Nobel Prize in Physiology or Medicine

The Nobel Prize in Physiology or Medicine (Swedish: Nobelpriset i fysiologi eller medicin), administered by the Nobel Foundation, is awarded yearly for outstanding discoveries in the fields of life sciences and medicine. It is one of five Nobel Prizes established in his will in 1895 by Swedish chemist Alfred Nobel, the inventor of dynamite. Nobel was interested in experimental physiology and wanted to establish a prize for scientific progress through laboratory discoveries. The Nobel Prize is presented at an annual ceremony on 10 December, the anniversary of Nobel's death, along with a diploma and a certificate for the monetary award. The front side of the medal displays the same profile of Alfred Nobel depicted on the medals for Physics, Chemistry, and Literature. The reverse side is unique to this medal. The most recent Nobel prize was announced by Karolinska Institute on 1 October 2018, and has been awarded to American James P. Allison and Japanese Tasuku Honjo – for their discovery of cancer therapy by inhibition of negative immune regulation.[2]

As of 2015, 106 Nobel Prizes in Physiology or Medicine have been awarded to 198 men and 12 women. The first one was awarded in 1901 to the German physiologist Emil von Behring, for his work on serum therapy and the development of a vaccine against diphtheria. The first woman to receive the Nobel Prize in Physiology or Medicine, Gerty Cori, received it in 1947 for her role in elucidating the metabolism of glucose, important in many aspects of medicine, including treatment of diabetes.

Some awards have been controversial. This includes one to António Egas Moniz in 1949 for the prefrontal lobotomy, bestowed despite protests from the medical establishment. Other controversies resulted from disagreements over who was included in the award. The 1952 prize to Selman Waksman was litigated in court, and half the patent rights awarded to his co-discoverer Albert Schatz who was not recognized by the prize. The 1962 prize awarded to James D. Watson, Francis Crick and Maurice Wilkins for their work on DNA structure and properties did not acknowledge the contributing work from others, such as Oswald Avery and Rosalind Franklin who had died by the time of the nomination. Since the Nobel Prize rules forbid nominations of the deceased, longevity is an asset, considering prizes are awarded as long as 50 years after the discovery. Also forbidden is awarding any one prize to more than three recipients. In the last half century there has been an increasing tendency for scientists to work as teams, resulting in controversial exclusions.

The Nobel Prize in Physiology or Medicine
A golden medallion with an embossed image of a bearded man facing left in profile. To the left of the man is the text "ALFR•" then "NOBEL", and on the right, the text (smaller) "NAT•" then "MDCCCXXXIII" above, followed by (smaller) "OB•" then "MDCCCXCVI" below.
Awarded forDiscoveries in physiology or medicine that led to benefit for humankind
LocationStockholm, Sweden
Presented byNobel Assembly at Karolinska Institutet
Reward(s)9 million SEK (2017)[1]
First awarded1901
Currently held byJames P. Allison, Tasuku Honjo (2018)
Websitenobelprize.org

Background

AlfredNobel adjusted
Nobel was interested in experimental physiology and set up his own laboratories.

Alfred Nobel was born on 21 October 1833 in Stockholm, Sweden, into a family of engineers.[3] He was a chemist, engineer and inventor who amassed a fortune during his lifetime, most of it from his 355 inventions of which dynamite is the most famous.[4] He was interested in experimental physiology and set up his own labs in France and Italy to conduct experiments in blood transfusions. Keeping abreast of scientific findings, he was generous in his donations to Ivan Pavlov's laboratory in Russia, and was optimistic about the progress resulting from scientific discoveries made in laboratories.[5]

In 1888, Nobel was surprised to read his own obituary, titled "The merchant of death is dead", in a French newspaper. As it happened, it was Nobel's brother Ludvig who had died, but Nobel, unhappy with the content of the obituary and concerned that his legacy would reflect poorly on him, was inspired to change his will.[6] In his last will, Nobel requested that his money be used to create a series of prizes for those who confer the "greatest benefit on mankind" in physics, chemistry, peace, physiology or medicine, and literature.[7] Though Nobel wrote several wills during his lifetime, the last was written a little over a year before he died at the age of 63.[8] Because his will was contested, it was not approved by the Storting (Norwegian Parliament) until 26 April 1897.[9]

After Nobel's death, the Nobel Foundation was set up to manage the assets of the bequest.[10] In 1900, the Nobel Foundation's newly created statutes were promulgated by Swedish King Oscar II.[11][12] According to Nobel's will, the Karolinska Institute in Sweden, a medical school and research center, is responsible for the Prize in Physiology or Medicine.[13] Today, the prize is commonly referred to as the Nobel Prize in Medicine.[14]

