Golgi and the Spanish biologistSantiago Ramón y Cajal were jointly awarded the Nobel Prize in Physiology or Medicine in 1906 "in recognition of their work on the structure of the nervous system".[2]
Camillo Golgi was born on 7 July 1843 in the village of Corteno near Brescia, in theprovince of Brescia (Lombardy), at the time Kingdom of Lombardy–Venetia, today Italy. The village is now namedCorteno Golgi in his honour. His father Alessandro Golgi was a physician and district medical officer, originally from Pavia. In 1860, he entered theUniversity of Pavia to study medicine, and earned his medical degree in 1865.[3] He did an internship at the San Matteo Hospital (now IRCCS Policlinico San Matteo Foundation). During his internship he briefly worked as a civil physician in the Italian Army, and as assistant surgeon at the Novara Hospital (now Azienda Ospedaliero Universitaria Maggiore della Carità di Novara). At the same time he was also involved in the medical team for investigatingcholera epidemic in villages around Pavia.[4]
In 1867, he resumed his academic study under the supervision ofCesare Lombroso. Lombroso was a renowned scientist inmedical psychology such as genius, madness and criminality. Inspired by Lombroso, Golgi wrote a thesis on the etiology ofmental disorders, from which he obtained hisM.D. in 1868.[5] He became more interested in experimental medicine, and started attending the Institute of General Pathology headed by Giulio Bizzozero. Three years his junior, Bizzozero was an eloquent teacher and experimenter, who specialised in histology of the nervous system and the properties ofbone marrow. The most important research publications of Golgi were directly or indirectly influenced by Bizzozero. The two became so close that they lived in the same building; and Golgi later married Bizzozero's niece, Lina Aletti.[6] By 1872, Golgi was an established clinician and histopathologist. He, however, had no opportunity as a tenured professor in Pavia to pursue teaching and research in neurology.[5]
Financial pressure prompted him to join the Hospital of the Chronically Ill (Pio Luogo degli Incurabili) inAbbiategrasso, near Milan, as Chief Medical Officer in 1872. To continue research, he set up a simple laboratory on his own in a refurbished hospital kitchen, and it was there that he started making his most notable discoveries. His major achievement was the development of staining technique for nerve tissue called the black reaction (later theGolgi's method). He published his major works between 1875 and 1885 in the journalRivista sperimentale di Freniatria e di medicina legale.[7] In 1875, he joined the faculty of histology at the University of Pavia. In 1879, he was appointed Chair of Anatomy at theUniversity of Siena. But the next year, he returned to the University of Pavia as full Professor of histology.[8] From 1879 he also became Professor of General Pathology as well as Honorary Chief (Primario ad honorarem) at the San Matteo Hospital. He served as Rector of the University of Pavia twice, first between 1893 and 1896, and second between 1901 and 1909. During theFirst World War (1914–1917), he directed the military hospital Collegio Borrmeo at Pavia. He retired in 1918 and continued to research in his private laboratory till 1923. He died on 21 January 1926.[5]
Golgi and his wife Lina Aletti had no children, and they adopted Golgi's niece Carolina.[6]
Golgi was irreligious in his later life and became an agnostic atheist. One of his former students attempted an unsuccessfuldeathbed conversion on him.[9][10]
The first illustration by Golgi of the nervous system. Vertical section of the olfactory bulb of a dog (in 1875).
Thecentral nervous system was difficult to study during Golgi's time because the cells were hard to identify. The availabletissue staining techniques were useless for studyingnervous tissue. While working as chief medical officer at the Hospital of the Chronically Ill, he experimented with metal impregnation of nervous tissue, using mainlysilver (silver staining). In early 1873, he discovered a method of staining nervous tissue that would stain a limited number of cells at random in their entirety. He first treated the tissue with potassium dichromate to harden it, and then with silver nitrate. Under the microscope, the outline of the neuron became distinct from the surrounding tissue and cells. The silver chromate precipitate, as a reaction product, selectively stains only some cellular components randomly, sparing other cell parts. The silver chromate particles create a stark black deposit on thesoma (nerve cell body) as well as on theaxon and alldendrites, providing an exceedingly clear and well-contrasted picture ofneuron against a yellow background. This makes it easier to trace the structure of the nerve cells in the brain for the first time.[6] Since cells are selective stained in black, he called the processla reazione nera ("the black reaction"), but today it is calledGolgi's method or the Golgi stain.[11] On 16 February 1873, he wrote to his friend Niccolò Manfredi:
I am delighted that I have found a new reaction to demonstrate, even to the blind, the structure of the interstitial stroma of the cerebral cortex.
