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PNAS Classics

PNAS Classics are landmark papers from the archives of PNAS—significant scientific reports that have withstood the test of time. The authors of PNAS Classics opened new avenues of research within their respective fields. These groundbreaking discoveries offer insights into how the best science is practiced. Each Classic article is accompanied by a Classic Perspective written by a scientist in the field and a Classic Profile written by a science writer. Together these newly commissioned articles recall the impact of these discoveries, describe how they have advanced the field, and set them in current scientific context.


Biochemistry: Phillip Sharp

Image courtesy of M. Scott Brauer, with permission of MIT News.

Perspective

Arnold J. Berk
Discovery of RNA splicing and genes in pieces
PNAS 2016 113:801
[Full Text] [PDF]

Classic Article

Susan M. Berget, Claire Moore, and Phillip A. Sharp
Spliced segments at the 5′ terminus of adenovirus 2 late mRNA
PNAS 1977 74:3171
[Full Text] [PDF]


Mathematics: Gérard Debreu

Image courtesy of Konrad Jacobs, copyright Mathematisches Forschungsinstitut Oberwolfach.

Commentary

Partha Sarathi Dasgupta, and Eric S. Maskin
Debreu’s social equilibrium existence theorem
PNAS 2015 112:15769
[Full Text] [PDF]

Classic Article

Gérard Debreu
A Social Equilibrium Existence Theorem
PNAS 1952 38:886
[Full Text] [PDF]


Medical Sciences: Robert C. Gallo

Image courtesy of the National Institute of Allergy and Infectious Diseases.

Perspective

John M. Coffin
The discovery of HTLV-1, the first pathogenic human retrovirus
PNAS 2015 112:15525
[Full Text] [PDF]

Classic Article

Bernard J. Poiesz, Francis W. Ruscetti, Adi F. Gazdar, Paul A. Bunn, John D. Minna, and Robert C. Gallo
Detection and isolation of type C retrovirus particles from fresh and cultured lymphocytes of a patient with cutaneous T-cell lymphoma
PNAS 1980 77:7415
[Full Text] [PDF]


Biochemistry: Peter H. Duesberg and Peter K. Vogt

Image courtesy of Mark Dastrup.

Commentary

Klaus Bister
Discovery of oncogenes: The advent of molecular cancer research
PNAS 2015 112:15259
[Full Text] [PDF]

Classic Article

Peter H. Duesberg and Peter K. Vogt
Differences between the Ribonucleic Acids of Transforming and Nontransforming Avian Tumor Viruses
PNAS 1970 67:1673
[Full Text] [PDF]


Neurobiology: Marcus E. Raichle

Image reprinted with permission from Elsevier.

Perspective

Manu S. Goyal, Siddarth Venkatesh, Jeffrey Milbrandt, Jeffrey I. Gordon, and Marcus E. Raichle
Feeding the brain and nurturing the mind: Linking nutrition and the gut microbiota to brain development
PNAS 2015 112:14105
[Full Text] [PDF]

Core Concept

Helen H. Shen
Core Concept: Resting-state connectivity
PNAS 2015 112:14115
[Full Text] [PDF]

Classic Article

Marcus E. Raichle, Ann Mary MacLeod, Abraham Z. Snyder, William J. Powers, Debra A. Gusnard, and Gordon L. Shulman
A default mode of brain function
PNAS 2001 98:676
[Full Text] [PDF]


Mathematics: Lloyd Shapley

Image courtesy of Shutterstock/PerseoMedusa.

Perspective

Eilon Solan and Nicolas Vieille
Stochastic games
PNAS 2015 112:13743
[Full Text] [PDF]

Classic Article

L. S. Shapley
Stochastic games
PNAS 1953 39:1095
[Full Text] [PDF]


Chemistry: Linus Pauling and Charles D. Coryell

Image courtesy of J. Barton.

Perspective

Kara L. Bren, Richard Eisenberg, and Harry B. Gray
Discovery of the magnetic behavior of hemoglobin: A beginning of bioinorganic chemistry
PNAS 2015 112:13123
[Full Text] [PDF]

Classic Article

Linus Pauling and Charles D. Coryell
The Magnetic Properties and Structure of the Hemochromogens and Related Substances
PNAS 1936 22:159
[Full Text] [PDF]

Classic Article

Linus Pauling and Charles D. Coryell
The Magnetic Properties and Structure of Hemoglobin, Oxyhemoglobin and Carbonmonoxyhemoglobin
PNAS 1936 22:210
[Full Text] [PDF]


Biology: Alfred J. Lotka

Image courtesy of MetLife Archives.

