organelle and gained the respect it deserved (3). Taking into account Palade’s suggestion that phosphate-buffered osmium tetroxide should be used, Bethesda National Cancer Institute scientists Albert Dalton and Felix (21) soon demonstrated the detailed electron microscopic structure of the GA in epididymis cells. These researchers described the organelle as a structure in the cytoplasm consisting of folded, smooth-surfaced sacs and numerous vesicles and vacuoles with the staining technique developed by Camillo Golgi (21).

For many years, scientists focused on morphology rather than the function of the GA. It was known almost from the beginning by light microscopic observations that this organelle developed well in secreting cells. However, the role of this organelle in secretion and glycosylation was not elucidated until the 1960s (22). Palade suggested that the GA was associated with the vectorial transport of secretory proteins in exocrine pancreatic cells, and that vesicular transport also occured between the sacs of this organelle (23). Fleischer et al. (24), Morre et al. (25), and Neutra and Leblond (26) emphasized the important role of the GA in glycoprotein synthesis. The results of the autoradiographic studies of Godman and Lane showing the uptake of sulfate into the GA suggested that this organelle had an important role in sulfation and therefore in glycoprotein biosynthesis (27). Between 1967-1975, various researchers conducted numerous studies on the role of the GA in the secretory pathway and vesicular transport (28-30). In the 1980s, studies were carried out emphasizing the importance of mannose-6 phosphate in the exiting of lysosomal enzymes from the GA and in targeting them (31-34). Between 1981 and 1984, Rothman et al. studied substance transport in the Golgi sacs (35-37). The COPII protein cover was determined by Duden et al., Seratini et al., and Waters et al. in 1991, and by Barlow et al. in 1994 (22).

In parallel with the technological developments, it was possible to reach detailed information about the location of the GA, its morphological, functional and pathological features. Extraordinary new information is being obtained about morphological and functional properties of the GA with the discovery of new genes, the development of advanced technology techniques such as green fluorescent protein based live cell imaging techniques, dynamic live cell imaging techniques, high resolution electron microscopy techniques that can create three-dimensional structure, and correlative microscopy techniques (CLEM) (38,39). In particular, CLEM provides great advantages for obtaining new findings about substance trafficking, targeting and signaling mechanisms in Golgi sacs.

Naming the Golgi Apparatus

The discovery of the detailed morphological features of the GA with the use of electron microscopes caused this organelle to be given various names such as “Golgi body”, “Golgi zone”, “Golgi substance”, and “Golgi net”. In 1910, Carlo Besta named this organelle “Golgi apparatus” (40), but this name was officially entered into the scientific literature in 1913, using it in Nusbaum’s article (41). The definition of  “Golgi complex” entered the scientific literature in 1956 with the study of Dalton and Felix (42). Today, both the names “Golgi apparatus” and “Golgi complex” are used, but the “Golgi apparatus” is mostly preferred. Scientists are accustomed to the word “Golgi” not only because of this organelle, but also because of Golgi’s silver implantation techniques, Golgi type I and type II cells, Golgi cells in the cerebellum, and Golgi tendon organ. Especially in recent years, many new terms have entered the literature in parallel with the fact that techniques that provide detailed information and three-dimensional imaging provide new information about the features of this organelle and its region. These are definitions such as “Golgi receptor”, “Golgi strip”, “Golgi cluster”, “Golgi skeleton”, “Golgi sac”, “Golgi tubule”, “Golgi vesicle”, and “Golgi vacuole” (43). Although Camillo Golgi’s name was mentioned in these common uses, many researchers do not know who this scientist actually was, how, where and under what conditions he lived. This review article was written with respect to Camillo Golgi and the scientists in his team, who provided the recognition of the GA, despite the limited possibilities that could not be compared with today’s technological facilities, and other scientists who played very important roles in the recognition of the GA with their research, although their names were not remembered.

Peer-review: Externally peer reviewed.

Financial Disclosure: The author declared that this study received no financial support.