Nomination and selection

It was important to Nobel that the prize be awarded for a "discovery" and that it be of "greatest benefit on mankind".[15] Per the provisions of the will, only select persons are eligible to nominate individuals for the award. These include members of academies around the world, professors of medicine in Sweden, Denmark, Norway, Iceland, and Finland, as well as professors of selected universities and research institutions in other countries. Past Nobel laureates may also nominate.[16] Until 1977, all professors of Karolinska Institute together decided on the Nobel Prize in Physiology or Medicine. That year, changes in Swedish law forced the Institute to make public any documents pertaining to the Nobel Prize and it was considered necessary to establish a legally independent body for the Prize work. Therefore, the Nobel Assembly was constituted, consisting of 50 professors at Karolinska Institute. It elects the Nobel Committee with 5 members who evaluate the nominees, the Secretary who is in charge of the organization, and each year 10 adjunct members to assist in the evaluation of candidates. In 1968, a provision was added that no more than three persons may share a Nobel prize.[17]

True to its mandate, the Committee has chosen researchers working in the basic sciences over those who have made applied science contributions. Harvey Cushing, a pioneering American neurosurgeon who identified Cushing's syndrome, was not awarded the prize, nor was Sigmund Freud, as his psychoanalysis lacks hypotheses that can be experimentally confirmed.[18] The public expected Jonas Salk or Albert Sabin to receive the prize for their development of the polio vaccines, but instead the award went to John Enders, Thomas Weller, and Frederick Robbins whose basic discovery that the polio virus could reproduce in monkey cells in laboratory preparations made the vaccines possible.[19]

Through the 1930s, there were frequent prize laureates in classical physiology, but after that the field began fragmenting into specialties. The last classical physiology laureates were John Eccles, Alan Hodgkin, and Andrew Huxley in 1963 for their findings regarding "unitary electrical events in the central and peripheral nervous system."[20]

Prizes

A Medicine or Physiology Nobel Prize laureate earns a gold medal, a diploma bearing a citation, and a sum of money.[21] These are awarded during the prize ceremony at the Stockholm Concert Hall.

Medals

The Nobel Prize medals, minted by Myntverket[22] in Sweden, are registered trademarks of the Nobel Foundation. Each medal features an image of Alfred Nobel in left profile on the obverse (front) side of the medal. The Nobel Prize medals for Physics, Chemistry, Physiology or Medicine, and Literature have identical obverses, showing the image of Alfred Nobel and the years of his birth and death (1833–1896). Before 1980, the medals were made of 23-karat gold; since then the medals are of 18-karat green gold, plated with 23-karat gold.[23]

The medal awarded by the Karolinska Institute displays an image of "the Genius of Medicine holding an open book in her lap, collecting the water pouring out from a rock in order to quench a sick girl's thirst." The medal is inscribed with words taken from Virgil's Aeneid and reads: Inventas vitam juvat excoluisse per artes, which translates to "inventions enhance life which is beautified through art."[24]

Diplomas

Nobel laureates receive a diploma directly from the King of Sweden. Each diploma is uniquely designed by the prize-awarding institutions for the laureate that receives it. In the case of the Nobel Prize in Physiology or Medicine, that is the Nobel Assembly at Karolinska Institute. Well-known artists and calligraphers from Sweden are commissioned to create it.[25] The diploma contains a picture and text which states the name of the laureate and a citation as to why they received the prize.[25]

Award money

At the awards ceremony, the laureate is given a document indicating the award sum. The amount of the cash award may differ from year to year, based on the funding available from the Nobel Foundation. For example, in 2009 the total cash awarded was 10 million SEK (US$1.4 million),[26] but in 2012, the amount was 8 million Swedish Krona, or US$1.1 million.[27] If there are two laureates in a particular category, the award grant is divided equally between the recipients, but if there are three, the awarding committee may opt to divide the grant equally, or award half to one recipient and a quarter to each of the two others.[28][29][30][31]

Ceremony and banquet

The awards are bestowed at a gala ceremony followed by a banquet.[32] The Nobel Banquet is an extravagant affair with the menu, planned months ahead of time, kept secret until the day of the event. The Nobel Foundation chooses the menu after tasting and testing selections submitted by selected chefs of international repute. Currently it is a three-course dinner, although it was originally six courses in 1901. Each Nobel Prize laureate may bring up to 16 guests. Sweden's royal family attends, and typically the Prime Minister and other members of the government attend as well as representatives of the Nobel family.[33]

Laureates

Lorenz and Tinbergen1
Nikolaas Tinbergen (left) and Konrad Lorenz (right) were awarded (with Karl von Frisch) for their discoveries concerning animal behavior.[34]

The first Nobel Prize in Physiology or Medicine was awarded in 1901 to the German physiologist Emil Adolf von Behring.[35] Behring's discovery of serum therapy in the development of the diphtheria and tetanus vaccines put "in the hands of the physician a victorious weapon against illness and deaths".[36][37] In 1902, the award went to Ronald Ross for his work on malaria, "by which he has shown how it enters the organism and thereby has laid the foundation for successful research on this disease and methods of combating it".[38] He identified the mosquito as the transmitter of malaria, and worked tirelessly on measures to prevent malaria worldwide.[39][40] The 1903 prize was awarded to Niels Ryberg Finsen, the first Faroese laureate, "in recognition of his contribution to the treatment of diseases, especially lupus vulgaris, with concentrated light radiation, whereby he has opened a new avenue for medical science".[41][42] He died within a year after receiving the prize at the age of 43.[43] Ivan Pavlov, whose work Nobel admired and supported, received the prize in 1904 for his work on the physiology of digestion.[44]

