Leland H. Hartwell

Leland Hartwell was studying Saccharomyces cerevisiae, the baker’s yeast, when he discovered CDC (cell division cycle) genes. CDC genes are responsible for the regulation of the cell cycle and mutations in the gene cause cancerous growth in cells.

Hartwell showed how the cells grow and divide. His research showed that gaps in a cell cycle i.e. a pause allows the repairing of the damaged DNA. His work allowed the understanding of normal and abnormal cell growth and multiplication. In recognition of this research Hartwell received the Nobel Prize in Physiology or Medicine 2001 sharing it with Tim Hunt and Paul Nurse.

Hartwell also received the Albert Lasker Award for Basic Medical Research 1998, Medal of Merit 2003 and the Komen Brinker Award for Scientific Distinction.

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Biologist BIOLOGIST (cell cycle)
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1973 Eli Lilly Award in Microbiology and Immunology

1983-1984 Guggenheim Awardee and American Cancer Society Scholar

1990-present American Cancer Society Research Professor Genetics

1990 National Institutes of Health Merit Award

1991 General Motors Sloan Award

1991 Hoffman LaRoche Mattia Award

1992 Gairdner Foundation International Award

1992 University of Chicago Simon Shubitz Award

1993 Brandeis University Rosenstiel Award

1994 Sloan-Kettering Cancer Center Katherine Berkan Judd Award

1994 Genetics Society of America Medal

1995 MGH Warren Triennial Prize

1995 Columbia University Horwitz Award

1995 Keith Porter Award, Amer Soc Cell Biology

1996 Passano Award

1996 Carnegie Mellon Dickson Prize

1998 Albert Lasker Basic Medical Research Award

1998 Brinker International Award for Basic Science – Susan G. Komen Breast Cancer Foundation

1999 California Institute of Technology – Distinguished Alumni Award

1999 City of Medicine Award

1999 American Cancer Society Medal of Honor

2000 Léopold Griffuel Prize – Association pour la Recherche sur le Cancer, France

2000 The Massry Prize – The Meira and Shaul G. Massry Foundation

2001 The Nobel Prize in Physiology or Medicine

Hartwell, L; Mankoff, D; Paulovich, A; Ramsey, S; Swisher, E Cancer biomarkers: a systems approach. Nature biotechnology 2006;24(8):905-8.

Aebersold, R; Anderson, L; Caprioli, R; Druker, B; Hartwell, L; Smith, R Perspective: a program to improve protein biomarker discovery for cancer. Journal of proteome research 2005;4(4):1104-9.

Hartwell, LH Yeast and cancer. Bioscience reports 2004;24(4-5):523-44.

Hartwell, L Genetics. Robust interactions. Science (New York, N.Y.) 2004;303(5659):774-5.

Etzioni, R; Urban, N; Ramsey, S; McIntosh, M; Schwartz, S; Reid, B; Radich, J; Anderson, G; Hartwell, L The case for early detection. Nature reviews. Cancer 2003;3(4):243-52.

Hartwell, LH Nobel Lecture. Yeast and cancer. Bioscience reports 2002;22(3-4):373-94.

Hartwell, L Interview: Leland Hartwell, PhD, Nobel Prize for Medicine winner. MedGenMed : Medscape general medicine 2001;3(4):3.

Hartman, JL; Garvik, B; Hartwell, L Principles for the buffering of genetic variation. Science (New York, N.Y.) 2001;291(5506):1001-4.

Emili, A; Schieltz, DM; Yates, JR; Hartwell, LH Dynamic interaction of DNA damage checkpoint protein Rad53 with chromatin assembly factor Asf1. Molecular cell 2001;7(1):13-20.

Simon, JA; Szankasi, P; Nguyen, DK; Ludlow, C; Dunstan, HM; Roberts, CJ; Jensen, EL; Hartwell, LH; Friend, SH Differential toxicities of anticancer agents among DNA repair and checkpoint mutants of Saccharomyces cerevisiae. Cancer research 2000;60(2):328-33.

Hartwell, LH; Hopfield, JJ; Leibler, S; Murray, AW From molecular to modular cell biology. Nature 1999;402(6761 Suppl):C47-52.