  1. Golgi C. The impossible interview with the man of the hidden biological structures. Interview by Paolo Mazzarello. J Hist Neurosci 2006;15:318-25. 
  2. Raju TN. The Nobel chronicles. 1901: Emil Adolf von Behring (1854-1917). Lancet 1998;352:75. 
  3. Mazzarello P, Garbarino C, Calligaro A. How Camillo Golgi became “the Golgi”. FEBS Lett 2009;583:3732-7. 
  4. Dröscher A. Camillo Golgi and the discovery of the Golgi apparatus. Histochem Cell Biol 1998;109:425-30. 
  5. Mazzarello P, Calligaro A, Vannini V, Muscatello U. The sarcoplasmic reticulum: its discovery and rediscovery. Nat Rev Mol Cell Biol 2003;4:69-74. 
  6. Golgi C. On the structure of the nerve cells of the spinal ganglia. 1898. J Microsc 1989;155:9-14.
  7. Golgi C. On the structure of nerve cells. 1898. J Microsc 1989;155:3-7. 
  8. Mazzarello P, Bentivoglio M. The centenarian Golgi apparatus. Nature 1998;392:543-4. 
  9. Pensa A. Sopra una fina particolarità di struttura di alcune cellule dele capsule soprarenali. Bollettino della Società Medico-Chirurgica di Pavia 1899: 76–85.
  10. Negri A. Di una fina particolarità di struttura delle cellule di alcune ghiandole dei mammiferi. Bollettino della Società Medico-Chirurgica di Pavia 1900: 61–70.
  11. Gemelli E. Contributo alla conoscenza sulla struttura della ghiandola pituitaria nei mammiferi. Bollettino della Società Medico-Chirurgica di Pavia 1900; 231–40.
  12. Golgi C. Di un metodo per la facile e pronta dimostrazione dell’apparato reticolare interno delle cellule nervose. Bollettino della Società Medico-Chirurgica di Pavia 1908; 81-7.
  13. Golgi C. Sur une fine particularite´ de la structure de l’e´pithe´lium de la muqueuse gastrique et intestinale de quelques verte´bre´s. Arch Ital Biol 1909;51:213-45.
  14. D’Agata G. Sulle modificazioni dell’apparato reticolare interno nell’epitelio della mucosa gastrica. Bollettino della Società Medico-Chirurgica di Pavia 1910; 517-22.
  15. Trautmann JC. Camillo Golgi (1843–1926) und die Entdeckung des ‘‘apparato reticolare interno” Golgi-Apparat (Dissertation). Lübeck; 1988.
  16. Bentivoglio M, Mazzarello P. The pathway to the cell and its organelles: one hundred years of the Golgi apparatus. Endeavour 1998;22:101-5.
  17. Dröscher A. The history of the Golgi apparatus in neurones from its discovery in 1898 to electron microscopy. Brain Res Bull 1998;47:199-203.
  18. Palade GE, Claude A. The nature of the Golgi apparatus; identification of the Golgi apparatus with a complex of myelin figures. J Morphol 1949;85:71-111.
  19. Palade GE, Claude A. The nature of the Golgi apparatus; parallelism between intercellular myelin figures and Golgi apparatus in somatic cells. J Morphol 1949;85:35-69. 
  20. Pease DC, Baker RF. Electron microscopy of nervous tissue. Anat Rec 1951;110:505-29. 
  21. Dalton AJ, Felix MD. Cytologic and cytochemical characteristics of the Golgi substance of epithelial cells of the epididymis in situ, in homogenates and after isolation. Am J Anat 1954;94:171-207. 
  22. Farquhar MG, Palade GE. The Golgi apparatus: 100 years of progress and controversy. Trends Cell Biol 1998;8:2-10. 
  23. Palade G. Intracellular aspects of the process of protein synthesis. Science. 1975;189:347-58.
  24. Fleischer B, Fleischer S, Ozawa H. Isolation and characterization of Golgi membranes from bovine liver. J Cell Biol 1969;43:59-79.