Subsequently, those selecting the recipients have exercised wide latitude in determining what falls under the umbrella of Physiology or Medicine. The awarding of the prize in 1973 to Nikolaas Tinbergen, Konrad Lorenz, and Karl von Frisch for their observations of animal behavioral patterns could be considered a prize in the behavioral sciences rather than medicine or physiology.[14] Tinbergen expressed surprise in his Nobel Prize acceptance speech at "the unconventional decision of the Nobel Foundation to award this year's prize 'for Physiology or Medicine' to three men who had until recently been regarded as 'mere animal watchers'".[45]

Gerty Theresa Cori
In 1947, Gerty Cori was the first woman to be awarded the Prize in Physiology or Medicine.

Laureates have been awarded the Nobel Prize in a wide range of fields that relate to physiology or medicine. As of 2010, eight Prizes have been awarded for contributions in the field of signal transduction through G proteins and second messengers. 13 have been awarded for contributions in the field of neurobiology[46] and 13 have been awarded for contributions in Intermediary metabolism.[47] The 100 Nobel Prizes in Physiology or Medicine have been awarded to 195 individuals through 2009.[48][49] Twelve women have received the prize: Gerty Cori (1947), Rosalyn Yalow (1977), Barbara McClintock (1983), Rita Levi-Montalcini (1986), Gertrude B. Elion (1988), Christiane Nüsslein-Volhard (1995), Linda B. Buck (2004), Françoise Barré-Sinoussi (2008), Elizabeth H. Blackburn (2009), Carol W. Greider (2009), May-Britt Moser (2014) and Youyou Tu (2015).[50] Only one woman, Barbara McClintock, has received an unshared prize in this category, for the discovery of genetic transposition.[48][51] Mario Capecchi, Martin Evans, and Oliver Smithies was awarded the prize in 2007 for the discovery of a gene targeting procedure (a type of genetic recombination) for introducing homologous recombination in mice, employing embryonic stem cells through the development of the knockout mouse.[52][53] There have been 37 times when the Nobel Prize in Physiology or Medicine was awarded to a single individual, 31 times when it was shared by two, and 33 times there were three laureates (the maximum allowed).

In 2009, the Nobel Prize was awarded to Elizabeth Blackburn, Carol W. Greider and Jack W. Szostak of the United States for discovering the process by which chromosomes are protected by telomeres (regions of repetitive DNA at the ends of chromosomes) and the enzyme telomerase; they shared the prize of 10,000,000 SEK (slightly more than 1 million, or US$1.4 million).[54] Rita Levi-Montalcini, an Italian neurologist, who together with colleague Stanley Cohen, received the 1986 Nobel Prize in Physiology or Medicine for their discovery of Nerve growth factor (NGF), was the first Nobel laureate to reach the 100th birthday.[49]

Time factor and death

RMSt
Ralph M. Steinman was the first posthumous recipient of the Prize in Physiology or Medicine.

Because of the length of time that may pass before the significance of a discovery becomes apparent, some prizes are awarded many years after the initial discovery. Barbara McClintock made her discoveries in 1944, before the structure of the DNA molecule was known; she was not awarded the prize until 1983. Similarly, in 1916 Peyton Rous discovered the role of tumor viruses in chickens, but was not awarded the prize until 50 years later, in 1966.[55] Nobel laureate Carol Greider's research leading to the prize was conducted over 20 years before. She noted that the passage of time is an advantage in the medical sciences, as it may take many years for the significance of a discovery to become apparent.[56]

In 2011, Canadian immunologist Ralph M. Steinman was awarded the prize; however, unknown to the committee, he had died three days before the announcement. The committee decided that since the prize was awarded "in good faith," it would be allowed to stand.

Controversial inclusions and exclusions

Some of the awards have been controversial. The person who was deserving of the 1923 prize for the discovery of insulin as a central hormone for controlling diabetes (awarded only a year after its discovery)[57] has been heatedly debated. It was shared between Frederick Banting and John Macleod; this infuriated Banting who regarded Macleod's involvement as minimal. Macleod was the department head at the University of Toronto but otherwise was not directly involved in the findings. Banting thought his laboratory partner Charles Best, who had shared in the laboratory work of discovery, should have shared the prize with him as well. In fairness, he decided to give half of his prize money to Best. Macleod on his part felt the biochemist James Collip, who joined the laboratory team later, deserved to be included in the award and shared his prize money with him.[57] Some maintain that Nicolae Paulescu, a Romanian professor of physiology at the University of Medicine and Pharmacy in Bucharest, was the first to isolate insulin, in 1916, although his pancrein was an impure aqueous extract unfit for human treatment similar to the one used previously by Israel Kleiner.[58][59][60] When Banting published the paper that brought him the Nobel,[61] Paulescu already held a patent for his discovery (10 April 1922, patent no. 6254 (8322) "Pancreina şi procedeul fabricaţiei ei"/"Pancrein and the process of making it", from the Romanian Ministry of Industry and Trade).[62][63][64]

