Just To Start Things OFF ,,,,,,im going to post up a small biography for Mr Robbert W Allard,,,
hopefully we all learn a thing or 2 ...
thankyou for any contrabutions you guys might make during this thread,,,,thankyou for just being here
N A T I O N A L A C A D E M Y O F S C I E N C E S
R O B E R T W A Y N E A L L A R D
1 9 1 9 – 2 0 0 3
A Biographical Memoir by
M I C H A E L T . C L E G G
Any opinions expressed in this memoir are those of the author
and do not necessarily reflect the views of the
National Academy of Sciences.
Biographical Memoirs
COPYRIGHT 2006
NATIONAL ACADEMY OF SCIENCES
WASHINGTON, D.C.
ROBERT WAYNE ALLARD
September 3, 1919–March 25, 2003
BY MICHAEL T. CLEGG
ROBERT (“BOB”) WAYNE ALLARD made wide-ranging contributions
to both basic and applied plant genetics. He began
as a plant breeder and wrote one of the most successful
plant-breeding texts of his era, but his most important contributions
were in evolutionary genetics. He was a founder
of experimental plant population genetics and he infused
the field with high standards of experimental and theoretical
rigor. His investigations ranged from elegant experiments
to dissect the genetic factors responsible for quantitative
genetic variation, to the study of gene-environment
interactions, to the analysis of selection in long-term experimental
barley populations. But his most significant work
was encompassed in a series of papers on the genetics of
inbreeding populations, where he overturned conventional
dogma by showing that inbreeding plant populations have
substantial levels of genetic variation. In the course of his
work on inbreeding species, he turned to the characterization
of the genetics of wild and naturalized species and
contributed to the origins of the field of plant ecological
genetics. He was also a teacher par excellence, training more
than 50 Ph.D. students and an even larger number of
postdoctoral students over a career that spanned more than
4 BIOGRAPHICAL MEMOIRS
50 years, and he led the University of California, Davis,
Genetics Department to preeminence during the 1960s and
1970s.
EARLY INFLUENCES
Bob Allard was born in the San Fernando Valley of California
on September 3, 1919. In the years between the two
world wars the San Fernando Valley was largely pastoral
and Bob’s early years were spent on the family farm. Around
1930 his father relocated his farming operation to the San
Joaquin Valley about 15 miles west of Modesto. Like many
farmers of the era, Bob’s father cooperated with University
of California agricultural researchers by dedicating a portion
of his land to experimental trials. A UC Berkeley plant
breeder named W. W. Mackie maintained plots of lima and
common beans on the Allard farm, and Bob was assigned
the task of assisting Mackie with the maintenance of the
experimental plots. This turned out to be the formative
experience of Bob’s young life, because Mackie instilled in
Bob a lifelong fascination with the causes of phenotypic
variation. In an oral history interview, Bob much later recalled
that Mackie introduced him to the new science of
Mendelian genetics during this period, thereby contributing
to his later choice of scientific career.
In recounting these early experiences, Bob would passionately
describe the pleasure he took in listening to Mackie
and in hearing his theories about genetic variation and its
practical exploitation. Bob was not a man to dwell on the
past; he strongly preferred to look toward the future. His
occasional recollections of Mackie were exceptional and reflected
the enduring impact of this period on his later scientific
development. According to Bob’s much later memories,
Mackie was also interested in the ecological bases of
adaptation and he introduced Bob to other plant species
ROBERT WAYNE ALLARD 5
common in their Central Valley environment, including the
slender wild oat (Avena barbata) that would later feature
importantly in some of Bob’s research.
It seems likely that Mackie influenced Bob’s decision to
enter UC Davis as a student of agriculture in the fall of
1937. During his undergraduate years Bob worked as a student
assistant for Coit Suneson of the U.S. Department of
Agriculture, and this also had an enduring impact on Bob.
Suneson, along with Harry Harlan and Gus Wiebe, was engaged
in developing bulk populations of wheat and barley,
known as composite cross populations. The theory at the
time was that bulk populations would both act as a reservoir
for useful genetic variation while at the same time evolving
toward greater adaptation under standard agricultural
conditions. Years later Bob would use these composite cross
populations as a powerful resource for studies in experimental
population genetics. These early experiences did
much to define Bob’s approaches to plant genetics and
they serve to illustrate the powerful impact that scientific
mentors can have on young minds.