  25. Morre J, Merlin LM, Keenan TW. Localization of glycosyl transferase activities in a Golgi apparatus-rich fraction isolated from rat liver. Biochem Biophys Res Commun 1969;37:813-9. 
  26. Neutra M, Leblond CP. Synthesis of the carbohydrate of mucus in the golgi complex as shown by electron microscope radioautography of goblet cells from rats injected with glucose-H3. J Cell Biol 1966;30:119-36. 
  27. Godman GC, Lane N. On The Site Of Sulfation In The Chondrocyte. J Cell Biol 1964;21:353-66.
  28. Jamieson JD, Palade GE. Synthesis, intracellular transport, and discharge of secretory proteins in stimulated pancreatic exocrine cells. J Cell Biol 1971;50:135-58.
  29. Kern HF, Jamieson JD, Palade GE. Der Einfluss von Kobaltchlorid auf den Sekretionsprozess der exokrinen Pankreaszelle [Effect of cobalt chloride on secretory process of the exocrine pancreatic cell]. Verh Anat Ges 1972;67:501-5. 
  30. Tartakoff AM, Jamieson JD, Scheele GA, Palade GE. Studies on the pancreas of the guinea pig. Parallel processing and discharge of exocrine proteins. J Biol Chem 1975;250:2671-7. 
  31. Sly WS. Receptor-mediated transport of acid hydrolases to lysosomes. Curr Top Cell Regul 1985;26:27-38. 
  32. Willingham MC, Pastan IH, Sahagian GG, Jourdian GW, Neufeld EF. Morphologic study of the internalization of a lysosomal enzyme by the mannose 6-phosphate receptor in cultured Chinese hamster ovary cells. Proc Natl Acad Sci U S A 1981;78:6967-71.
  33. Gabel CA, Goldberg DE, Kornfeld S. Identification and characterization of cells deficient in the mannose 6-phosphate receptor: evidence for an alternate pathway for lysosomal enzyme targeting. Proc Natl Acad Sci U S A 1983;80:775-9. 
  34. Gabel CA, Goldberg DE, Kornfeld S. Lysosomal enzyme oligosaccharide phosphorylation in mouse lymphoma cells: specificity and kinetics of binding to the mannose 6-phosphate receptor in vivo. J Cell Biol 1982;95:536-42. 
  35. Rothman JE. The Golgi apparatus: roles for distinct ‘cis’ and ‘trans’ compartments. Ciba Found Symp 1982;(92):120-37. 
  36. Dunphy WG, Fries E, Urbani LJ, Rothman JE. Early and late functions associated with the Golgi apparatus reside in distinct compartments. Proc Natl Acad Sci U S A 1981;78:7453-7.
  37. Rothman JE, Miller RL, Urbani LJ. Intercompartmental transport in the Golgi complex is a dissociative process: facile transfer of membrane protein between two Golgi populations. J Cell Biol 1984;99:260-71.
  38. Polishchuk EV, Polishchuk RS. Analysis of Golgi complex function using correlative light-electron microscopy. Methods Cell Biol 2013;118:243-58.
  39. Eşrefoğlu M. Golgi apparatus: Morphology and function with recent facts. Bezmialem Science 2019;7:331-8.
  40. Besta C. Sull’apparato reticolare interno (apparato di Golgi) della cellula nervosa. Anatomischer Anzeiger 1910;36:476–86.
  41. Nusbaum J. Ueber den sogenannten inneren Golgischen Netzapparat und sein Verha¨ltnis zu den Mitochondrien, Chromidien und andern Zellstrukturen im Tierreich zusammenfassendes. Referat Arch Zellforsch 1913;10:359 –67.
  42. Dalton AJ, Felix MD. comparative study of the Golgi complex. J Biophys Biochem Cytol 1956;2:79-84. 
  43. Fabene PF, Bentivoglio M. 1898-1998: Camillo Golgi and “the Golgi”: one hundred years of terminological clones. Brain Res Bull 1998;47:195-8.