The Spanish neurophysiologist Fernando de Castro (1896-1967) was the first to describe arterial chemoreceptors and circumscribe them to the carotid body for the respiratory reflexes in 1926-1928. For many experts, this direct disciple of Santiago Ramón y Cajal deserved to share the Nobel Prize 1938 with the awarded Corneille Heymans, but at that time Spain was immersed in the Spanish Civil War and it seems that the Nobel Board even doubted if he was alive or not, being Madrid at the front since almost the beginning of the conflict. Heymans himself recognized the merits of De Castro for the Nobel Prize in different occasions, including a famous talk in Montevideo (Uruguay).[65]

Montagnier-Barré-Sinoussi-zur Hausen-press conference Dec 06th, 2008-1
Scandal and controversy resulted from the 2008 award to Harald zur Hausen for the discovery of HPV, and to Françoise Barré-Sinoussi and Luc Montagnier for discovering HIV.

In 1949, despite protests from the medical establishment, the Portuguese neurologist António Egas Moniz received the Physiology or Medicine Prize for his development of the prefrontal leucotomy, which he promoted by declaring the procedure's success just 10 days postoperative. Due largely to the publicity surrounding the award, it was prescribed without regard for modern medical ethics. Favorable results were reported by such publications as The New York Times. It is estimated that around 40,000 lobotomies were performed in the United States before the procedure's popularity faded.[66] Rosemary Kennedy, the sister of John F. Kennedy, was subjected to the procedure by their father; it incapacitated her to the extent that she needed to be institutionalized for the rest of her life.[67][68]

The 1952 prize, awarded solely to Selman Waksman for his discovery of streptomycin, omitted the recognition some felt due to his co-discoverer Albert Schatz.[69][70] There was litigation brought by Schatz against Waksman over the details and credit of the streptomycin discovery; Schatz was awarded a substantial settlement, and, together with Waksman, Schatz was to be officially recognized as a co-discoverer of streptomycin as concerned patent rights. However, he is not recognized as a Nobel Prize laureate.[69]

The 1962 Prize awarded to James D. Watson, Francis Crick, and Maurice Wilkins—for their work on DNA structure and properties—did not recognize contributing work from others, such as Alec Stokes and Herbert Wilson. In addition, Erwin Chargaff, Oswald Avery, and Rosalind Franklin (whose key DNA x-ray crystallography work was the most detailed yet least acknowledged among the three)[71] contributed directly to the ability of Watson and Crick to solve the structure of the DNA molecule. But Avery died in 1955, Franklin died in 1958 and posthumous nominations for the Nobel Prize are not permitted. However, recently unsealed files of the Nobel Prize nominations reveal that no one ever nominated Franklin for the prize when she was alive.[72] Wilkins' crucial contribution was to show Rosalind Franklin's key x-ray photos to Watson.[73] As a result of Watson's misrepresentations of Franklin and her role in the discovery of the double helix in his book The Double Helix, Franklin has come to be portrayed as a classic victim of sexism in science.[74][75] Chargaff, for his part, was not quiet about his exclusion from the prize, bitterly writing to other scientists about his disillusionment regarding the field of molecular biology.[73]

The 2008 award went to Harald zur Hausen in recognition of his discovery that human papillomavirus (HPV) can cause cervical cancer, and to Françoise Barré-Sinoussi and Luc Montagnier for discovering the human immunodeficiency virus (HIV).[76] Whether Robert Gallo or Luc Montagnier deserved more credit for the discovery of the virus that causes AIDS has been a matter of considerable controversy. As it was, Gallo was left out and not awarded a prize.[77][78] Additionally, there was scandal when it was learned that Harald zur Hausen was being investigated for having a financial interest in vaccines for the cervical cancer that HPV can cause. AstraZeneca, which with a stake in two lucrative HPV vaccines could benefit financially from the prize, had agreed to sponsor Nobel Media and Nobel Web. According to Times Online, two senior figures in the selection process that chose zur Hausen also had strong links with AstraZeneca.[79]

Limits on number of awardees

The provision that restricts the maximum number of nominees to three for any one prize, introduced in 1968, has caused considerable controversy.[17][80] From the 1950s onward, there has been an increasing trend to award the Nobel Prize in Physiology or Medicine to more than one person. There were 59 people who received the prize in the first 50 years of the last century, while 113 individuals received it between 1951 and 2000. This increase could be attributed to the rise of the international scientific community after World War II, resulting in more persons being responsible for the discovery, and nominated for, a particular prize. Also, current biomedical research is more often carried out by teams rather than by scientists working alone, making it unlikely that any one scientist, or even a few, is primarily responsible for a discovery;[19] this has meant that a prize nomination that would have to include more than three contributors is automatically excluded from consideration.[55] Also, deserving contributors may not be nominated at all because the restriction results in a cut off point of three nominees per prize, leading to controversial exclusions.[15]

Years without awards

There have been nine years in which the Nobel Prize in Physiology or Medicine was not awarded (1915–1918, 1921, 1925, 1940–1942). Most of these occurred during either World War I (1914–1918) or World War II (1939–1945).[49] In 1939, Adolf Hitler's Third Reich forbade Gerhard Domagk to accept his prize.[81] He was later able to receive the diploma and medal but not the money.[49][82]

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Sources

Books

Further reading

External links

Alfred P. Sloan Jr. Prize

The Alfred P. Sloan Jr. Prize was a $250,000

award given by the General Motors Cancer Research Foundation for outstanding oncological research.The prize was awarded annually from 1979 to 2005. Of the winners, 15 out of 37 have gone on to win either a Nobel Prize in Physiology or Medicine or a Nobel Prize in Chemistry.

in 2006, due to budget constraints, the Alfred P. Sloan Jr. prize, the Charles K. Kettering prize, and the Charles S. Mott Prize were consolidated into a single General Motors Cancer Research Award which also had a value of $250,000.

