Nobel Prize in Physiology or Medicine

for the discovery of microRNA and its role in post-transcriptional gene regulation

Victor Ambros

United States of America

Gary Ruvkun

United States of America

for their discoveries concerning nucleoside base modifications that enabled the development of effective mRNA vaccines against COVID-19

Katalin Karikó

Hungary, United States of America

Drew Weissman

United States of America

for his discoveries concerning the genomes of extinct hominins and human evolution

Svante Pääbo

Sweden

for the discovery of receptors for temperature and touch

David Julius

United States of America

Ardem Patapoutian

United States of America

for the discovery of the Hepatitis C virus

Harvey J. Alter

United States of America

Michael Houghton

United Kingdom of Great Britain and Northern Ireland

Charles M. Rice

United States of America

for their discoveries of how cells sense and adapt to oxygen availability

William Kaelin Jr.

United States of America

Peter J. Ratcliffe

United Kingdom of Great Britain and Northern Ireland

Gregg L. Semenza

United States of America

for the discovery of immune-checkpoint inhibitors and their application to cancer therapy

James P. Allison

United States of America

Tasuku Honjo

Japan

for the discovery of molecular mechanisms controlling the circadian rhythm

Jeffrey C. Hall

United States of America

Michael Rosbash

United States of America

Michael W. Young

United States of America

for the discoveries of mechanisms for autophagy

Yoshinori Ohsumi

Japan

for their discoveries concerning a novel therapy against infections caused by roundworm parasites

William C. Campbell

Ireland, United States of America

Satoshi Ōmura

Japan

for the discovery of cells that constitute a positioning system in the brain

John O'Keefe

United States of America, United Kingdom of Great Britain and Northern Ireland

May-Britt Moser

Norway

Edvard I. Moser

Norway

for the discoveries of the machinery that regulates vesicle traffic, the system that transports proteins to their correct destinations inside cells

Randy Schekman

United States of America

James Rothman

United States of America

Thomas C. Südhof

United States of America

for the discovery that mature cells can be reprogrammed to become pluripotent

John Gurdon

United Kingdom of Great Britain and Northern Ireland

Shinya Yamanaka

Japan

for their discoveries concerning the activation of innate immunity

Bruce Beutler

United States of America

Jules A. Hoffmann

France

for the development of in vitro fertilization (IVF) technology

Robert G. Edwards

United Kingdom of Great Britain and Northern Ireland

for the discovery of how chromosomes are protected by telomeres and the enzyme telomerase

Elizabeth H. Blackburn

United States of America, Australia

Carol W. Greider

United States of America

Jack W. Szostak

United States of America

Discovery of human papillomaviruses causing cervical cancer

Harald zur Hausen

Germany

for their discoveries of principles for introducing specific gene modifications in mice by the use of embryonic stem cells

Mario Capecchi

United States of America

Martin Evans

United Kingdom of Great Britain and Northern Ireland

Oliver Smithies

United States of America

for the discovery of RNA interference – gene silencing by double-stranded RNA

Andrew Fire

United States of America

Craig Mello

United States of America

for the discovery of the bacterium Helicobacter pylori and its role in gastritis and peptic ulcer disease

Barry Marshall

Australia

Robin Warren

Australia

for their discoveries of odorant receptors and the organization of the olfactory system

Richard Axel

United States of America

Linda B. Buck

United States of America

for their discoveries concerning magnetic resonance imaging

Paul Lauterbur

United States of America

Peter Mansfield

United Kingdom of Great Britain and Northern Ireland

for the discoveries concerning genetic regulation of organ development and programmed cell death

Sydney Brenner

United Kingdom of Great Britain and Northern Ireland

H. Robert Horvitz

United States of America

John E. Sulston

United Kingdom of Great Britain and Northern Ireland

for their discoveries of key regulators of the cell cycle

Leland H. Hartwell

United States of America

Tim Hunt

United Kingdom of Great Britain and Northern Ireland

Paul M. Nurse

United Kingdom of Great Britain and Northern Ireland

for their discoveries concerning signal transduction in the nervous system

Arvid Carlsson

Sweden

Paul Greengard

United States of America

Eric R. Kandel

United States of America

for the discovery that proteins have intrinsic signals that govern their transport and localization in the cell

Günter Blobel

United States of America

for the discovery of nitric oxide as a signalling molecule in the cardiovascular system

