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Camillo Golgi
Camillo Golgi
(Italian: [kaˈmillo ˈɡɔldʒi]; 7 July 1843 – 21 January 1926) was an Italian physician and pathologist known for his works on the central nervous system. He studied medicine at the University of Pavia
Pavia
(where he later spent most of his professional career) between 1860 and 1868 under the tutelage of Cesare Lombroso. Inspired by pathologist Giulio Bizzozero, he pursued research in nervous system. His discovery of a staining technique called black reaction (sometimes called Golgi's method
Golgi's method
or Golgi's staining in his honour) in 1873 was a major breakthrough in neuroscience. Several structures and phenomena in anatomy and physiology are named for him, including the Golgi apparatus, the Golgi tendon organ
Golgi tendon organ
and the Golgi tendon reflex. He is recognized as the greatest neuroscientist and biologist of his time.[1] Golgi and a Spanish biologist Santiago Ramón y Cajal
Santiago Ramón y Cajal
were jointly given the Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine
1906 "in recognition of their work on the structure of the nervous system".[2]

Contents

1 Biography

1.1 Personal life

2 Contributions

2.1 Black reaction or Golgi's staining 2.2 Nervous system 2.3 Kidney 2.4 Malaria 2.5 Cell organelle

3 Awards and legacy

3.1 Monuments in Pavia

4 Eponyms 5 See also 6 References 7 Further reading 8 External links

Biography[edit] Camillo Golgi
Camillo Golgi
was born in July 1843 in the village of Corteno, in the province of Brescia (Lombardy), Italy. The village is now named Corteno Golgi
Corteno Golgi
in his honour. His father Allessandro Golgi was a physician and district medical officer, originally from Pavia. In 1860, he entered the University of Pavia
Pavia
to study medicine, and earned his medical degree in 1865. 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 investigating cholera epidemic in villages around Pavia. In 1867, he resumed his academic study under the supervision of Cesare Lombroso. Lombroso was a renowned scientist in medical psychology such as genius, madness and criminality. Inspired by Lombroso, Golgi wrote a thesis on the etiology of mental disorders, from which he obtained his M.D. in 1868.[3] 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 of bone 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.[4] By 1872, Golgi was an established clinician and histopathologist. He, however, had no opportunity as a tenured professor in Pavia
Pavia
to pursue teaching and research in neurology.[3] Financial pressure prompted him to join the Hospital of the Chronically Ill (Pio Luogo degli Incurabili) in Abbiategrasso, 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 the Golgi's method). He published his major works between 1875 and 1885 in the journal Rivista sperimentale di Freniatria e di medicina legale.[5] In 1785, he joined the faculty of histology at the University of Pavia. In 1879, he was appointed Chair of Anatomy
Anatomy
at the University of Siena. But the next year, he returned to the University of Pavia
Pavia
as full Professor of histology.[6] 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
Pavia
twice, first between 1893 and 1896, and second between 1901 and 1909. During the First World War
First 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.[3] Personal life[edit] Golgi and his wife Lina Aletti had no children, and they adopted Golgi's niece Carolina.[4] Golgi was irreligious in his later life and became an agnostic atheist. One of his former students attempted an unsuccessful deathbed conversion on him.[7][8] Contributions[edit] Black reaction or Golgi's staining[edit]

The first illustration by Golgi of the nervous system. Vertical section of the olfactory bulb of a dog (in 1875).

Central nervous system
Central nervous system
was difficult to study during Golgi's time because the cells were hard to identify. The available tissue staining techniques were useless for studying nervous tissue. While working as chief medical officer at the Hospital of the Chronically Ill, he experimented with metal impregnation of nervous tissue, using mainly silver (silver staining). In the 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 microscope, the outline of the neuron became distinct from the surrounding tissue and cells. The silver chromate precipitate, as a reaction product, only selective stains some cellular components randomly, sparing other cell parts. The silver chromate particles create a stark black deposit on the soma (nerve cell body) as well as on the axon and all dendrites, providing an exceedingly clear and well-contrasted picture of neuron against a yellow background. This makes it easier to trace the structure of the nerve cells in the brain for the first time.[4] Since cells are selective stained in black, he called the process la reazione nera ("the black reaction"), but today it is called Golgi's method
Golgi's method
or the Golgi stain.[9] 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 the Gazzeta Medica Italiani on 2 August 1873.[10] Nervous system[edit]

Drawing by Camillo Golgi
Camillo Golgi
of a hippocampus stained with the silver nitrate method.