Marton, MJ; DeRisi, JL; Bennett, HA; Iyer, VR; Meyer, MR; Roberts, CJ; Stoughton, R; Burchard, J; Slade, D; Dai, H; Bassett, DE; Hartwell, LH; Brown, PO; Friend, SH Drug target validation and identification of secondary drug target effects using DNA microarrays. Nature medicine 1998;4(11):1293-301.

Nurse, P; Masui, Y; Hartwell, L Understanding the cell cycle. Nature medicine 1998;4(10):1103-6.

Paulovich, AG; Armour, CD; Hartwell, LH The Saccharomyces cerevisiae RAD9, RAD17, RAD24 and MEC3 genes are required for tolerating irreparable, ultraviolet-induced DNA damage. Genetics 1998;150(1):75-93.

Brown, PO; Hartwell, L Genomics and human disease–variations on variation. Nature genetics 1998;18(2):91-3.

Hartwell, LH; Szankasi, P; Roberts, CJ; Murray, AW; Friend, SH Integrating genetic approaches into the discovery of anticancer drugs. Science (New York, N.Y.) 1997;278(5340):1064-8.

Mihich, E; Hartwell, L Eighth Annual Pezcoller Symposium: genomic instability and immortality in cancer. Cancer research 1997;57(19):4437-41.

Toczyski, DP; Galgoczy, DJ; Hartwell, LH CDC5 and CKII control adaptation to the yeast DNA damage checkpoint. Cell 1997;90(6):1097-106.

Schrick, K; Garvik, B; Hartwell, LH Mating in Saccharomyces cerevisiae: the role of the pheromone signal transduction pathway in the chemotropic response to pheromone. Genetics 1997;147(1):19-32.

Dorer, R; Boone, C; Kimbrough, T; Kim, J; Hartwell, LH Genetic analysis of default mating behavior in Saccharomyces cerevisiae. Genetics 1997;146(1):39-55.

Wooden, JM; Hartwell, LH; Vasquez, B; Sibley, CH Analysis in yeast of antimalaria drugs that target the dihydrofolate reductase of Plasmodium falciparum. Molecular and biochemical parasitology 1997;85(1):25-40.

Paulovich, AG; Toczyski, DP; Hartwell, LH When checkpoints fail. Cell 1997;88(3):315-21.

Paulovich, AG; Margulies, RU; Garvik, BM; Hartwell, LH RAD9, RAD17, and RAD24 are required for S phase regulation in Saccharomyces cerevisiae in response to DNA damage. Genetics 1997;145(1):45-62.

Pryciak, PM; Hartwell, LH AKR1 encodes a candidate effector of the G beta gamma complex in the Saccharomyces cerevisiae pheromone response pathway and contributes to control of both cell shape and signal transduction. Molecular and cellular biology 1996;16(6):2614-26.

Garvik, B; Carson, M; Hartwell, L Single-stranded DNA arising at telomeres in cdc13 mutants may constitute a specific signal for the RAD9 checkpoint. Molecular and cellular biology 1995;15(11):6128-38.

Dorer, R; Pryciak, PM; Hartwell, LH Saccharomyces cerevisiae cells execute a default pathway to select a mate in the absence of pheromone gradients. The Journal of cell biology 1995;131(4):845-61.

Paulovich, AG; Hartwell, LH A checkpoint regulates the rate of progression through S phase in S. cerevisiae in response to DNA damage. Cell 1995;82(5):841-7.

Hartwell, L 1994 forbeck cancer forum on cell cycle checkpoints. Clinical cancer research : an official journal of the American Association for Cancer Research 1995;1(9):1067.

Hartwell, LH; Kastan, MB Cell cycle control and cancer. Science (New York, N.Y.) 1994;266(5192):1821-8.

Hartwell, L Cell cycle. cAMPing out. Nature 1994;371(6495):286.

Weinert, TA; Kiser, GL; Hartwell, LH Mitotic checkpoint genes in budding yeast and the dependence of mitosis on DNA replication and repair. Genes & development 1994;8(6):652-65.

Hartwell, L; Weinert, T; Kadyk, L; Garvik, B Cell cycle checkpoints, genomic integrity, and cancer. Cold Spring Harbor symposia on quantitative biology 1994;59():259-63.