The first and only winner of the General Motors Cancer Research Award was Napoleone Ferrara.After 2006 no more prizes were awarded.

Charité

The Charité – Universitätsmedizin Berlin is Europe's largest university clinic, affiliated with Humboldt University and Freie Universität Berlin. With numerous Collaborative Research Centers (CRC) of the Deutsche Forschungsgemeinschaft, including three Nobel Prize laureates working as research fellows in the Einstein Foundation, it is one of Germany's most research-intensive medical institutions. From 2012 to 2017, it was ranked by Focus as the best of over 1000 hospitals in Germany. More than half of all German Nobel Prize winners in Physiology or Medicine, including Emil von Behring, Robert Koch and Paul Ehrlich, worked at the Charité. Its medical school is one of the most prestigious and competitive in Germany with admission rates lower than 5%.

G protein

G proteins, also known as guanine nucleotide-binding proteins, are a family of proteins that act as molecular switches inside cells, and are involved in transmitting signals from a variety of stimuli outside a cell to its interior. Their activity is regulated by factors that control their ability to bind to and hydrolyze guanosine triphosphate (GTP) to guanosine diphosphate (GDP). When they are bound to GTP, they are 'on', and, when they are bound to GDP, they are 'off'. G proteins belong to the larger group of enzymes called GTPases.

There are two classes of G proteins. The first function as monomeric small GTPases (small G-proteins), while the second function as heterotrimeric G protein complexes. The latter class of complexes is made up of alpha (α), beta (β) and gamma (γ) subunits. In addition, the beta and gamma subunits can form a stable dimeric complex referred to as the beta-gamma complex

.

G proteins located within the cell are activated by G protein-coupled receptors (GPCRs) that span the cell membrane. Signaling molecules bind to a domain of the GPCR located outside the cell, and an intracellular GPCR domain then in turn activates a particular G protein. Some inactive-state GPCRs have also been shown to be "pre-coupled" with G proteins. The G protein activates a cascade of further signaling events that finally results in a change in cell function. G protein-coupled receptor and G proteins working together transmit signals from many hormones, neurotransmitters, and other signaling factors. G proteins regulate metabolic enzymes, ion channels, transporter proteins, and other parts of the cell machinery, controlling transcription, motility, contractility, and secretion, which in turn regulate diverse systemic functions such as embryonic development, learning and memory, and homeostasis.

Johns Hopkins School of Medicine

The Johns Hopkins University School of Medicine (JHUSOM), located in Baltimore, Maryland, United States (founded in 1893) is the academic medical teaching and research arm of the Johns Hopkins University, founded in 1876. The School of Medicine shares a campus with the Johns Hopkins Hospital, established in 1889. Johns Hopkins has consistently ranked among the top medical schools in the United States, in the number of research grants awarded by the National Institutes of Health. Its main teaching hospital, the Johns Hopkins Hospital, was ranked the number one hospital in the United States for 22 years by U.S. News & World Report.

List of Jewish Nobel laureates

As of 2017, Nobel Prizes have been awarded to 902 individuals, of whom 203 or 22.5% were Jews, although the total Jewish population comprises less than 0.2% of the world's population. This means the percentage of Jewish Nobel laureates is at least 112.5 times or 11,250% above average. Various theories have been made to explain this phenomenon, which has received considerable attention. Prominent late Israeli academics Dr. Elay Ben-Gal and Professor Yeshayahu Leibowitz, curious about the phenomenon, started to form an encyclopedia of Jewish Nobel laureates and interview as many as possible about their life and work.