In 1871, a German anatomist Joseph von Gerlach
Joseph 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, namely axon from the dendrites. 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 the reticular theory which states that the brain is a single network of nerve fibres, and not of discrete cells.[11][12] Although Golgi's earlier works between 1873 and 1885 clearly depicted the axonal connections of cerebellar cortex and olfactory 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.[13][14] Golgi's theory was challenged by Ramón y Cajal, who used the same technique developed by Golgi. According to Ramón y Cajal's neurone theory, the nervous system is but a collection of individual cells, the neurones, which are interconnected to form a network.[15] In addition to this, Golgi was the first to give clear descriptions of the structure of the cerebellum, hippocampus, spinal cord, olfactory lobe, as well as striatal and cortical lesions in a case of chorea. In 1878, he also discovered a receptor organ that senses changes in muscle tension, and is now known as Golgi tendon organ
Golgi tendon organ
or Golgi receptor; and Golgi-Mazzoni corpuscles (pressure transductors).[16] He further developed a stain specific for myelin (a specialised portion of axon) using potassium dichromate and mercuric chloride. Using this he discovered the myelin annular apparatus, often called the horny funnel of Golgi-Rezzonico.[3] Kidney[edit] Golgi studied kidney function during 1882 to 1889. In 1882, he published his observations on the mechanism of renal 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 from tubulointerstitial nephritis, and he noted that the process was an essential part of repairing the kidney tissue. He was the first to dissect out intact nephrons, and show that the distal tubulus (loop of Henle) of the nephron returns to its originating glomerulus, 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).[17] Malaria[edit] A French Army physician Charles Louis Alphonse Laveran
Charles Louis Alphonse Laveran
discovered that malaria was caused by microscopic parasite (now called Plasmodium falciparum) in 1880. But scientists were sceptical until Golgi intervened. It was Golgi who helped him prove that malarial parasite was a microscopic protozoan. From 1885, Golgi studied the malarial parasite and its transmission. He established two types of malaria, tertian and quartan fevers caused by Plasmodium vivax
Plasmodium vivax
and Plasmodium malariae respectively.[18] In 1886, he discovered that malarial fever (paroxysm) was produced by the asexual stage in the human blood (called erythocytic cycle, or Golgi cycle).[19] In 1889-1890, Golgi and Ettore Marchiafava
Ettore Marchiafava
described the differences between benign tertian malaria and malignant tertian malaria (the latter caused by P. falciparum). By 1898, along with Giovanni Battista Grassi, Amico Bignami, Giuseppe Bastianelli, Angelo Celli
Angelo Celli
and Marchiafava, he confirmed that malaria was transmitted by Anopheline mosquito.[20] Cell organelle[edit] An organelle in eukaryotice cells now known as Golgi apparatus
Golgi apparatus
or Golgi complex, or sometimes simply as Golgi, was discovered by Camillo Golgi.[21] Golgi modified his black reaction using osmium dichromate solution with which he stained the nerve cells (Purkinje cells) of the cerebellum of an owl.[22] He noticed thread-like networks inside the cells and named them apparato 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.[23] After the same was confirmed by his assistant Emilio Veratti, he published it in the Bollettino della Società medico-chirurgica di Pavia.[24] 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.[22] It was only firmly established after 50 years of its discovery, when electron microscopes were developed.[25] Awards and legacy[edit] Golgi, together with Santiago Ramón y Cajal, received the Nobel Prize in Physiology
Physiology
or Medicine in 1906 for his studies of the structure of the nervous system. In 1900 he was named senator by King Umberto I.[26] In 1913 he became foreign member of the Royal Netherlands Academy of Arts and Sciences.[27] He received honorary doctorate from the University of Cambridge, University of Geneva, Kristiania University College, National and Kapodistrian University of Athens, and Paris-Sorbonne University. He was commemorated in stamp by European Community in 1994.[16] Monuments in Pavia
Pavia
[edit]

Marble statue of Golgi at the University of Pavia

In Pavia
Pavia
several landmarks stand as Golgi’s memory.