Dorer, R; Pryciak, P; Schrick, K; Hartwell, LH The induction of cell polarity by pheromone in Saccharomyces cerevisiae. Harvey lectures 1994;90():95-104.

Brown, MT; Goetsch, L; Hartwell, LH MIF2 is required for mitotic spindle integrity during anaphase spindle elongation in Saccharomyces cerevisiae. The Journal of cell biology 1993;123(2):387-403.

Weinert, TA; Hartwell, LH Cell cycle arrest of cdc mutants and specificity of the RAD9 checkpoint. Genetics 1993;134(1):63-80.

Kadyk, LC; Hartwell, LH Replication-dependent sister chromatid recombination in rad1 mutants of Saccharomyces cerevisiae. Genetics 1993;133(3):469-87.

Hartwell, L Defects in a cell cycle checkpoint may be responsible for the genomic instability of cancer cells. Cell 1992;71(4):543-6.

Kadyk, LC; Hartwell, LH Sister chromatids are preferred over homologs as substrates for recombinational repair in Saccharomyces cerevisiae. Genetics 1992;132(2):387-402.

Hartwell, L Bringing the basic scientist into human disease research. Molecular biology of the cell 1992;3(8):837-8.

Hartwell, LH Alfred P. Sloan, Jr. Prize. Role of yeast in cancer research. Cancer 1992;69(10):2615-21.

Hartwell, LH Twenty-five years of cell cycle genetics. Genetics 1991;129(4):975-80.

Jackson, CL; Konopka, JB; Hartwell, LH S. cerevisiae alpha pheromone receptors activate a novel signal transduction pathway for mating partner discrimination. Cell 1991;67(2):389-402.

Hartwell, L Yeast embryology. Pathways of morphogenesis. Nature 1991;352(6337):663-4.

Neiman, PE; Hartwell, LH Malignant instability. Workshop on Genetic Instability and its Role in Carcinogenesis sponsored by the Programs in Molecular Medicine of the Fred Hutchinson Cancer Research Center and the University of Washington, Seattle, WA, USA, January 11-12, 1991. The New biologist 1991;3(4):347-51.

Brown, M; Garvik, B; Hartwell, L; Kadyk, L; Seeley, T; Weinert, T Fidelity of mitotic chromosome transmission. Cold Spring Harbor symposia on quantitative biology 1991;56():359-65.

Weinert, TA; Hartwell, LH Characterization of RAD9 of Saccharomyces cerevisiae and evidence that its function acts posttranslationally in cell cycle arrest after DNA damage. Molecular and cellular biology 1990;10(12):6554-64.

Jackson, CL; Hartwell, LH Courtship in S. cerevisiae: both cell types choose mating partners by responding to the strongest pheromone signal. Cell 1990;63(5):1039-51.

Jackson, CL; Hartwell, LH Courtship in Saccharomyces cerevisiae: an early cell-cell interaction during mating. Molecular and cellular biology 1990;10(5):2202-13.

Hartwell, LH; Weinert, TA Checkpoints: controls that ensure the order of cell cycle events. Science (New York, N.Y.) 1989;246(4930):629-34.

Burke, D; Gasdaska, P; Hartwell, L Dominant effects of tubulin overexpression in Saccharomyces cerevisiae. Molecular and cellular biology 1989;9(3):1049-59.

Weinert, T; Hartwell, L Control of G2 delay by the rad9 gene of Saccharomyces cerevisiae. Journal of cell science. Supplement 1989;12():145-8.

Konopka, JB; Jenness, DD; Hartwell, LH The C-terminus of the S. cerevisiae alpha-pheromone receptor mediates an adaptive response to pheromone. Cell 1988;54(5):609-20.

Weinert, TA; Hartwell, LH The RAD9 gene controls the cell cycle response to DNA damage in Saccharomyces cerevisiae. Science (New York, N.Y.) 1988;241(4863):317-22.

Cross, F; Hartwell, LH; Jackson, C; Konopka, JB Conjugation in Saccharomyces cerevisiae. Annual review of cell biology 1988;4():429-57.