Robert F. Furchgott

United States of America

Louis J. Ignarro

United States of America

Ferid Murad

United States of America

for the discovery of Prions – a new biological principle of infection

Stanley B. Prusiner

United States of America

for their discoveries concerning the specificity of the cell-mediated immune defence

Peter C. Doherty

Australia

Rolf M. Zinkernagel

Switzerland

for their discoveries concerning the genetic control of early embryonic development

Edward B. Lewis

United States of America

Christiane Nüsslein-Volhard

Germany

Eric F. Wieschaus

United States of America

for their discovery of G-proteins and the role of these proteins in signal transduction in cells

Alfred G. Gilman

United States of America

Martin Rodbell

United States of America

for the discovery of split genes

Richard J. Roberts

United Kingdom of Great Britain and Northern Ireland

Phillip Allen Sharp

United States of America

for the discoveries concerning reversible protein phosphorylation as a biological regulatory mechanism

Edmond H. Fischer

Switzerland, United States of America

Edwin G. Krebs

United States of America

for their discoveries concerning the function of single ion channels in cells

Erwin Neher

Germany

Bert Sakmann

Germany

for their discoveries concerning organ and cell transplantation in the treatment of human disease

Joseph E. Murray

United States of America

E. Donnall Thomas

United States of America

for the discovery that the oncogenes of retroviruses are of cellular origin

J. Michael Bishop

United States of America

Harold E. Varmus

United States of America

for their discoveries of important principles for drug treatment

James W. Black

United Kingdom of Great Britain and Northern Ireland

Gertrude B. Elion

United States of America

George H. Hitchings

United States of America

for his discovery of the genetic principle for generation of antibody diversity

Susumu Tonegawa

Japan

for their discoveries of growth factors

Rita Levi-Montalcini

Italy, United States of America

Stanley Cohen

United States of America

for their discoveries concerning the regulation of cholesterol metabolism

Michael Stuart Brown

United States of America

Joseph L. Goldstein

United States of America

for theories concerning the specificity in development and control of the immune system and the discovery of the principle for production of monoclonal antibodies

Niels Kaj Jerne

Denmark

Georges J. F. Köhler

Germany

César Milstein

Argentina, United Kingdom of Great Britain and Northern Ireland

for the discovery of mobile genetic elements

Barbara McClintock

United States of America

discoveries concerning prostaglandins and related biologically active substances

Sune Bergström

Sweden

Bengt I. Samuelsson

Sweden

John Robert Vane

United Kingdom of Great Britain and Northern Ireland

for discoveries concerning the functional specialization of the cerebral hemispheres

Roger Wolcott Sperry

United States of America

for their discoveries concerning genetically determined structures on the cell surface that regulate immunological reactions

Baruj Benacerraf

United States of America

Jean Dausset

France

George Davis Snell

United States of America

for the development of computer assisted tomography (CT)

Godfrey Hounsfield

United Kingdom of Great Britain and Northern Ireland

Allan McLeod Cormack

United States of America

for the discovery of restriction enzymes and their application to problems of molecular genetics

Daniel Nathans

United States of America

Hamilton O. Smith

United States of America

Werner Arber

Switzerland

for their discoveries concerning the peptide hormone production of the brain

Roger Guillemin

United States of America

Andrew Schally

United States of America

for their discoveries concerning new mechanisms for the origin and dissemination of infectious diseases

Baruch Samuel Blumberg

United States of America

Daniel Carleton Gajdusek

United States of America

for their discoveries concerning the interaction between tumour viruses and the genetic material of the cell

Renato Dulbecco

United States of America

Howard Martin Temin

United States of America

David Baltimore

United States of America

for their discoveries concerning the structural and functional organization of the cell

Albert Claude

Belgium

Christian de Duve

Belgium

George Emil Palade

United States of America

for their discoveries concerning organization and elicitation of individual and social behaviour patterns

Konrad Lorenz

Austria

Karl von Frisch

Germany

Nikolaas Tinbergen

United Kingdom of Great Britain and Northern Ireland

for the discoveries concerning the chemical structure of antibodies

Gerald Edelman

United States of America

Rodney Robert Porter

United Kingdom of Great Britain and Northern Ireland

for his discoveries concerning the mechanisms of the action of hormones

Earl Wilbur Sutherland Jr.