A marble statue, in a yard of the old buildings of the University of Pavia, at N.65 of the central "Strada Nuova". On the basement, there is the following inscription in Italian language: " Camillo Golgi
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
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 hundreds 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
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
Bartolomeo Panizza
and Adelchi Negri. Golgi's museum was created in 2012, in the ancient Palazzo Botta of the University of Pavia
Pavia
at N.10 of Piazza Antoniotto Botta reconstructs the study of Camillo Golgi
Camillo Golgi
and its laboratories with furniture and original instruments.[28]

Eponyms[edit]

The organelle Golgi apparatus
Golgi apparatus
or Golgi complex The sensory receptor Golgi tendon organ Golgi's method
Golgi's method
or Golgi stain, a nervous tissue staining technique The enzyme Golgi alpha-mannosidase II Golgi cells of the cerebellum Golgi I nerve cells (with long axons) Golgi II nerve cells (with short or no axons)

See also[edit]

List of pathologists

References[edit]

^ Gerd Kempermann MD (2001). Adult Neurogenesis (2nd ed.). Oxford University Press. p. 616. ISBN 978-0199729692.  ^ "The Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine
1906". www.nobelprize.org. Retrieved 22 December 2017.  ^ a b c d Mazzarello, Paolo (1999). "Camillo Golgi's Scientific Biography". Journal of the History of the Neurosciences. 8 (2): 121–131. doi:10.1076/jhin.8.2.121.1836. PMID 11624293.  ^ a b c Bentivoglio, M. (2014). "Golgi, Camillo". In Daroff, Robert B.; Aminoff, Michael J. Encyclopedia of the Neurological Sciences (Second edition. ed.). Burlington: Elsevier Science. pp. 464–466. ISBN 9780123851581.  ^ Drouin, Emmanuel; Piloquet, Philippe; Péréon, Yann (2015). "The first illustrations of neurons by Camillo Golgi". The Lancet Neurology. 14 (6): 567. doi:10.1016/S1474-4422(15)00051-4. PMID 25987274.  ^ Zanobio, Bruno. " Camillo Golgi
Camillo Golgi
facts, information, pictures". www.encyclopedia.com. Retrieved 22 December 2017.  ^ Paolo Mazzarello; Henry A. Buchtel; Aldo Badiani (1999). The hidden structure: a scientific biography of Camillo Golgi. Oxford University Press. p. 34. ISBN 978-0-19-852444-1. 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. PMID 16995603.  ^ DeFelipe, Javier (2015). "The dendritic spine story: an intriguing process of discovery". Frontiers in Neuroanatomy. 9: 14. doi:10.3389/fnana.2015.00014. PMC 4350409 . PMID 25798090.  ^ Marina Bentivoglio (20 April 1998). "Life and Discoveries of Camillo Golgi". Nobelprize.org. Nobel Media. Retrieved 23 August 2013.  ^ Cimino G (1999). " Reticular theory
Reticular theory
versus neuron theory in the work of Camillo Golgi". Physis Riv Int Stor Sci. 36 (2): 431–472. PMID 11640243.  ^ Raviola E, Mazzarello P (2011). "The diffuse nervous network of Camillo Golgi: facts and fiction". Brain Res Rev. 66 (1–2): 75–82. doi:10.1016/j.brainresrev.2010.09.005. PMID 20840856.  ^ Marina Bentivoglio (20 April 1998). "Life and Discoveries of Camillo Golgi". Nobelprize.org. Nobel Media. Retrieved 23 August 2013.  ^ Bock, Ortwin (2013). "Cajal, Golgi, Nansen, Schäfer and the Neuron Doctrine". Endeavour. 37 (4): 228–234. doi:10.1016/j.endeavour.2013.06.006. PMID 23870749.  ^ a b Mazzarello, P. (1998). " Camillo Golgi
Camillo Golgi
(1843-1926)". Journal of Neurology, Neurosurgery & Psychiatry. 64 (2): 212–212. doi:10.1136/jnnp.64.2.212.  ^ Dal Canton, Ilaria; Calligaro, Alessandro L.; Dal Canton, Francesca; Frosio-Roncalli, Moris; Calligaro, Alberto (1999). "Contributions of Camillo Golgi
Camillo Golgi
toRenal Histology and Embryology". American Journal of Nephrology. 19 (2): 304–307. doi:10.1159/000013465. PMID 10213832.  ^ 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.  ^ Antinori, Spinello; Galimberti, Laura; Milazzo, Laura; Corbellino, Mario (2012). "Biology of human malaria plasmodia including Plasmodium knowlesi". Mediterranean Journal of Hematology and Infectious Diseases. 4 (1): 2012013. doi:10.4084/MJHID.2012.013. PMC 3340990 . PMID 22550559.  ^ Cox, Francis EG (2010). "History of the discovery of the malaria parasites and their vectors". Parasites & Vectors. 3 (1): 5. doi:10.1186/1756-3305-3-5. PMC 2825508 . PMID 20205846.  ^ 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. PMID 11624302.  ^ a b Dröscher, Ariane (1998). "The history of the golgi apparatus in neurones from its discovery in 1898 to electron microscopy". Brain Research Bulletin. 47 (3): 199–203. doi:10.1016/S0361-9230(98)00080-X. PMID 9865850.  ^ Mazzarello, Paolo; Garbarino, Carla; Calligaro, Alberto (2009). "How Camillo Golgi
Camillo Golgi
became "the Golgi"". FEBS Letters. 583 (23): 3732–3737. doi:10.1016/j.febslet.2009.10.018. PMID 19833130.  ^ Dröscher, A (1998). " Camillo Golgi
Camillo Golgi
and the discovery of the Golgi apparatus". Histochemistry and Cell Biology. 109 (5-6): 425–30. doi:10.1007/s004180050245. PMID 9681625.  ^ 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. PMID 10933465.  ^ GOLGI Camillo. Italian senate
Italian senate
website ^ "C. Golgi (1844 - 1926)". Royal Netherlands Academy of Arts and Sciences. Retrieved 19 July 2015.  ^ Spizzi, Dante. "Museo Camillo Golgi". museocamillogolgi.unipv.eu (in Italian). Retrieved 23 December 2017. 