His discovery was published in theGazzeta Medica Italiani on 2 August 1873.[12]
Drawing by Camillo Golgi of ahippocampus stained with the silver nitrate method
In 1871, a German anatomistJoseph von Gerlach postulated that the brain is a complex "protoplasmic network", in the form of a continuous network called the reticulum. Using his black reaction, Golgi could trace various regions of the cerebro-spinal axis, clearly distinguishing the different nervous projections, namelyaxon from thedendrites. He drew up a new classification of cells on the basis of the structure of their nervous prolongation. He described an extremely dense and intricate network, composed of a web of intertwined branches of axons coming from different cell layers ("diffuse nervous network"). This network structure, which emerges from the axons, is essentially different from that hypothesized by Gerlach. It was the main organ of the central nervous system according to Golgi. Thus, Golgi presented thereticular theory which states that the brain is a single network of nerve fibres, and not of discrete cells.[13][14] Although Golgi's earlier works between 1873 and 1885 clearly depicted the axonal connections ofcerebellar cortex andolfactory bulb as independent of one another, his later works including the Nobel Lecture showed the entire granular layer of the cerebellar cortex occupied by a network of branching and anastomosing nerve processes. This was due to his strong conviction in the reticular theory.[15][13] Golgi's theory was challenged by Ramón y Cajal, who used the same technique developed by Golgi. According to Ramón y Cajal'sneurone theory, the nervous system is but a collection of individual cells, the neurones, which are interconnected to form a network.[16]
In addition to this, Golgi was the first to give clear descriptions of the structure of thecerebellum,hippocampus,spinal cord,olfactory lobe, as well asstriatal and cortical lesions in a case ofchorea. In 1878, he also discovered a receptor organ that senses changes in muscle tension, and is now known asGolgi tendon organ or Golgi receptor; and Golgi-Mazzoni corpuscles (pressure transductors).[17] He further developed a stain specific formyelin (a specialised membrane which wraps around the axon) usingpotassium dichromate andmercuric chloride. Using this he discovered the myelin annular apparatus, often called the horny funnel of Golgi-Rezzonico.[5]
Golgi studied kidney function during 1882 to 1889. In 1882, he published his observations on the mechanism ofrenal hypertrophy, which he understood to be due to renal cell proliferation. In 1884, he described tubular cell mitoses in the kidney of a person suffering fromtubulointerstitial nephritis, and he noted that the process was an essential part of repairing the kidney tissue. He was the first to dissect out intactnephrons, and show that the distal tubulus (loop of Henle) of the nephron returns to its originatingglomerulus, a finding that he published in 1889 ("Annotazioni intorno all'Istologia dei reni dell'uomo e di altri mammifieri e sull'istogenesi dei canalicoli oriniferi".Rendiconti R. Acad. Lincei 5: 545–557, 1889).[18]
A French Army physicianCharles Louis Alphonse Laveran discovered thatmalaria was caused by microscopic parasite (now calledPlasmodium falciparum) in 1880. But scientists were sceptical until Golgi intervened. It was Golgi who helped him prove that malarial parasite was a microscopicprotozoan. From 1885, Golgi studied the malarial parasite and its transmission. He established two types of malaria, tertian and quartanfevers caused byPlasmodium vivax andPlasmodium malariae respectively.[19] In 1886, he discovered that malarial fever (paroxysm) was produced by the asexual stage in the human blood (called erythrocytic cycle, or Golgi cycle).[20] In 1889–1890, Golgi andEttore Marchiafava described the differences between benign tertian malaria and malignant tertian malaria (the latter caused byP. falciparum). By 1898, along withGiovanni Battista Grassi,Amico Bignami,Giuseppe Bastianelli,Angelo Celli and Marchiafava, he confirmed that malaria was transmitted byanopheline mosquitoes.[21]
An organelle in eukaryotic cells now known asGolgi apparatus or Golgi complex, or sometimes simply as Golgi, was discovered by Camillo Golgi.[22] Golgi modified his black reaction using osmium dichromate solution with which he stained the nerve cells (Purkinje cells) of the cerebellum of a barn owl.[23] He noticed thread-like networks inside the cells and named themapparato reticolare interno (internal reticular apparatus). Recognising them to be unique cellular components, he presented his discovery before the Medical-Surgical Society of Pavia in April 1898.[24] After the same was confirmed by his assistant Emilio Veratti, he published it in theBollettino della Società medico-chirurgica di Pavia.[25] However, most scientists disputed his discovery as nothing but a staining artefact. Their microscopes were not powerful enough to identify the organelles. By the 1930s, Golgi's description was largely rejected.[23] It was only firmly established 50 years after its discovery, when electron microscopes were developed.[26]
Golgi, together withSantiago Ramón y Cajal, received theNobel Prize in Physiology or Medicine in 1906 for his studies of the structure of the nervous system. Prior to the work that earned Cajal and Golgi their prize, there was a debate over how nerve impulses were conducted in the brain. One camp thought that neurons fused (between axon terminals of cell #1 and dendrites of cell #2), so that the brain essentially became one contiguous cell, a superneuron; this would result in asyncytium, forming the basis of the obsoletereticular theory. The other camp thought that neurons didn't fuse, but met in very close proximity, and this was called theneuron doctrine. Resolution of the cellular components at the point of interaction between cells was impossible at the time, and would only become possible much later with the invention of theelectron microscope in the 1950s. But with Golgi's stain, Cajal was able to decisively demonstrate the outline of a complete neuron, to the exclusion of adjacent neurons, thereby demonstrating that a universal neural syncytium did not exist. Cajal and Golgi, being from opposing camps, came to hate each other and would not speak. When they arrived inStockholm to receive their joint prize, Cajal managed to put their differences aside enough to praise Golgi, but Golgi did not reciprocate.[28]
In 1900 he was namedsenator by KingUmberto I.[29] In 1913 he became foreign member of theRoyal Netherlands Academy of Arts and Sciences.[30] He received honorary doctorates from the University of Cambridge, University of Geneva, Kristiania University College, National and Kapodistrian University of Athens, and Paris-Sorbonne University. In 1994, the European Community commemorated him with postage stamps.[17]
Marble statue of Golgi at the University of PaviaCamillo Golgi's house in Pavia
In Pavia several landmarks stand as Golgi's memory.
