Robert W. Allard

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Robert Wayne Allard
Robert W. Allard.jpg
Robert Wayne Allard

(1919-09-03)September 3, 1919
DiedMarch 25, 2003(2003-03-25) (aged 83)
Spouse(s)Ann Catherine Wilson Allard
ChildrenSusan Ann
Thomas Wilson (deceased)
Jane Ellen
Gillian Elizabeth
Stacie Roberta
AwardsDeKalb-Pfizer Distinguished Career
Crop Science Science of America
Member, National Academy of Sciences 1973
Fellow, American Academy of Arts and Sciences 1985

Robert "Bob" Wayne Allard (September 3, 1919 – March 25, 2003) was an American plant breeder and plant population geneticist who is widely regarded as one of the leading plant population geneticists of the 20th century.[1] Allard became Chair of the Genetics Department at University of California, Davis in 1967; he was elected to the National Academy of Sciences in 1973, and was awarded the DeKalb-Pfizer Distinguished Career Award and the Crop Science Science of America Award. He was honored as the Nilsson-Ehle Lecturer of the Mendelian Society of Sweden and as the Wilhelmine Key lecturer of the American Genetic Association. He also served as president of the Genetics Society of America, the American Genetic Association and the American Society of Naturalists.[2]

As Chair of the Department of Genetics at U.C. Davis he played a major role in bringing Theodosius Dobzhansky and Francisco J. Ayala to the Genetics Department in the early 1970s.[3] With G. Ledyard Stebbins there as well, the department became preeminent in both plant and animal evolution. He trained 56 doctorate students and more than 100 masters students over his career.


Allard Staff U.C. Davis Staff Editor 1939

Allard entered U.C. Davis as an undergraduate in 1937, and attended the University of Wisconsin–Madison for graduate training. His PhD work was interrupted by World War II, where he engaged in biowarfare research. After the War he returned to U.W. Madison to defend his Ph.D. thesis on wheat cytogenetics.[4][5]

R.W. Allard as U.S. Navy Ensign, 1944

Early career and research[edit]

He joined the faculty at U.C. Davis in 1946, where he was hired as a plant breeder in the Agronomy Department. He worked on developing varieties of lima beans, in addition to studying the inheritance of its seed coat polymorphisms as well as this plant's basic genetics.[6] He also worked on wheat diseases.[7]

In the 1950s he also started doing research in the area of quantitative genetics,[8] and by the 1960s he had broadened his use of experimental plants to include barley.[9][10]

Additionally at this time he published perhaps his most important publication, the 1960 book "Principles of Plant Breeding",[11] which was translated into 17 languages, and was the principal plant breeding book for at least a generation. A second edition was published in 1999.[12]

His ever-expanding interests led him into plant population genetics, a field that he essentially founded. His particular interest was in the population genetics of Inbreeding species, and included wild plants such as Collinsia, Avena barbata and Avena fatua. His interest in inbreeding species probably stemmed from his interactions with Ledyard Stebbins, who was also at U.C. Davis, and had predicted that wild inbreeding plant populations should be devoid of genetic variation. As a plant breeder who had done selection on inbreeding species in cultivated plants, Allard knew that there was ample genetic variation within "pure lines." It is easy to see how he would have become intrigued with inbreeding wild species. And indeed starting in the early 1960s he worked with experimental populations of wheat, barley, and lima beans as well as the wild plants Collinsia, Avena barbata, and Avena fatua investigating a wide variety of issues important to the population genetics of inbreeding species. He published numerous foundational papers on the effects of mating systems, selection, gene-environment interactions, linkage disequilibrium, and genetic drift on the genetic variation of inbreeding plant populations.[13][14][15][16][17][18][19]

Starting at the latter part of the 1960s and to the end of his career, Allard and co-workers focused primarily in a new area for his lab, that of ecological genetics, particularly in the plant Avena barbata. This work was greatly assisted at the end of the 1960s and throughout the 1970s and 1980s by the use of molecular markers known as isozymes combined with traditional quantitative genetics and morphological single-marker traits[20][21][22][23] [24][25][26][27][28][29][30][31][32][33][34] as well as more advanced molecular techniques as these became available in the 1980s and 1990s and were applied to Avena barbata[35] and other species such as wild and cultivated barley[36][37] and pines.[38]

An over-riding theme of Allard's lifetime work was the demonstration of "favorable epistatic combinations of alleles of different loci", or multilocus gene complexes in wild and cultivated plants that were assembled in and adapted to specific habitats. To illustrate this theme his last two papers focused on two in-breeding species (the slender wild oat Avena barbata and barley Hordeum vulgare) and one outcrossing species, corn (Zea maize).[39][40]