Further reading[edit]

Mazzarello, Paolo (2010), Golgi: A Biography of the Founder of Modern Neuroscience, translated by Badiani, Aldo; Buchtel, Henry A., New York: Oxford University Press, ISBN 9780195337846  Mironov, Alexander A.; Margit, Pavelka (2006). The Golgi Apparatus State of Art After 110 Years of Camillo's Discovery. Dordrecht: Springer. ISBN 3211763104.  Morré, D. James; Mollenhauer, Hilton H. (2009). The Golgi Apparatus: The First 100 Years. New York: Springer. ISBN 0387743472.  De Carlos, Juan A; Borrell, José (2007), "A historical reflection of the contributions of Cajal and Golgi to the foundations of neuroscience.", Brain Research Reviews (published August 2007), 55 (1), pp. 8–16, doi:10.1016/j.brainresrev.2007.03.010, PMID 17490748  Muscatello, Umberto (2007), "Golgi's contribution to medicine.", Brain Research Reviews (published August 2007), 55 (1), pp. 3–7, doi:10.1016/j.brainresrev.2007.03.007, PMID 17462742  Kruger, Lawrence (2007), "The sensory neuron and the triumph of Camillo Golgi", Brain Research Reviews (published October 2007), 55 (2), pp. 406–10, doi:10.1016/j.brainresrev.2007.01.008, PMID 17408565  Fabene, P F; Bentivoglio, M (1998), "1898–1998: Camillo Golgi
Camillo Golgi
and "the Golgi": one hundred years of terminological clones.", Brain Res. Bull. (published October 1998), 47 (3), pp. 195–8, doi:10.1016/S0361-9230(98)00079-3, PMID 9865849  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, 40 (6), pp. 483–96, PMID 9778732  Farquhar, M G; Palade, G E (1998), "The Golgi apparatus: 100 years of progress and controversy.", Trends Cell Biol. (published January 1998), 8 (1), pp. 2–10, doi:10.1016/S0962-8924(97)01187-2, PMID 9695800 

Wikimedia Commons has media related to Camillo Golgi.