After finishing his undergraduate training, Bob entered
the graduate program at the University of Wisconsin, Madison.
Certainly the biggest thing that happened to him at
Madison was meeting and marrying Ann, his wife of 59
years. On the rare occasions when Bob would talk about his
graduate school days, his chief recollection was being called
into World War II service just prior to the scheduled date
for his final dissertation defense. It seems that the university
would not reschedule the defense, and Bob had to return
to Madison after the war to defend his thesis. Bob’s
Ph.D. research was on wheat cytogenetics, and aside from
publishing his dissertation work following the war, he never
returned to this topic. There were strong influences at Madison,
including Rubush G. Shands (his major professor),
6 BIOGRAPHICAL MEMOIRS
Charles E. Allen, and R. A. Brink, but I always had the
feeling that Bob had a clear idea of his future research
directions by the time he left Davis.
After entering World War II service, Bob was sent to
the Naval Supply School at Harvard. Later he was assigned
to a research unit at Fort Detrick, Maryland, where he was
engaged in work on biowarfare, a subject he never discussed,
except to say that he had been in a research unit for part of
the war. Still, this provided Bob’s only postdoctoral training
and broadened his research experience.
In 1946 Bob returned to UC Davis as assistant professor
of agronomy and assistant geneticist in the Agricultural Experiment
Station, and he remained at Davis affiliated with
the Agronomy Department throughout his career. He was
hired as a bean breeder and his particular focus was on the
improvement of the lima bean. At that time Davis was a
branch of the Berkeley College of Agriculture and had little
autonomy. It was also a very small school with fewer than
800 students, most of whom were there for two-year terminal
degrees in agriculture. Bob was an important player in
a faculty generation that turned Davis from a small satellite
agriculture campus into a thriving and world-renowned university.
From the beginning Bob’s work blended both basic genetics
and practical plant improvement. In the initial years
he focused on both the identification of disease-resistance
genes and applications of the backcross method of breeding
for the incorporation of disease resistance into elite
lines of lima beans. The search for disease resistance genes
led to an extended field trip to Central and South America
to collect wild relatives and primitive land race materials as
genetic resources for future breeding efforts. He later published
an article for the California Dry Bean Research Conference
on plant exploring in Latin America. The conservaROBERT
WAYNE ALLARD 7
tion of genetic resources remained an abiding interest, one
that was communicated to a number of Bob’s students.
At heart Bob was a geneticist, and along with his practical
work on lima bean improvement, he began to develop
genetic markers in lima beans. These were largely seed coat
markers based on an amazing range of seed coat color patterns.
Bob and his early students patiently dissected the
inheritance of these discrete color polymorphisms and then
employed them as markers to study adaptive change in the
lima bean. A particularly fascinating aspect of the color
patterns was the interactions between different genetic factors
that lead to the mosaic patterns evident on the seed
coats. We learn and generalize from our empirical experiences,
and these are based on the materials that we choose
to study. In Bob’s case the theme of gene interaction, based
in part on his observations of seed coat color patterns in
the lima bean, continued to dominate his thinking throughout
his career.
The practical side of Bob’s program prospered in these
early years. He released a number of new varieties of lima
beans; one variety, “Mackie,” was named for his childhood
mentor. He also began work on a novel plant-breeding text.
The book, Principles of Plant Breeding, published in 1960
had an enormous impact and was ultimately translated into
17 languages. It remained the premier plant-breeding text
for a generation. The book was novel because it emphasized
genetic principles rather than methods and this contributed
to its great success. Bob was also a very fine writer
and this, too, contributed to the wide acceptance of Principles
of Plant Breeding. He took great pains with everything
he wrote, and the result was always a model of clarity
and precision. Bob would not put his name on a paper
until he had worked through it carefully, reanalyzed the
data, and improved the exposition. He did not believe in
8 BIOGRAPHICAL MEMOIRS
honorary authorships and he was very economical with citations.
His practice was to cite only essential supporting papers.
For many years Bob’s plant-breeding colleagues urged
him to write a second edition of Principles of Plant Breeding,
and he promised to do so, but it was not until 1999,
almost 15 years after Bob’s retirement and just four years
before his death that a second edition was published. Bob
admitted that the second edition was really an entirely new
book that contained little carried over from the parent book
published 39 years earlier. The second book is really a plant
population genetics book that synthesizes a life’s study of
plant evolution. It is uniquely Bob, both in the lucidity of
the writing and in the presentation and articulation of his
vision of evolutionary genetics.