  1. ^ Clegg, Michael T. (2006). Robert Wayne Allard:1919-2003. Washington D.C.: National Academy of Sciences. p. 21.
  2. ^ "Yolo County Obituaries-A". The California USGenWeb Project Yolo County. Retrieved 22 May 2017.
  3. ^ Clegg, Michael T. (2006). Robert Wayne Allard:1919-2003. Washington D.C.: National Academy of Sciences. p. 21.
  4. ^ Allard, Robert Wayne (1946). A cytogenetic study of the effect of backcrossing to common wheat in a hybrid between Triticum vulgare Vill. and Triticum timopheevi Zhuk. University of Wisconsin, Madison.
  5. ^ Allard, R. W. (1949). "A cytogenetic study dealing with the transfer of genes from Triticum timopheevi to common wheat by backcrossing". Journal of Agricultural Research. 78 (3–4): 33–64. PMID 18113659.
  6. ^ Allard, R.W.; Clement, W.M. (1959). "Linkage in lima beans". Journal of Heredity. 50 (2): 63–67. doi:10.1093/oxfordjournals.jhered.a106876.
  7. ^ R.G., Shands; Allard, R.W. (1954). "The inheritance of resistance to stem rust and powdery mildew in cytologically stable wheats derived from Triticum timopheevi". Phytopathology. 44: 266–274.
  8. ^ Alllard, Robert (1956). "The analysis of genic-environmental interactions by means of diallele crosses". Genetics. 41 (3): 305–318. PMC 1209783. PMID 17247629.
  9. ^ Fasoulas, A.C.; Allard, R.W. (1962). "Nonallelic Gene Interactions in the Inheritance of Quantitative Characters in Barley". Genetics. 47 (7): 899–907. PMC 1210382. PMID 17248119.
  10. ^ Wehrhahn, C; Allard, R.W. (1965). "The Detection and Measurement of the Effects of Individual Genes Involved in the Inheritance of a Quantitative Character in Wheat". Genetics. 51 (1): 109–119. PMC 1210759. PMID 17248230.
  11. ^ Allard, R.W. (1960). "Principles of Plant Breeding". Soil Science. 91 (6): 414. Bibcode:1961SoilS..91..414A. doi:10.1097/00010694-196106000-00017.
  12. ^ Allard, R.W. (1999). Principles of Plant Breeding, 2nd Edition. John Wiley & Sons. p. 264. ISBN 978-0-471-02309-8.
  13. ^ Jain, S.K. (1960). "Population studies in predominantly self-pollinated species. I. Evidence for heterozygote advantage in a closed population of barley". Proc. Natl. Acad. Sci. U.S.A. 46: 1371–77. doi:10.1073/pnas.46.10.1371.
  14. ^ Workman, P.L.; Allard, R.W. (1963). "Population studies in predominantly self-pollinated species. III. A matrix model for mixed selfing and random outcrossing". Proc. Natl. Acad. Sci. U.S.A. 48 (8): 1318–1325. Bibcode:1962PNAS...48.1318W. doi:10.1073/pnas.48.8.1318. PMC 220951. PMID 16590980.
  15. ^ Weil, J.; Allard, R.W. (1964). "The mating system and genetic variability in natural populations of Collinsia heterophylla". Evolution. 18 (4): 515–525. doi:10.1111/j.1558-5646.1964.tb01668.x.
  16. ^ Imam, A.G.; Allard, R.W. (1965). "Population Studies in Predominantly Self-Pollinated Species. VI. Genetic Variability between and within Natural Populations of Wild Oats from Differing Habitats in California". Genetics. 51 (1): 49–62. PMC 1210767. PMID 17248232.
  17. ^ Harding, J.; Allard, R.W.; Smelzer, D.G. (1966). "Population studies in predominantly self-pollinated species. IX. Frequency dependent selection in Phaseolus lunatus". Proc. Natl. Acad. Sci. U.S.A. 56 (1): 99–104. Bibcode:1966PNAS...56...99H. doi:10.1073/pnas.56.1.99. PMC 285681. PMID 5229860.
  18. ^ Jain, S. K.; Marshall, D.R. (1967). "Population studies in predominantly self-pollinating species. X. Variation in natural populations of Avena fatua and Avena barbata". American Naturalist. 101 (917): 19–33. doi:10.1086/282465.
  19. ^ Allard, R.W.; Jain, S.K.; Workman, P.L. (1967). "The genetics of inbreeding populations". Adv. Genetics. Advances in Genetics. 14: 55–131. doi:10.1016/S0065-2660(08)60425-3. ISBN 9780120176144.
  20. ^ Marshall, D.R.; Allard, R.W. (1969). "The genetics of electrophoretic variants in Avena: I. The esterase e4 e9 e10 phosphatase p5 and anodal peroxidase apx5 loci in a. barbata". Journal of Heredity. 60: 17–19. doi:10.1093/oxfordjournals.jhered.a107919.
  21. ^ Marshall, D.R.; Allard, R.W. (1970). "Maintenance of Isozyme Polymorphisms in Natural Populations of Avena barbata". Genetics. 66 (2): 393–399. PMC 1212502. PMID 17248512.
  22. ^ Hamrick, J.L.; Allard, R.W. (1972). "Microgeographical Variation in Allozyme Frequencies in Avena barbata". Proceedings of the National Academy of Sciences of the United States of America. 69 (8): 2100–2104. Bibcode:1972PNAS...69.2100H. doi:10.1073/pnas.69.8.2100. PMC 426877. PMID 16592002.