A marble statue, in a yard of the old buildings of theUniversity of Pavia, at N.65 of the central "Strada Nuova". On the basement, there is the following inscription in Italian language: "Camillo Golgi / patologo sommo / della scienza istologica / antesignano e maestro / la segreta struttura / del tessuto nervoso / con intenta vigilia / sorprese e descrisse / qui operò / qui vive / guida e luce ai venturi / MDCCCXLIII – MCMXXVI" (Camillo Golgi / outstanding pathologist / of histological science / precursor and master / the secret structure / of the nervous tissue / with strenuous effort / discovered and described / here he worked / here he lives / here he guides and enlightens future scholars / 1843 – 1926).
"Golgi’s home", also in Strada Nuova, at N.77, a few hundred meters away from the University, just in front to the historical "Teatro Fraschini". It is the home in which Golgi spent the most of his family life, with his wife Lina.
Golgi's tomb is in the Monumental Cemetery of Pavia (viale San Giovannino), along the central lane, just before the big monument to the fallen of the First World War. It is a very simple granite grave, with a bronze medallion representing the scientist's profile. Near Golgi's tomb, apart from his wife, two other important Italian medical scientists are buried:Bartolomeo Panizza andAdelchi Negri.
Golgi's museum was created in 2012, in the ancient Palazzo Botta of theUniversity of Pavia at N.10 of Piazza Antoniotto Botta reconstructs the study of Camillo Golgi and its laboratories with furniture and original instruments.[31]
^Paolo Mazzarello; Henry A. Buchtel; Aldo Badiani (1999). The hidden structure: a scientific biography of Camillo Golgi. Oxford University Press. p. 34.ISBN978-0-19-852444-1. It was probably during this period that Golgi became agnostic (or even frankly atheistic), remaining for the rest of his life completely alien to the religious experience.
^Rapport, Richard L. Nerve Endings: The Discovery of the Synapse. New York: W.W. Norton, 2005. Print.
^Chu, NS (2006). "[Centennial of the nobel prize for Golgi and Cajal—founding of modern neuroscience and irony of discovery]".Acta Neurologica Taiwanica.15 (3):217–222.PMID16995603.
^Dal Canton, Ilaria; Calligaro, Alessandro L.; Dal Canton, Francesca; Frosio-Roncalli, Moris; Calligaro, Alberto (1999). "Contributions of Camillo Golgi to Renal Histology and Embryology".American Journal of Nephrology.19 (2):304–307.doi:10.1159/000013465.PMID10213832.S2CID29666037.
^Golgi C. (1889). "Sul ciclo evolutivo dei parassiti malarici nella febbre terzana : diagnosi differenziale tra i parassiti endoglobulari malarici della terzana e quelli della quartana" [On the cycle of development of malarial parasites in tertian fever: differential diagnosis between the intracellular parasites of tertian and quartant fever].Archivio per le Scienza Mediche.13:173–196.
^Bentivoglio, Marina (1999). "The Discovery of the Golgi Apparatus".Journal of the History of the Neurosciences.8 (2):202–208.doi:10.1076/jhin.8.2.202.1833.PMID11624302.
^Bentivoglio, M; Mazzarello, P (1998). "One hundred years of the Golgi apparatus: history of a disputed cell organelle".Italian Journal of Neurological Sciences.19 (4):241–247.doi:10.1007/bf02427612.PMID10933465.S2CID31879493.
^Pauly, John Edward; Basmajian, John V.; Christensen, A. Kent; Jollie, William P.; Kelly, Douglas E., eds. (1987).The American Association of Anatomists, 1888-1987: essays on the history of anatomy in America and a report on the membership: past and present. Baltimore: Williams & Wilkins.ISBN978-0-683-06800-9.
^Sapolsky, RM. Behave: The Biology of Humans at our Best and Worst, New York: Penguin Books, 2017, p688
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Mironov, A A; Komissarchik, Ia Iu; Mironov, A A; Snigirevskaia, E S; Luini, A (1998), "[Current concept of structure and function of the Golgi apparatus. On the 100-anniversary of the discovery by Camillo Golgi]",Tsitologiia, vol. 40, no. 6, pp. 483–96,PMID9778732
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