QUANTITATIVE GENETICS
The field of quantitative genetics had a large impact on
agricultural research in the 1940s and 1950s. The origins of
quantitative genetics derive from R. A. Fisher’s 1918 paper
reconciling Mendelian genetics and Darwinian evolution
by natural selection. Quantitative, or biometrical, genetics
aims to partition phenotypic variation into genetic and environmental
components and it provides a scientific basis
for designing efficient schemes for selection. In later years
Bob recalled that while a student at Wisconsin, he had been
influenced by Sewall Wright, who along with R. A. Fisher
was the other great architect of the field of quantitative
genetics. It is clear from Bob’s later recollections that he
was anxious to move beyond lima bean breeding by mastering
the skills of quantitative genetics. During the academic
year 1954-1955, Bob found the opportunity to hone his skills
in statistics and quantitative genetics by taking a year’s sabbatical
leave in Birmingham, England, to work with KenROBERT
WAYNE ALLARD 9
neth Mather, one of the era’s leaders in quantitative genetics.
A few years later, in 1960, he returned to England to
work at Oxford with Norman J. T. Bailey, a leading statistician
in the field of mathematical genetics. These sabbaticals
had an enduring impact on Bob’s research directions.
In the middle 1950s Bob began to publish papers that
attacked various biometrical issues of the day. One paper
was devoted to maximum likelihood estimators for recombination,
others focused on the analysis of various diallele
crosses, and still others concerned the problem of estimating
gene-environment interactions. He began publishing
more frequently in broadly based genetics journals rather
than in agricultural journals so that his papers would reach
a broader audience of geneticists. He also continued to
publish on applied topics throughout his career. One paper
of this period that deserves special mention is an elegant
dissection of the genetics of heading time in wheat
(1965). In this paper Bob showed that a major gene controlled
heading date, but he went beyond this to show how
the remaining phenotypic variation in heading date could
be resolved into additional genetic components, revealing
the influence of multiple genetic factors of unequal effect.
The paper pushed the approaches of quantitative genetics
to their experimental limits. By this time Bob’s research
had evolved beyond the lima bean to exploit other plant
species more appropriate for investigating basic questions
of quantitative genetics. By the early 1960s Bob’s lab was
regarded as a leading lab for the study of plant quantitative
genetics. Even as he achieved this goal, Bob was moving in
new directions.
THE GENETICS OF INBREEDING POPULATIONS
Stimulated in part by his colleague G. Ledyard Stebbins,
Bob began to investigate the genetics of inbreeding spe10
BIOGRAPHICAL MEMOIRS
cies. In his classic 1950 book, Variation and Evolution in
Plants, Stebbins had claimed that inbreeding plant populations
should be largely devoid of genetic variation. The
argument put forward by Stebbins was that inbreeding leads
to homozygosity and the superior inbred type should outcompete
all other lines leading to a homogeneous population.
Bob knew from his plant-breeding experiences that
inbreeding crops, such as lima beans, had large stores of
genetic variation and showed rapid genetic responses to
selection. Stebbins had repeated what was the conventional
dogma of the time, but this provided the stimulus for Bob
to begin what became a classic series of experiments to
characterize genetic variation in inbreeding plant species.
Stebbins, for his part, encouraged this effort to look more
deeply at the genetics of inbreeding species. The quest led
Bob into an entirely new field, ecological genetics, which
sought to combine population genetics with ecology, where
Bob played a foundational role. It also began a series of
papers on population studies in predominantly self-pollinated
species that spanned a period of more than 20 years.
The studies of inbreeding populations led Bob from
quantitative genetics into population genetics. Bob quickly
established the leading program on plant population genetics
of the 1960s, and he and his students found novel
ways of approaching the fundamental questions of this field.
One important innovation harked back to the composite
cross populations of his early undergraduate days. At the
time, population genetics was dominated by Drosophila, partly
because the short generation time of Drosophila permitted
experiments over many generations, thereby allowing the
direct observation of evolutionary changes in gene frequencies.