United States of America

for their discoveries concerning the humoral transmitters in the nerve terminals and the mechanisms for their storage, release and inactivation

Bernard Katz

United Kingdom of Great Britain and Northern Ireland

Ulf von Euler

Sweden

Julius Axelrod

United States of America

for their discoveries concerning the replication mechanism and the genetic structure of viruses

Max Delbrück

United States of America

Alfred Hershey

United States of America

Salvador Luria

United States of America

for their interpretation of the genetic code and its function in protein synthesis

Robert W. Holley

United States of America

Har Gobind Khorana

United States of America

Marshall Warren Nirenberg

United States of America

for their discoveries concerning the primary physiological and chemical visual processes in the eye

Ragnar Granit

Sweden

Haldan Keffer Hartline

United States of America

George Wald

United States of America

discovery of tumour-inducing viruses

for their discoveries concerning genetic control of enzyme and virus synthesis

François Jacob

France

André Michel Lwoff

France

Jacques Monod

France

for their discoveries concerning the mechanism and regulation of cholesterol and fatty acid metabolism

Konrad Emil Bloch

United States of America

Feodor Felix Konrad Lynen

West Germany

for their discoveries concerning the ionic mechanisms involved in excitation and inhibition in the peripheral and central portions of the nerve cell membrane

John Carew Eccles

Australia

Alan Lloyd Hodgkin

United Kingdom of Great Britain and Northern Ireland

Andrew Huxley

United Kingdom of Great Britain and Northern Ireland

for their discoveries concerning the molecular structure of nucleic acids and its significance for information transfer in living material

James Dewey Watson

United States of America

Francis Harry Compton Crick

United Kingdom of Great Britain and Northern Ireland

Maurice Hugh Frederick Wilkins

United Kingdom of Great Britain and Northern Ireland, New Zealand

for his discoveries of the physical mechanism of stimulation within the cochlea

Georg von Békésy

United States of America

for the discovery of acquired immunological tolerance

Frank Macfarlane Burnet

Australia

Peter Medawar

United Kingdom of Great Britain and Northern Ireland

for their discovery of the mechanisms in the biological synthesis of ribonucleic acid and deoxyribonucleic acid

Severo Ochoa de Albornoz

United States of America

Arthur Kornberg

United States of America

for their discovery that genes act by regulating definite chemical events

George Wells Beadle

United States of America

Edward Lawrie Tatum

United States of America

for his discoveries relating to synthetic compounds that inhibit the action of certain body substances, and especially their action on the vascular system and the skeletal muscles

Daniel Bovet

Italy

for their discoveries concerning heart catheterization and pathological changes in the circulatory system

André Frédéric Cournand

United States of America

Dickinson W. Richards

United States of America

Werner Forssmann

West Germany

for his discoveries concerning the nature and mode of action of oxidation enzymes

Hugo Theorell

Sweden

for the discovery of the ability of poliomyelitis viruses to grow in cultures of various types of tissue

John Franklin Enders

United States of America

Thomas Huckle Weller

United States of America

Frederick Chapman Robbins

United States of America

for the discovery of the citric acid cycle

Hans Adolf Krebs

United Kingdom of Great Britain and Northern Ireland

for his discovery of streptomycin, the first antibiotic effective against tuberculosis

Selman Waksman

United States of America

for his discoveries concerning yellow fever and how to combat it

Max Theiler

Union of South Africa

for their discoveries relating to the hormones of the adrenal cortex, their structure and biological effects

Edward Calvin Kendall

United States of America

Tadeusz Reichstein

Switzerland

Philip Showalter Hench

United States of America

for his discovery of the functional organization of the interbrain as a coordinator of the activities of the internal organs

Walter Rudolf Hess

Switzerland

for his discovery of the high efficiency of DDT as a contact poison against several arthropods

Paul Hermann Müller

Switzerland

for their discovery of the course of the catalytic conversion of glycogen

Carl Ferdinand Cori

United States of America

Gerty Theresa Cori

United States of America

for the discovery of the production of mutations by means of X-ray irradiation

Hermann Joseph Muller

United States of America

for the discovery of penicillin and its curative effect in various infectious diseases

Alexander Fleming

United Kingdom of Great Britain and Northern Ireland

Ernst Boris Chain

United Kingdom of Great Britain and Northern Ireland

Howard Walter Florey

Australia

for their discoveries relating to the highly differentiated functions of single nerve fibres