Koshland, D; Hartwell, LH The structure of sister minichromosome DNA before anaphase in Saccharomyces cerevisiae. Science (New York, N.Y.) 1987;238(4834):1713-6.

Jenness, DD; Goldman, BS; Hartwell, LH Saccharomyces cerevisiae mutants unresponsive to alpha-factor pheromone: alpha-factor binding and extragenic suppression. Molecular and cellular biology 1987;7(4):1311-9.

Koshland, D; Rutledge, L; Fitzgerald-Hayes, M; Hartwell, LH A genetic analysis of dicentric minichromosomes in Saccharomyces cerevisiae. Cell 1987;48(5):801-12.

Meeks-Wagner, D; Wood, JS; Garvik, B; Hartwell, LH Isolation of two genes that affect mitotic chromosome transmission in S. cerevisiae. Cell 1986;44(1):53-63.

Meeks-Wagner, D; Hartwell, LH Normal stoichiometry of histone dimer sets is necessary for high fidelity of mitotic chromosome transmission. Cell 1986;44(1):43-52.

Jenness, DD; Burkholder, AC; Hartwell, LH Binding of alpha-factor pheromone to Saccharomyces cerevisiae a cells: dissociation constant and number of binding sites. Molecular and cellular biology 1986;6(1):318-20.

Burkholder, AC; Hartwell, LH The yeast alpha-factor receptor: structural properties deduced from the sequence of the STE2 gene. Nucleic acids research 1985;13(23):8463-75.

Carson, MJ; Hartwell, L CDC17: an essential gene that prevents telomere elongation in yeast. Cell 1985;42(1):249-57.

Hartwell, LH; Smith, D Altered fidelity of mitotic chromosome transmission in cell cycle mutants of S. cerevisiae. Genetics 1985;110(3):381-95.

Koshland, D; Kent, JC; Hartwell, LH Genetic analysis of the mitotic transmission of minichromosomes. Cell 1985;40(2):393-403.

Jenness, DD; Burkholder, AC; Hartwell, LH Binding of alpha-factor pheromone to yeast a cells: chemical and genetic evidence for an alpha-factor receptor. Cell 1983;35(2 Pt 1):521-9.

Dutcher, SK; Hartwell, LH Test for temporal or spatial restrictions in gene product function during the cell division cycle. Molecular and cellular biology 1983;3(7):1255-65.

Dutcher, SK; Hartwell, LH Genes that act before conjugation to prepare the Saccharomyces cerevisiae nucleus for caryogamy. Cell 1983;33(1):203-10.

Wood, JS; Hartwell, LH A dependent pathway of gene functions leading to chromosome segregation in Saccharomyces cerevisiae. The Journal of cell biology 1982;94(3):718-26.

Dutcher, SK; Hartwell, LH The role of S. cerevisiae cell division cycle genes in nuclear fusion. Genetics 1982;100(2):175-84.

Hartwell, LH Mutants of Saccharomyces cerevisiae unresponsive to cell division control by polypeptide mating hormone. The Journal of cell biology 1980;85(3):811-22.

Hartwell, LH Cell division from a genetic perspective. The Journal of cell biology 1978;77(3):627-37.

Hartwell, LH; Unger, MW Unequal division in Saccharomyces cerevisiae and its implications for the control of cell division. The Journal of cell biology 1977;75(2 Pt 1):422-35.

Reid, BJ; Hartwell, LH Regulation of mating in the cell cycle of Saccharomyces cerevisiae. The Journal of cell biology 1977;75(2 Pt 1):355-65.

Johnston, GC; Pringle, JR; Hartwell, LH Coordination of growth with cell division in the yeast Saccharomyces cerevisiae. Experimental cell research 1977;105(1):79-98.

Hartwell, LH Sequential function of gene products relative to DNA synthesis in the yeast cell cycle. Journal of molecular biology 1976;104(4):803-17.

Unger, MW; Hartwell, LH Control of cell division in Saccharomyces cerevisiae by methionyl-tRNA. Proceedings of the National Academy of Sciences of the United States of America 1976;73(5):1664-8.

Hartwell, LH Saccharomyces cerevisiae cell cycle. Bacteriological reviews 1974;38(2):164-98.