  23. ^ Clegg, M.T.; Allard, R.W. (1972). "Patterns of Genetic Differentiation in the Slender Wild Oat Species Avena barbata". Proceedings of the National Academy of Sciences of the United States of America. 69 (7): 1820–1824. Bibcode:1972PNAS...69.1820C. doi:10.1073/pnas.69.7.1820. PMC 426810. PMID 16591999.
  24. ^ Allard, R.W.; Babbel, G.R.; Clegg, M.T.; Kahler, A.L. (1972). "Evidence for coadaptation in Avena barbata". Proceedings of the National Academy of Sciences of the United States of America. 69 (10): 3043–3048. Bibcode:1972PNAS...69.3043A. doi:10.1073/pnas.69.10.3043. PMC 389703. PMID 4342975.
  25. ^ Miller, R.D. (1977). "Genetic Variability in the Slender Wild Oat Avena barbata in California". Ph.D. Dissertation, University of California, Davis.
  26. ^ Allard, R.W.; Miller, R.D.; Kahler, A.L. (1978). The relationship between degree of environmental heterogeneity and genetic polymorphism. IN: 'Structure and functioning of plant populations'. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeling Naturkunde, Tweede Reeks, deel 70.
  27. ^ Hakim-Elahi, A. (1980). "Temporal changes in the population structure of the slender wild oat (Avena barbata) as measured by allozyme polymorphisms". Ph.D. Dissertation, University of California, Davis.
  28. ^ Price, S.C. (1980). "Polymorphism and phenomorphism in the tetraploid slender wild oat Avena barbata". Ph.D. Dissertation University of California, Davis.
  29. ^ Hutchinson, E.S. (1982). "Genetic markers and ecotypic differentiation of Avena bartata Pott ex Link". Ph.D. Dissertation, University of California, Davis.
  30. ^ Pinero, D. (1982). "Correlations between enzyme phenotypes and physical environment in California populations of Avena barbata and Avena fatua". Ph.D. Dissertation, University of California, Davis.
  31. ^ Cluster, P.D. (1984). "Correlation between genetic variation for allo-zyme markers and quantitative characters in Avena barbata Pott. ex. Link". Ph.D. Dissertation, University of California, Davis.
  32. ^ Price, S.C.; Shumaker, K.M.; Kahler, A.L.; Allard, R.W.; Hill, J.E. (1984). "Estimates of population differentiation obtained from enzyme polymorphisms and quantitative characters". Journal of Heredity. 75 (2): 141–142. doi:10.1093/oxfordjournals.jhered.a109889.
  33. ^ Price, S.C.; Hill, J.E.; Allard, R.W. (1988). "The Morphological and Physiological Response of Slender Oat (Avena barbata) to the Herbicides Barban and Difenzoquat". Weed Science. 36: 60–69. doi:10.1017/s0043174500074464.
  34. ^ Garcia, F.J.; Perez de la Vega, M.; Allard, R.W. (1989). "Allelic and genotypic composition of ancestral Spanish and colonial Californian gene pools of Avena barbata: Evolutionary implications". Genetics. 122 (3): 687–694. PMC 1203742. PMID 2759424.
  35. ^ Cluster, P.D.; Allard, R.W. (1995). "Evolution of ribosomal DNA (rDNA) genetic structure in colonial Californian populations of Avena barbata". Genetics. 139 (2): 941–954. PMC 1206392. PMID 7713443.
  36. ^ Saghai-Maroof, M.A.; Soliman, K.M.; Jorgensen, R.A. (1984). "Ribosomal DNA spacer-length polymorphisms in barley: mendelian inheritance, chromosomal location, and population dynamics". Proceedings of the National Academy of Sciences of the United States of America. 81 (24): 8014–8018. Bibcode:1984PNAS...81.8014S. doi:10.1073/pnas.81.24.8014. PMC 392284. PMID 6096873.
  37. ^ Neale, D.B.; Saghai-Maroof, M.A.; Allard, R.W.; Zhang, Q.; Jorgensen, R.A. (1988). "Chloroplast DNA diversity in populations of wild and cultivated barley". Genetics. 120 (4): 1105–1110. PMC 1203573. PMID 2906305.
  38. ^ Wagner, D.B.; Furnier, G.R.; Saghai-Maroof, M.A.; Williams, S.M.; Dancik, B.P.; Allard, R.W. (1987). "Chloroplast DNA polymorphisms in lodgepole and jack pines and their hybrids". Proceedings of the National Academy of Sciences of the United States of America. 84 (7): 2097–2100. Bibcode:1987PNAS...84.2097W. doi:10.1073/pnas.84.7.2097. PMC 304592. PMID 3470779.
  39. ^ Allard, R.W. (1996). "Genetic Basis of the Evolution of Adaptedness in Plants". Euphytica. 92 (1–2): 1–11. doi:10.1007/bf00022822.
  40. ^ Allard, R.W. (1999). "History of Plant Population Genetics". Annual Review of Genetics. 33: 1–27. doi:10.1146/annurev.genet.33.1.1. PMID 10690402.

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