Joseph Erlanger

United States of America

Herbert Spencer Gasser

United States of America

for the discovery of vitamin K

Henrik Carl Peter Dam

Denmark

for the discovery of the antibacterial effects of prontosil

Gerhard Domagk

Greater German Reich

for the discovery of the role played by the sinus and aortic mechanisms in the regulation of respiration

Corneille Jean François Heymans

Belgium

for his discoveries in connection with the biological combustion processes, with special reference to vitamin C and the catalysis of fumaric acid

Albert Szent-Györgyi de Nagyrápolt

Kingdom of Hungary

for their discoveries relating to chemical transmission of nerve impulses

Henry Hallett Dale

United Kingdom of Great Britain and Northern Ireland

Otto Loewi

United States of America

for his discovery of the organizer effect in embryonic development

Hans Spemann

Greater German Reich

for their discoveries concerning liver therapy in cases of anaemia

George Hoyt Whipple

United States of America

George Richards Minot

United States of America

William Parry Murphy

United States of America

for his discoveries concerning the role played by the chromosome in heredity

Thomas Hunt Morgan

United States of America

for their discoveries regarding the functions of neurons

Charles Scott Sherrington

United Kingdom of Great Britain and Northern Ireland

Edgar Douglas Adrian

United Kingdom of Great Britain and Northern Ireland

for his discovery of the nature and mode of action of the respiratory enzyme

Otto Heinrich Warburg

German Reich

for the discovery of human blood groups

Karl Landsteiner

Austria

Discovery of the antineuritic vitamin (vitamin B1)

Christiaan Eijkman

Netherlands

for his work on typhus

Charles Jules Henri Nicolle

France

for his discovery of the therapeutic value of malaria inoculation in the treatment of dementia paralytica

Julius Wagner-Jauregg

Austria

for his discovery of the Spiroptera carcinoma and the idea that a parasitic infection can induce cancer

Johannes Andreas Grib Fibiger

Denmark

for his discovery of the mechanism of the electrocardiogram

Willem Einthoven

Netherlands

for the discovery of insulin

Frederick Grant Banting

Canada

John James Rickard Macleod

United Kingdom of Great Britain and Northern Ireland

for his discovery relating to the production of heat in the muscle

Archibald Vivian Hill

United Kingdom of Great Britain and Northern Ireland

for his discovery of the capillary motor regulating mechanism

Schack August Steenberg Krogh

Denmark

for his discoveries relating to immunity

Jules Bordet

Belgium

for his work on the physiology and pathology of the vestibular apparatus

Róbert Bárány

Austro-Hungarian Empire

for elucidating the mechanisms and dangers of anaphylaxis (acute hypersensitivity reactions)

Charles Robert Richet

France

for his work on vascular suture and the transplantation of blood vessels and organs

Alexis Carrel

France

for his work on the dioptrics of the eye

Allvar Gullstrand

Sweden

for contributions to our knowledge of cell chemistry made through his work on proteins, including the nucleic substances

Albrecht Kossel

German Empire

for his work on the physiology, pathology and surgery of the thyroid gland

Emil Theodor Kocher

Switzerland

for their work on immunity

Ilya Ilyich Mechnikov

Russian Empire

Paul Ehrlich

German Empire

for his work on the role played by protozoa in causing diseases

Charles Louis Alphonse Laveran

France

for their work on the structure of the nervous system

Camillo Golgi

Kingdom of Italy

Santiago Ramón y Cajal

Spain

for his investigations and discoveries in relation to tuberculosis

Robert Koch

German Empire

for his work on the physiology of digestion, through which knowledge on vital aspects of the subject has been transformed and enlarged

Ivan Petrovich Pavlov

Russian Empire

for his contribution to the treatment of diseases, especially lupus vulgaris, with concentrated light radiation, whereby he has opened a new avenue for medical science

Niels Ryberg Finsen

Denmark

for his work on malaria, by which he showed how it enters the organism and thereby laid the foundation for successful research on this disease and methods of combating it

Ronald Ross

United Kingdom of Great Britain and Northern Ireland

for his work on serum therapy, especially its application against diphtheria, by which he has opened a new road in the domain of medical science and thereby placed in the hands of the physician a victorious weapon against illness and deaths

Emil Adolf von Behring

German Empire