Hereford, LM; Hartwell, LH Sequential gene function in the initiation of Saccharomyces cerevisiae DNA synthesis. Journal of molecular biology 1974;84(3):445-61.

Hartwell, LH; Culotti, J; Pringle, JR; Reid, BJ Genetic control of the cell division cycle in yeast. Science (New York, N.Y.) 1974;183(4120):46-51.

Hartwell, LH Three additional genes required for deoxyribonucleic acid synthesis in Saccharomyces cerevisiae. Journal of bacteriology 1973;115(3):966-74.

Hereford, LM; Hartwell, LH Role of protein synthesis in the replication of yeast DNA. Nature: New biology 1973;244(135):129-31.

Hartwell, LH; Mortimer, RK; Culotti, J; Culotti, M Genetic Control of the Cell Division Cycle in Yeast: V. Genetic Analysis of cdc Mutants. Genetics 1973;74(2):267-86.

Shulman, RW; Hartwell, LH; Warner, JR Synthesis of ribosomal proteins during the yeast cell cycle. Journal of molecular biology 1973;73(4):513-25.

Bücking-Throm, E; Duntze, W; Hartwell, LH; Manney, TR Reversible arrest of haploid yeast cells in the initiation of DNA synthesis by a diffusible sex factor. Experimental cell research 1973;76(1):99-110.

Hartwell, LH Synchronization of haploid yeast cell cycles, a prelude to conjugation. Experimental cell research 1973;76(1):111-7.

Hereford, LM; Hartwell, LH Defective DNA synthesis in permeabilized yeast mutants. Nature: New biology 1971;234(49):171-2.

Hartwell, LH Genetic control of the cell division cycle in yeast. IV. Genes controlling bud emergence and cytokinesis. Experimental cell research 1971;69(2):265-76.

Culotti, J; Hartwell, LH Genetic control of the cell division cycle in yeast. 3. Seven genes controlling nuclear division. Experimental cell research 1971;67(2):389-401.

Hartwell, LH Genetic control of the cell division cycle in yeast. II. Genes controlling DNA replication and its initiation. Journal of molecular biology 1971;59(1):183-94.

Hartwell, LH Periodic density fluctuation during the yeast cell cycle and the selection of synchronous cultures. Journal of bacteriology 1970;104(3):1280-5.

Hartwell, LH; Culotti, J; Reid, B Genetic control of the cell-division cycle in yeast. I. Detection of mutants. Proceedings of the National Academy of Sciences of the United States of America 1970;66(2):352-9.

Martin, TE; Hartwell, LH Resistance of active yeast ribosomes to dissociation by KCl. The Journal of biological chemistry 1970;245(6):1504-6.

Hartwell, LH; McLaughlin, CS; Warner, JR Identification of ten genes that control ribosome formation in yeast. Molecular & general genetics : MGG 1970;109(1):42-56.

Hartwell, LH; Hutchison, HT; Holland, TM; McLaughlin, CS The effect of cycloheximide upon polyribosome stability in two yeast mutants defective respectively in the initiation of polypeptide chains and in messenger RNA synthesis. Molecular & general genetics : MGG 1970;106(4):347-61.

Hartwell, LH Biochemical genetics of yeast. Annual review of genetics 1970;4():373-96.

McLaughlin, CS; Magee, PT; Hartwell, LH Role of isoleucyl-transfer ribonucleic acid synthetase in ribonucleic acid synthesis and enzyme repression in yeast. Journal of bacteriology 1969;100(2):579-84.

Hutchison, HT; Hartwell, LH; McLaughlin, CS Temperature-sensitive yeast mutant defective in ribonucleic acid production. Journal of bacteriology 1969;99(3):807-14.

McLaughlin, CS; Hartwell, LH A mutant of yeast with a defective methionyl-tRNA synthetase. Genetics 1969;61(3):557-66.

Hartwell, LH; McLaughlin, CS A mutant of yeast apparently defective in the initiation of protein synthesis. Proceedings of the National Academy of Sciences of the United States of America 1969;62(2):468-74.

McLaughlin, CS; Hartwell, LH Mutants of yeast defective i

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  • Postings

    President and Director
    Fred Hutchinson Cancer Research Center in Seattle, Washington