List of Nobel laureates affiliated with Imperial College London

The Nobel Prizes are awarded annually by the Royal Swedish Academy of Sciences, the Karolinska Institute, and the Norwegian Nobel Committee to individuals who make outstanding contributions in the fields of chemistry, physics, literature, peace, and physiology or medicine. They were established by the 1895 will of Alfred Nobel, which dictates that the awards should be administered by the Nobel Foundation. Another prize, the Nobel Memorial Prize in Economic Sciences, was established in 1968 by the Sveriges Riksbank, the central bank of Sweden, for contributors to the field of economics. Each prize is awarded by a separate committee: the Royal Swedish Academy of Sciences awards the Prizes in Physics, Chemistry, and Economics, the Karolinska Institute awards the Prize in Physiology or Medicine, and the Norwegian Nobel Committee awards the Prize in Peace. Each recipient receives a medal, a diploma and a cash prize that has varied throughout the years. In 1901, the winners of the first Nobel Prizes were given 150,782 SEK, which is equal to 7,731,004 SEK in December 2007. In 2008, the winners were awarded a prize amount of 10,000,000 SEK. The awards are presented in Stockholm in an annual ceremony on December 10, the anniversary of Nobel's death.As of 2009, there have been 15 Nobel laureates affiliated with Imperial College London. Imperial College considers laureates who attended the university as undergraduate students, graduate students or were members of the faculty as affiliated laureates. Frederick Gowland Hopkins, who attended the Royal School of Mines from 1881 to 1883, was the first Imperial College-affiliated laureate, winning the Nobel Prize in Physiology or Medicine in 1929. One Nobel Prize was shared by two Imperial College laureates; Ernst Boris Chain and Alexander Fleming won the 1945 Nobel Prize in Physiology or Medicine. Thirteen of the fifteen Imperial College laureates were members of the faculty. Nine laureates were fellows of the college. No Imperial College laureate has won the Nobel Prize in Literature, the Nobel Peace Prize, or the Nobel Memorial Prize in Economic Sciences.

List of Nobel laureates affiliated with Johns Hopkins University

The Nobel Prizes are awarded annually by the Royal Swedish Academy of Sciences, the Karolinska Institute, and the Norwegian Nobel Committee to individuals who make outstanding contributions in the fields of chemistry, physics, literature, peace, and physiology or medicine. They were established by the 1895 will of Alfred Nobel, which dictates that the awards should be administered by the Nobel Foundation. Another prize, the Nobel Memorial Prize in Economic Sciences, was established in 1968 by the Sveriges Riksbank, the central bank of Sweden, for contributors to the field of economics. Each prize is awarded by a separate committee: the Royal Swedish Academy of Sciences awards the Prizes in Physics, Chemistry, and Economics, the Karolinska Institute awards the Prize in Physiology or Medicine, and the Norwegian Nobel Committee awards the Prize in Peace. Each recipient receives a medal, a diploma and a cash prize that has varied throughout the years. In 1901, the winners of the first Nobel Prizes were given 150,782 SEK, which is equal to 7,731,004 SEK in December 2007. In 2008, the winners were awarded a prize amount of 10,000,000 SEK. The awards are presented in Stockholm in an annual ceremony on December 10, the anniversary of Nobel's death.As of 2011, there have been 36 Nobel laureates affiliated with Johns Hopkins University. Johns Hopkins considers laureates who attended the university as undergraduate students, graduate students or were members of the faculty as affiliated laureates. Woodrow Wilson, who received his Ph.D. from Johns Hopkins in 1886, was the first Johns Hopkins-affiliated laureate, winning the Nobel Peace Prize in 1919. Four Nobel Prizes were shared by Johns Hopkins laureates: George Minot and George Whipple won the 1934 Nobel Prize in Physiology or Medicine, Joseph Erlanger and Herbert Spencer Gasser won the 1944 Nobel Prize in Physiology in Medicine, Daniel Nathans and Hamilton O. Smith won the 1978 Nobel Prize in Physiology or Medicine, and David H. Hubel and Torsten N. Wiesel won the 1981 Nobel Prize in Physiology or Medicine. Eighteen Johns Hopkins laureates have won the Nobel Prize in Physiology or Medicine, more than any other category. Twenty-four laureates were members of the Johns Hopkins faculty, five laureates received their Ph.D. at Johns Hopkins, eight laureates received their M.D. at Johns Hopkins, and two laureates, Francis Peyton Rous and Martin Rodbell, received their undergraduate degrees at Johns Hopkins.

List of Nobel laureates affiliated with Washington University in St. Louis

The Nobel Prizes are awarded annually by the Royal Swedish Academy of Sciences, the Karolinska Institute, and the Norwegian Nobel Committee to individuals who make outstanding contributions in the fields of chemistry, physics, literature, peace, and physiology or medicine. They were established by the 1895 will of Alfred Nobel, which dictates that the awards should be administered by the Nobel Foundation. Another prize, the Nobel Memorial Prize in Economic Sciences, was established in 1968 by the Sveriges Riksbank, the central bank of Sweden, for contributors to the field of economics. Each prize is awarded by a separate committee; the Royal Swedish Academy of Sciences awards the Prizes in Physics, Chemistry, and Economics, the Karolinska Institute awards the Prize in Physiology or Medicine, and the Norwegian Nobel Committee awards the Prize in Peace. Each recipient receives a medal, a diploma and a cash prize that has varied throughout the years. In 1901, the winners of the first Nobel Prizes were given 150,782 SEK, which is equal to 7,731,004 SEK in December 2007. In 2008, the winners were awarded a prize amount of 10,000,000 SEK. The awards are presented in Stockholm in an annual ceremony on December 10, the anniversary of Nobel's death.As of 2014, there have been 23 laureates affiliated with Washington University in St. Louis. Washington University considers laureates who attended the university as undergraduate students, graduate students or were members of the faculty as affiliated laureates. Arthur Compton, the chancellor of the university from 1945 to 1953, was the first laureate affiliated with the university, winning the Nobel Prize in Physics in 1927. Four Nobel Prizes were shared by Washington University laureates; Joseph Erlanger and Herbert Spencer Gasser won the 1944 Nobel Prize in Physiology or Medicine, Carl Ferdinand Cori and wife Gerty Cori won the 1947 Nobel Prize in Physiology or Medicine, Arthur Kornberg and Severo Ochoa won the 1959 Nobel Prize in Physiology or Medicine, and Daniel Nathans and George Davis Snell won the 1980 Nobel Prize in Physiology or Medicine. Seventeen Washington University laureates have won the Nobel Prize in Physiology or Medicine, more than any other category. With the exception of Daniel Nathans, who received his M.D. from Washington University and William E. Moerner who received his undergraduate degrees from the university, all Washington University laureates have been members of the university faculty.