Bob had become the custodian of the composite cross
populations, and he quickly realized that the populations
he had helped synthesize in his youth would allow a mulROBERT
WAYNE ALLARD 11
tiple generation approach in longer-lived annual plant species
as well. The basic reason rested on the fact that seed
could be stored over a number of years, allowing an investigator
to analyze gene frequencies in past generations. To
see how this worked it is necessary to describe the system
for propagating the composite cross populations. The practice
was to advance the populations each year by growing a
new generation under standard agricultural conditions at
Davis, while also storing a portion of seed from each year’s
harvest for several years. The saved seed would then be
rejuvenated by growing out a new generation every five years
or so. This provided a parallel series of populations that
represented early, intermediate, and late generations. By
the early 1960s the oldest populations had about a 30-year
history and the youngest had a history of only five or six
generations. Because of this scheme, the barley and wheat
composite cross populations provided a unique resource to
follow changes in phenotypic traits, gene frequencies, and
disease resistance loci over 30 or more generations.
Bob used every tool available to study genetic change in
the composite cross populations, beginning with simple
morphological polymorphisms and quantitative characters
and moving on to isozymes and finally to restriction fragment
length polymorphisms (RFLPs) near the end of his
career. Bob was among the first to adopt the isozyme method
when it appeared in the middle 1960s. Isozymes had an
enormous impact, because for the first time they allowed
the investigator to sample a large number of genes that
coded for various enzymatic proteins. Prior to this, students
of population genetics were limited to morphological variants,
such as the seed coat color polymorphisms of lima
bean or to quantitative traits where the underlying genes
were impossible to resolve. Isozymes allowed one to sample
many individual gene products and to ask questions about
12 BIOGRAPHICAL MEMOIRS
genome-wide levels of genetic variation. RFLPs offered the
advantages of isozymes while also permitting the investigator
to measure variation for portions of the genome that do
not code for enzymatic proteins. Throughout his career
Bob was always among the first to adopt new approaches to
address scientific questions. He was undaunted by obstacles
or by the investment of effort associated with acquiring new
technologies.
Regardless of the experimental approach employed in
studying the composite cross populations, substantial changes
in trait or gene frequencies were always observed over time,
and these were too large to be ascribed to genetic drift,
leaving selection as the only plausible explanation. The next
natural question was, could selection be quantified at individual
loci? Theodosius Dobzhansky and Sewall Wright had
developed approaches to the quantification of selection on
inversion polymorphisms in Drosophila pseudoobscura, but
these depended on the assumption of random mating. The
basic estimation technique was to derive transition equations
that predicted genotypic frequencies in one generation
based on their frequencies in previous generations after
accounting for the mating process. A set of weights that
mapped the predicted frequencies onto the observed frequencies
quantified the strength of selection.
Barley is a predominantly self-fertilizing plant, so the
random mating assumption could not be employed. A quantitative
theory of mating and a method to estimate the parameters
of such a quantitative model was required. A quantitative
theory, known as the mixed-mating model, which
allowed for a mixture of self-fertilization and random outcrossing,
had been published in 1951 by Fyfe and Bailey
(the same Bailey that Bob had worked with on sabbatical in
Oxford, England). Bob and his students employed this model
to estimate the single outcrossing parameter that indexed
ROBERT WAYNE ALLARD 13
the mixed-mating model and to derive transition equations
to estimate selection in the composite cross and other populations.
The technique for estimating the proportion of outcrossing
relied on another important property of plants;
the fact that one can easily collect numerous progeny of a
single maternal plant as seed. With the use of marker genes
it was possible to estimate the fraction of self-fertilization
and its complement—the fraction of outcrossing—from family
structured data.
Armed with a quantitative characterization of the mating
process one could quantify selection at individual marker
loci. But self-fertilization has another important consequence
that rendered it impossible to attribute selection to the marker
loci actually observed. Because self-fertilization leads to
homozygosity, effective recombination is greatly reduced and
any statistical associations among different loci decay slowly
over time. Populations like the composite cross populations,
with a relatively short history, would still retain statistical
associations between loci from their initial composition. Bob
and his students initiated the theoretical study of the behavior
of linkage disequilibrium (the technical term for
correlations between loci in allelic state) in mixed-mating
systems in the middle 1960s. At a time when computer simulations
were just beginning to be applied to genetic problems,
they published an important simulation study describing
the complex behavior of linkage disequilibrium in predominantly
self-fertilizing populations. Later estimates of the
magnitude of linkage disequilibrium in the composite cross
and other populations showed that it was typically large.