External links[edit]

Life and Discoveries of Camillo Golgi The Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine
1906 Some places and memories related to Camillo Golgi The museum in Corteno, now called Corteno Golgi, dedicated to Golgi. Includes a gallery of images of his birthplace. The Museum Camillo Golgi
Camillo Golgi
in Pavia Biography at Encyclopædia Britannica Biography at Encyclopedia.com Profile at Whonamedit?

v t e

Laureates of the Nobel Prize in Physiology
Physiology
or Medicine

1901–1925

1901 Emil Behring 1902 Ronald Ross 1903 Niels Finsen 1904 Ivan Pavlov 1905 Robert Koch 1906 Camillo Golgi
Camillo Golgi
/ Santiago Ramón y Cajal 1907 Alphonse Laveran 1908 Élie Metchnikoff
Élie Metchnikoff
/ Paul Ehrlich 1909 Emil Kocher 1910 Albrecht Kossel 1911 Allvar Gullstrand 1912 Alexis Carrel 1913 Charles Richet 1914 Róbert Bárány 1915 1916 1917 1918 1919 Jules Bordet 1920 August Krogh 1921 1922 Archibald Hill
Archibald Hill
/ Otto Meyerhof 1923 Frederick Banting
Frederick Banting
/ John Macleod 1924 Willem Einthoven 1925

1926–1950

1926 Johannes Fibiger 1927 Julius Wagner-Jauregg 1928 Charles Nicolle 1929 Christiaan Eijkman
Christiaan Eijkman
/ Frederick Gowland Hopkins 1930 Karl Landsteiner 1931 Otto Warburg 1932 Charles Scott Sherrington
Charles Scott Sherrington
/ Edgar Adrian 1933 Thomas Morgan 1934 George Whipple
George Whipple
/ George Minot
George Minot
/ William Murphy 1935 Hans Spemann 1936 Henry Dale / Otto Loewi 1937 Albert Szent-Györgyi 1938 Corneille Heymans 1939 Gerhard Domagk 1940 1941 1942 1943 Henrik Dam
Henrik Dam
/ Edward Doisy 1944 Joseph Erlanger
Joseph Erlanger
/ Herbert Gasser 1945 Alexander Fleming
Alexander Fleming
/ Ernst Chain
Ernst Chain
/ Howard Florey 1946 Hermann Muller 1947 Carl Cori / Gerty Cori
Gerty Cori
/ Bernardo Houssay 1948 Paul Müller 1949 Walter Hess / António Egas Moniz 1950 Edward Kendall / Tadeusz Reichstein
Tadeusz Reichstein
/ Philip Hench

1951–1975

1951 Max Theiler 1952 Selman Waksman 1953 Hans Krebs / Fritz Lipmann 1954 John Enders / Thomas Weller / Frederick Robbins 1955 Hugo Theorell 1956 André Cournand / Werner Forssmann
Werner Forssmann
/ Dickinson W. Richards 1957 Daniel Bovet 1958 George Beadle / Edward Tatum
Edward Tatum
/ Joshua Lederberg 1959 Severo Ochoa
Severo Ochoa
/ Arthur Kornberg 1960 Frank Burnet / Peter Medawar 1961 Georg von Békésy 1962 Francis Crick
Francis Crick
/ James Watson
James Watson
/ Maurice Wilkins 1963 John Eccles / Alan Hodgkin / Andrew Huxley 1964 Konrad Bloch / Feodor Lynen 1965 François Jacob
François Jacob
/ André Lwoff / Jacques Monod 1966 Francis Rous / Charles B. Huggins 1967 Ragnar Granit
Ragnar Granit
/ Haldan Hartline / George Wald 1968 Robert W. Holley
Robert W. Holley
/ Har Khorana / Marshall Nirenberg 1969 Max Delbrück
Max Delbrück
/ Alfred Hershey
Alfred Hershey
/ Salvador Luria 1970 Bernard Katz / Ulf von Euler
Ulf von Euler
/ Julius Axelrod 1971 Earl Sutherland Jr. 1972 Gerald Edelman
Gerald Edelman
/ Rodney Porter 1973 Karl von Frisch
Karl von Frisch
/ Konrad Lorenz
Konrad Lorenz
/ Nikolaas Tinbergen 1974 Albert Claude
Albert Claude
/ Christian de Duve
Christian de Duve
/ George Palade 1975 David Baltimore
David Baltimore
/ Renato Dulbecco
Renato Dulbecco
/ Howard Temin