Profile

Sharon Kingsland
Alfred J. Lotka and the origins of theoretical population ecology
PNAS 2015 112:9493
[Full Text] [PDF]

Perspective

John W. Terborgh
Toward a trophic theory of species diversity
PNAS 2015 112:11415
[Full Text] [PDF]

Classic Article

Alfred J. Lotka
Analytical Note on Certain Rhythmic Relations in Organic Systems
PNAS 1920 6:410
[Full Text] [PDF]


Biochemistry: Tony Hunter and Bartholomew M. Sefton

Image courtesy of Shutterstock/bitsite.

Perspective

Tony Hunter
Discovering the first tyrosine kinase
PNAS 2015 112:7877
[Full Text] [PDF]

Inner Workings

Jessica Marshall
Inner Workings: Tyrosine kinases, their discovery and impact
PNAS 2015 112:7886
[Full Text] [PDF]

Classic Article

Tony Hunter and Bartholomew M. Sefton
Transforming gene product of Rous sarcoma virus phosphorylates tyrosine
PNAS 1980 77:1311
[Full Text] [PDF]


Astronomy: Edwin Hubble

Image courtesy of NASA.

Commentary

Neta A. Bahcall
Hubble’s Law and the expanding universe
PNAS 2015 112:3173
[Full Text] [PDF]

Inner Workings

David J. Harris
Inner Workings: Hubble’s quarter century in orbit has opened a universe of possibilities
PNAS 2015 112:3176
[Full Text] [PDF]

Classic Article

Edwin Hubble
A relation between distance and radial velocity among extra-galactic nebulae
PNAS 1929 15:168
[Full Text] [PDF]


Mathematics: George D. Birkhoff and J. v. Neumann

Image courtesy of Goethe University Frankfurt.

Perspective

Calvin C. Moore
Ergodic theorem, ergodic theory, and statistical mechanics
PNAS 2015 112:1907
[Full Text] [PDF]

Core Concept

Steven Ashley
Core Concept: Ergodic theory plays a key role in multiple fields
PNAS 2015 112:1914
[Full Text] [PDF]

Classic Article

J. v. Neumann
Proof of the Quasi-Ergodic Hypothesis
PNAS 1932 18:70
[Full Text] [PDF]

Classic Article

George D. Birkhoff
Proof of the Ergodic Theorem
PNAS 1931 17:656
[Full Text] [PDF]


Astronomy: W. Baade and F. Zwicky

Image courtesy of ESA/NASA.

Commentary

Adam S. Burrows
Baade and Zwicky: “Super-novae,” neutron stars, and cosmic rays
PNAS 2015 112:1242
[Full Text] [PDF]

Core Concept

Maggie McKee
Core Concept: Supernovae and the accelerating universe
PNAS 2015 112:1243
[Full Text] [PDF]

Classic Article

W. Baade and F. Zwicky
On Super-Novae
PNAS 1934 20:254
[Full Text] [PDF]

Classic Article

W. Baade and F. Zwicky
Cosmic Rays from Super-Novae
PNAS 1934 20:259
[Full Text] [PDF]


Evolution: Carl R. Woese and George E. Fox

Image courtesy of William Petri Jr. (University of Virginia, Charlottesville, VA).

Perspective

Jeremiah J. Faith, Jean-Frédéric Colombel, and Jeffrey I. Gordon
Identifying strains that contribute to complex diseases through the study of microbial inheritance
PNAS 2015 112:633
[Full Text] [PDF]

Inner Workings

Elie Dolgin
Inner Workings: Malnutrition, gutted
PNAS 2015 112:641
[Full Text] [PDF]

Retrospective

Larry Gold
The kingdoms of Carl Woese
PNAS 2013 9:3206
[Full Text] [PDF]

Classic Article

Carl R. Woese and George E. Fox
Phylogenetic structure of the prokaryotic domain: The primary kingdoms
PNAS 1977 74:5088
[Full Text] [PDF]


Physiology: M. S. Gazzaniga, J. E. Bogen, and R. W. Sperry

Commentary

Michael S. Gazzaniga
The split-brain: Rooting consciousness in biology
PNAS 2014 111:18093
[Full Text] [PDF]

Inner Workings

Helen H. Shen
Inner Workings: Discovering the split mind
PNAS 2014 111:18097
[Full Text] [PDF]

Classic Article

M. S. Gazzaniga, J. E. Bogen, and R. W. Sperry
Some functional effects of sectioning the cerebral commissures in man
PNAS 1962 48:1765
[Full Text] [PDF]


Cell Biology: Peter Novick and Randy Schekman

Logbook image courtesy of Peter Novick.