List of Nobel laureates affiliated with the University of Edinburgh

This list of Nobel laureates affiliated with the University of Edinburgh includes academic staff and researchers as well as graduates and non-graduate former students of the University of Edinburgh in Scotland, who were bestowed with the Nobel Prize and the Nobel Memorial Prize in Economic Sciences.

As of 2017, 23 Nobel laureates have been affiliated with the University. They have been affiliated as students, researchers, administrators and professors, and they have won Nobel Prizes in all categories.

It does not include those whose only affiliation with the university is (i) giving the Gifford Lectures (e.g. Niels Bohr, who gave the lecture entitled Causality and Complementarity: Epistemological Lessons of Studies in Atomic Physics in 1949) or (ii) the conferral of an honorary degree (e.g. August Krogh).

List of Nobel laureates in Physiology or Medicine

The Nobel Prize in Physiology or Medicine (Swedish: Nobelpriset i fysiologi eller medicin) is awarded annually by the Swedish Karolinska Institute to scientists and doctors in the various fields of physiology or medicine. It is one of the five Nobel Prizes established by the 1895 will of Alfred Nobel (who died in 1896), awarded for outstanding contributions in chemistry, physics, literature, peace, and physiology or medicine. As dictated by Nobel's will, the award is administered by the Nobel Foundation and awarded by a committee that consists of five members and an executive secretary elected by the Karolinska Institute. While commonly referred to as the Nobel Prize in Medicine, Nobel specifically stated that the prize be awarded for "physiology or medicine" in his will. Because of this, the prize can be awarded in a broader range of fields. The first Nobel Prize in Physiology or Medicine was awarded in 1901 to Emil Adolf von Behring, of Germany. Each recipient receives a medal, a diploma and a monetary award that has varied throughout the years. In 1901, von Behring received 150,782 SEK, which was equal to 7,731,004 SEK in December 2008. The award is presented in Stockholm at an annual ceremony on 10 December, the anniversary of Nobel's death.Laureates have won the Nobel Prize in a wide range of fields that relate to physiology or medicine. As of 2009, 8 Prizes have been awarded for contributions in the field of signal transduction by G proteins and second messengers, 13 have been awarded for contributions in the field of neurobiology and 13 have been awarded for contributions in intermediary metabolism. In 1939 Gerhard Domagk, a German, was not allowed by his government to accept the prize. He later received a medal and diploma, but not the money. As of 2018, the prize has been awarded to 216 individuals, twelve of them were women: Gerty Cori (1947), Rosalyn Yalow (1977), Barbara McClintock (1983), Rita Levi-Montalcini (1986), Gertrude B. Elion (1988), Christiane Nüsslein-Volhard (1995), Linda B. Buck (2004), Françoise Barré-Sinoussi (2008), Elizabeth H. Blackburn (2009), Carol W. Greider (2009), May-Britt Moser (2014) and Tu Youyou (2015). There have been nine years in which the Nobel Prize in Physiology or Medicine was not awarded (1915–1918, 1921, 1925, 1940–1942).

List of RNA biologists

For related information, see the articles on History of RNA Biology, History of Molecular Biology, and History of Genetics.

List of biologists

This is a list of notable biologists with a biography in Wikipedia. It includes zoologists, botanists, ornithologists, malacologists, naturalists and other specialities.

List of female Nobel laureates

As of 2018, Nobel Prizes have been awarded to 853 men, 51 women (Marie Curie won it twice), and 24 unique organizations.The distribution of female Nobel Laureates is as follows:

seventeen women have won the Nobel Peace Prize,

fourteen have won the Nobel Prize in Literature,

twelve have won the Nobel Prize in Physiology or Medicine,

five have won the Nobel Prize in Chemistry,

three have won the Nobel Prize in Physics,

and one, Elinor Ostrom, has won the Nobel Memorial Prize in Economic Sciences.The first woman to win a Nobel Prize was Marie Curie, who won the Nobel Prize in Physics in 1903 with her husband, Pierre Curie, and Henri Becquerel. Curie is also the only woman to have won multiple Nobel Prizes; in 1911, she won the Nobel Prize in Chemistry. Curie's daughter, Irène Joliot-Curie, won the Nobel Prize in Chemistry in 1935, making the two the only mother-daughter pair to have won Nobel Prizes.The most Nobel Prizes awarded to women in a single year was in 2009, when five women became laureates in four categories.