1976–2000

1976 Baruch Blumberg / Daniel Gajdusek 1977 Roger Guillemin / Andrew Schally
Andrew Schally
/ Rosalyn Yalow 1978 Werner Arber
Werner Arber
/ Daniel Nathans
Daniel Nathans
/ Hamilton O. Smith 1979 Allan Cormack / Godfrey Hounsfield 1980 Baruj Benacerraf / Jean Dausset
Jean Dausset
/ George Snell 1981 Roger Sperry / David H. Hubel
David H. Hubel
/ Torsten Wiesel 1982 Sune Bergström
Sune Bergström
/ Bengt I. Samuelsson / John Vane 1983 Barbara McClintock 1984 Niels Jerne / Georges Köhler / César Milstein 1985 Michael Brown / Joseph L. Goldstein 1986 Stanley Cohen / Rita Levi-Montalcini 1987 Susumu Tonegawa 1988 James W. Black / Gertrude B. Elion
Gertrude B. Elion
/ George H. Hitchings 1989 J. Michael Bishop
J. Michael Bishop
/ Harold E. Varmus 1990 Joseph Murray
Joseph Murray
/ E. Donnall Thomas 1991 Erwin Neher
Erwin Neher
/ Bert Sakmann 1992 Edmond Fischer / Edwin G. Krebs 1993 Richard J. Roberts
Richard J. Roberts
/ Phillip Sharp 1994 Alfred G. Gilman
Alfred G. Gilman
/ Martin Rodbell 1995 Edward B. Lewis
Edward B. Lewis
/ Christiane Nüsslein-Volhard
Christiane Nüsslein-Volhard
/ Eric F. Wieschaus 1996 Peter C. Doherty
Peter C. Doherty
/ Rolf M. Zinkernagel 1997 Stanley B. Prusiner 1998 Robert F. Furchgott
Robert F. Furchgott
/ Louis Ignarro
Louis Ignarro
/ Ferid Murad 1999 Günter Blobel 2000 Arvid Carlsson
Arvid Carlsson
/ Paul Greengard
Paul Greengard
/ Eric Kandel

2001–present

2001 Leland H. Hartwell / Tim Hunt
Tim Hunt
/ Paul Nurse 2002 Sydney Brenner
Sydney Brenner
/ H. Robert Horvitz / John E. Sulston 2003 Paul Lauterbur
Paul Lauterbur
/ Peter Mansfield 2004 Richard Axel
Richard Axel
/ Linda B. Buck 2005 Barry Marshall
Barry Marshall
/ Robin Warren 2006 Andrew Fire / Craig Mello 2007 Mario Capecchi
Mario Capecchi
/ Martin Evans
Martin Evans
/ Oliver Smithies 2008 Harald zur Hausen
Harald zur Hausen
/ Luc Montagnier
Luc Montagnier
/ Françoise Barré-Sinoussi 2009 Elizabeth Blackburn
Elizabeth Blackburn
/ Carol W. Greider
Carol W. Greider
/ Jack W. Szostak 2010 Robert G. Edwards 2011 Bruce Beutler
Bruce Beutler
/ Jules A. Hoffmann / Ralph M. Steinman (posthumously) 2012 John B. Gurdon
John B. Gurdon
/ Shinya Yamanaka 2013 James Rothman
James Rothman
/ Randy Schekman
Randy Schekman
/ Thomas C. Südhof 2014 John O'Keefe / May-Britt Moser
May-Britt Moser
/ Edvard Moser 2015 William C. Campbell / Satoshi Ōmura
Satoshi Ōmura
/ Tu Youyou 2016 Yoshinori Ohsumi 2017 Jeffrey C. Hall, Michael Rosbash, Michael W. Young

Authority control

WorldCat Identities VIAF: 57421035 LCCN: n80145398 ISNI: 0000 0001 1856 2501 GND: 119273896 SUDOC: 066912008 BNF: cb13746673x (data) NKC: nlk20000077323 ICCU: ITICCURAVV71

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