Commentary

Peter J. Novick
A pathway of a hundred genes starts with a single mutant: Isolation of sec1-1
PNAS 2014 111:9019
[Full Text] [PDF]

Inner Workings

Jessica Marshall
Inner Workings: Protein secretion
PNAS 2014 111:9021
[Full Text] [PDF]

Classic Article

Peter Novick and Randy Schekman
Secretion and cell-surface growth are blocked in a temperature-sensitive mutant of Saccharomyces cerevisiae
PNAS 1979 76:1862
[Full Text] [PDF]


Biochemistry: Marshall W. Nirenberg and J. Heinrich Matthaei

Courtesy of Marshall W. Nirenberg from the National Library of Medicine.

Commentary

C. Thomas Caskey and Philip Leder
The RNA code: Nature’s Rosetta Stone
PNAS 2014 111:5758
[Full Text] [PDF]

Inner Workings

Jessica Marshall
The genetic code
PNAS 2014 111:5760
[Full Text] [PDF]

Classic Article

Marshall W. Nirenberg and J. Heinrich Matthaei
The dependence of cell-free protein synthesis in E. coli upon naturally occurring or synthetic polyribonucleotides
PNAS 1961 47:1588
[Full Text] [PDF]


Medical Sciences: Michael S. Brown and Joseph L. Goldstein

Classic Profile

Prashant Nair
Brown and Goldstein: The Cholesterol Chronicles
PNAS 2013 110:14829
[Full Text] [PDF]

Classic Perspective

Randy Schekman
Discovery of the cellular and molecular basis of cholesterol control
PNAS 2013 110:14833
[Full Text] [PDF]

Classic Article

Michael S. Brown and Joseph L. Goldstein
Familial hypercholesterolemia: Defective binding of lipoproteins to cultured fibroblasts associated with impaired regulation of 3-hydroxy-3-methylglutaryl coenzyme a reductase activity
PNAS 1974 71:788
[Full Text] [PDF]


Zoology: Rita Levi-Montalcini

Image courtesy of Presidenza della Repubblica.

Classic Profile

Nicholette Zeliadt
Rita Levi-Montalcini: NGF, the prototypical growth factor
PNAS 2013 110:4873
[Full Text] [PDF]

Classic Perspective

Antonino Cattaneo
Immunosympathectomy as the first phenotypic knockout with antibodies
PNAS 2013 110:4877
[Full Text] [PDF]

Retrospective

Leslie Lars Iversen
Rita Levi-Montalcini: Neuroscientist par excellence
PNAS 2013 13:4862
[Full Text] [PDF]

Classic Article

Rita Levi-Montalcini and Barbara Booker
Destruction of the sympathetic ganglia in mammals by an antiserum to a nerve-growth protein
PNAS 1960 46:384
[Full Text] [PDF]


Genetics: Barbara McClintock

Photo by Ross Meurer. Image courtesy of the Barbara McClintock Collection, Cold Spring Harbor Laboratory Library and Archives.

Classic Profile

Sandeep Ravindran
Barbara McClintock and the discovery of jumping genes
PNAS 2012 109:20198
[Full Text] [PDF]

Classic Perspective

Nina V. Fedoroff
McClintock’s challenge in the 21st century
PNAS 2012 109:20200
[Full Text] [PDF]

Classic Article

Barbara McClintock
The origin and behavior of mutable loci in maize
PNAS 1950 36:344
[Full Text] [PDF]


Evolution: Carl R. Woese and George E. Fox

Photo by Jason Lindley; used with permission of the College of Liberal Arts and Sciences, University of Illinois at Urbana–Champaign.

Classic Perspective

Norman R. Pace, Jan Sapp, and Nigel Goldenfeld
Phylogeny and beyond: Scientific, historical, and conceptual significance of the first tree of life
PNAS 2012 109:1011
[Full Text] [PDF]

Classic Profile

Prashant Nair
Woese and Fox: Life, rearranged
PNAS 2012 109:1019
[Full Text] [PDF]

Classic Article

Carl R. Woese and George E. Fox
Phylogenetic structure of the prokaryotic domain: The primary kingdoms
PNAS 1977 74:5088
[Full Text] [PDF]


Biochemistry: Paul Greengard

Classic Perspective

Solomon H. Snyder
What dopamine does in the brain
PNAS 2011 108:18869
[Full Text] [PDF]

Classic Profile

Sandeep Ravindran
Paul Greengard: Signals underlying moods, addictions, and brain disorders
PNAS 2011 108:18872
[Full Text] [PDF]

Classic Article

John W. Kebabian, Gary L. Petzold, and Paul Greengard
Dopamine-sensitive adenylate cyclase in caudate nucleus of rat brain, and its similarity to the "dopamine receptor"
PNAS 1972 69:2145
[Full Text] [PDF]


Physics: Julian Schwinger

Image courtesy of the Smithsonian Institution Archives (SIA).