The most recent women to be awarded a Nobel Prize were Donna Strickland in Physics, Frances Arnold in Chemistry, and Nadia Murad for Peace (2018).

List of neuroscientists

Many famous neuroscientists are from the 20th and 21st century, as neuroscience is a fairly new science. However many anatomists, physiologists, biologists, neurologists, psychiatrists and other physicians and psychologists are considered to be neuroscientists as well. This list compiles the names of all neuroscientists with a corresponding Wikipedia biographical article, and is not necessarily a reflection of their relative importance in the field.

Neuroscientist

A neuroscientist (or neurobiologist) is a scientist who has specialised knowledge in the field of neuroscience, the branch of biology that deals with the physiology, biochemistry, anatomy and molecular biology of neurons and neural circuits and especially their association with behaviour and learning.

Neuroscientists generally work as researchers within a college, university, government agency, or private industry setting. In research-oriented careers, neuroscientists typically spend their time designing and carrying out scientific experiments that contribute to the understanding of the nervous system and its function. They can engage in basic or applied research. Basic research seeks to add information to our current understanding of the nervous system, whereas applied research seeks to address a specific problem, such as developing a treatment for a neurological disorder. Biomedically-oriented neuroscientists typically engage in applied research. Neuroscientists also have a number of career opportunities outside the realm of research, including careers in industry, science writing, government program management, science advocacy, and education. These individuals most commonly hold doctorate degrees in the sciences, but may also hold a master's degree.

The Neuroscientists day is celebrated on August 13th.

Nobel Assembly at the Karolinska Institute

The Nobel Assembly at the Karolinska Institute is a body at Karolinska Institute which awards the Nobel Prize in Physiology or Medicine. It is headquartered in the Nobel Forum on the grounds of the Karolinska Institute campus. Originally the Nobel Assembly was not a formal body but rather the collective of all professors (holders of chairs, i.e. full professors) at Karolinska Institute. In 1977 the Nobel Assembly became a separate private organization hosted by Karolinska Institute. Until 1984 all Karolinska Institute professors belonged to the Assembly; from 1984 the membership is restricted to 50 Karolinska Institute professors.The main work involved in collecting nominations and screening nominees is performed by the Nobel Committee at the Karolinska Institute, which has five members. The Nobel Committee, which is appointed by the Nobel Assembly, is only empowered to recommend laureates, while the final decision rests with the Nobel Assembly.

Passano Foundation

The Passano Foundation, established in 1945, provides an annual award to a research scientist whose work – done in the United States – is thought to have immediate practical benefits. Many Passano laureates have subsequently won the Nobel Prize.

Robert W. Holley

Robert William Holley (January 28, 1922 – February 11, 1993) was an American biochemist. He shared the Nobel Prize in Physiology or Medicine in 1968 (with Har Gobind Khorana and Marshall Warren Nirenberg) for describing the structure of alanine transfer RNA, linking DNA and protein synthesis.

Holley was born in Urbana, Illinois, and graduated from Urbana High School in 1938. He went on to study chemistry at the University of Illinois at Urbana-Champaign, graduating in 1942 and commencing his PhD studies in organic chemistry at Cornell University. During World War II Holley spent two years working under Professor Vincent du Vigneaud at Cornell University Medical College, where he was involved in the first chemical synthesis of penicillin. Holley completed his PhD studies in 1947.Following his graduate studies Holley remained associated with Cornell. He became an Assistant Professor of organic chemistry in 1948, and was appointed as Professor of Biochemistry in 1962. He began his research on RNA after spending a year's sabbatical (1955–1956) studying with James F. Bonner at the California Institute of Technology.

Holley's research on RNA focused first on isolating transfer RNA (tRNA), and later on determining the sequence and structure of alanine tRNA, the molecule that incorporates the amino acid alanine into proteins. Holley's team of researchers determined the tRNA's structure by using two ribonucleases to split the tRNA molecule into pieces. Each enzyme split the molecule at location points for specific nucleotides. By a process of "puzzling out" the structure of the pieces split by the two different enzymes, then comparing the pieces from both enzyme splits, the team eventually determined the entire structure of the molecule.

The structure was completed in 1964, and was a key discovery in explaining the synthesis of proteins from messenger RNA. It was also the first nucleotide sequence of a ribonucleic acid ever determined. Holley was awarded the Nobel Prize in Physiology or Medicine in 1968 for this discovery, and Har Gobind Khorana and Marshall W. Nirenberg were also awarded the prize that year for contributions to the understanding of protein synthesis.

Using the Holley team's method, other scientists determined the structures of the remaining tRNA's. A few years later the method was modified to help track the sequence of nucleotides in various bacterial, plant, and human viruses.

In 1968 Holley became a resident fellow at the Salk Institute for Biological Studies in La Jolla, California.

Tadeusz Reichstein

Tadeusz Reichstein (20 July 1897 – 1 August 1996) was a Polish-Swiss chemist and the Nobel Prize in Physiology or Medicine laureate (1950).

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