Classic Perspective

Silvan S. Schweber
The sources of Schwinger's Green's functions
PNAS 2005 102:7783
[Full Text] [PDF]

Classic Article (Part I)

Julian Schwinger
On the Green's functions of quantized fields I
PNAS 1951 37:452
[Full Text] [PDF]

Classic Article (Part II)

Julian Schwinger
On the Green's functions of quantized fields II
PNAS 1951 37:455
[Full Text] [PDF]


Biochemistry: Kathleen Danna and Daniel Nathans

Image courtesy of Susie Fitzhugh.

Classic Perspective

Richard J. Roberts
How restriction enzymes became the workhorses of molecular biology
PNAS 2005 102:5905
[Full Text] [PDF]

Classic Profile

Christen Brownlee
Danna and Nathans Restriction enzymes and the boon to modern molecular biology
PNAS 2005 102:5909
[Full Text] [PDF]

Classic Article

Kathleen Danna and Daniel Nathans
Specific Cleavage of Simian Virus 40 DNA by Restriction Endonuclease of Hemophilus Influenzae
PNAS 1971 68:2913
[Full Text] [PDF]


Genetics: Matthew Meselson and Franklin W. Stahl

Image courtesy of Caltech Archives.

Classic Perspective

Philip C. Hanawalt
Density matters: The semiconservative replication of DNA
PNAS 2004 101:17889
[Full Text] [PDF]

Classic Profile

Tinsley H. Davis
Meselson and Stahl: The art of DNA replication
PNAS 2004 101:17895
[Full Text] [PDF]

Classic Article

Matthew Meselson and Franklin W. Stahl
The replication of DNA in Escherichia Coli
PNAS 1958 44:671
[Full Text] [PDF]


Mathematics: John F. Nash

Image courtesy of Getty Images.

Classic Profile

Erica Klarreich
The Mathematics of Strategy
[Full Text]

Classic Perspective

Charles A. Holt and Alvin E. Roth
The Nash equilibrium: A perspective
PNAS 2004 101:3999
[Full Text] [PDF]

Classic Article

John F. Nash
Equilibrium points in n-person games
PNAS 1950 1:48
[Full Text] [PDF]


Astronomy: Edwin Hubble

Image courtesy of Caltech Archives.

Classic Profile

Christen Brownlee
Hubble's guide to expanding the universe
[Full Text]

Classic Perspective

Robert P. Kirshner
Hubble's diagram and cosmic expansion
PNAS 2004 101:8
[Full Text] [PDF]

Classic Article

Edwin Hubble
A relation between distance and radial velocity among extra-galactic nebulae
PNAS 1929 15:168
[Full Text] [PDF]


Developmental Biology: Robert Briggs and Thomas J. King

Image courtesy of Dr. Kitai Kim and M. Willy Lensch.

Classic Profile

Christen Brownlee
Nuclear transfer: Bringing in the clones
[Full Text]

Classic Perspective

J. B. Gurdon and J. A. Byrne
The first half-century of
nuclear transplantation
PNAS 2003 100:8048
[Full Text] [PDF]

Classic Article

Robert Briggs and Thomas J. King
Transplantation of living nuclei from blastula cells into enucleated frogs' eggs
PNAS 1952 38:455
[Full Text] [PDF]


Chemistry: Linus Pauling, Robert B. Corey, and H. R. Branson

Image courtesy of Caltech Archives.

Classic Profile

Christen Brownlee
The protein papers
[Full Text]

Classic Perspective

David Eisenberg
The discovery of the α-helix and β-sheet, the principal structural features
of proteins
PNAS 2003 100:11207
[Full Text] [PDF]

Classic Article

Linus Pauling, Robert B. Corey, and H. R. Branson
The structure of proteins: Two hydrogen-bonded helical configurations of the polypeptide chain
PNAS 1951 37:205
[Full Text] [PDF]

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