The conclusion was that chromosomal segments containing
the marker loci were subject to strong selection in virtually
all observed cases, but that one could not resolve selection
to the level of individual loci.
14 BIOGRAPHICAL MEMOIRS
Bob was not satisfied with the study of artificial populations.
The question he sought to answer was the broader
question concerning levels of genetic diversity in inbreeding
populations of plants in nature. By the early 1960s he
had launched a program to study natural populations of
inbreeding plants, and this work included a broad variety
of species, including Avena species (wild oats), other grasses
native to California, such as fescue, and annual native California
dicots, such as Collinsia species. These efforts began
an intensive period of ecological genetics research that
spanned nearly two decades. Avena barbata, the slender
wild oat, was a particular target of investigation during this
period. A. barbata is a naturalized component of the California
oak savannah that was introduced into California
during the Spanish Mission period from the Mediterranean
basin (almost certainly from Spain). The time dimension is
known, and this meant that genetic changes over a twohundred-
to three-hundred-year period could be documented.
Near the end of his life Bob recalled having been introduced
to Avena barbata by W. W. Mackie; once again
this powerful early influence determined a scientific direction,
and it was a fortunate choice, because A. barbata showed
markedly different patterns of evolution in different regions
of California. As later shown by two of Bob’s Spanish students
(Marcelino Perez de la Vega and Pedro Garcia Garcia),
these changes were not replicated in Spain, so they must
have arisen since the introduction of A. barbata to California.
Particularly dramatic were contrasting patterns of isozyme
variation between the foothills of the arid Central Valley of
California and the cooler and moister intermontane valleys
of the costal strip. The arid regions were nearly monomorphic
for a single multilocus genotype, while populations
from the coastal regions exhibited substantial levels of variROBERT
WAYNE ALLARD 15
ability. I was fortunate to play a role in these findings, and
it was a wonderful way to start a research career.
INTERACTING GENETIC SYSTEMS
A pervasive theme of Bob’s research and writing was the
importance of interactions among genes, between genes
and environments, and even among genotypes within populations.
Bob believed that context was essential and that
marginal effects were less important. I recall Bob attributing
this belief in the importance of interactions to his early
mentor W. W. Mackie. Regardless of the source, it clearly
dominated Bob’s thinking. This view went counter to conventional
population genetics theory that is based on the
notion that complex systems can be characterized by marginal
gene frequency changes. It also went counter to quantitative
genetics theory where additive effects were thought
to account for most variation. To this day the importance
of interaction remains an open question.
Beginning in the middle 1950s, Bob published experimental
work on gene environmental interactions. In the
1960s he turned to the problem of interactions among genes
at different loci. His approach of measuring linkage disequilibrium
as a surrogate for gene interactions was stimulated
by the theoretical calculations of R. C. Lewontin and
K. Kojima giving the precise relationship between selection
and recombination required for nonzero linkage disequilibria.
These highly simplified models showed that only nonadditive
selection over loci could retard recombination and
maintain permanent linkage disequilibrium. Thus Bob focused
on the estimation of linkage disequilibria in experimental
plant populations as a means of detecting interactions.
It later became clear that the existence of linkage
disequilibrium is neither necessary nor sufficient for the
existence of interlocus interactions, especially in inbreed16
BIOGRAPHICAL MEMOIRS
ing systems. Despite this, Bob did show that correlations
among loci could be pervasive in inbreeding plant populations
and that this would in turn affect their evolutionary
potential.
Together with his students, Bob studied the impact of
neighboring genotypes on the fitness of individual plants.
This system of intergenotypic interactions creates a frequency
dependent pattern of selection and widens the conditions
for the maintenance of a genetic polymorphism. As with
much of Bob’s work, theoretical calculations were supplemented
by direct measurements from experimental populations
to provide a predictive framework. Bob was also a
strong proponent of the idea that genetic mixtures would
perform better than single pure lines in an agricultural
context, although the evidence to support this view has been
meager.
A LIFE’S ACCOMPLISHMENTS
As noted above, Bob worked in, and in some cases helped
found, several distinct but related areas of plant genetics.
He had an enduring impact on plant breeding, largely
through his book but also through his early work in biometrical
genetics. These contributions were later recognized
through the DeKalb-Pfizer distinguished career award of
the Crop Science Society and the Crop Science Award of
the American Society of Agronomy. Bob was elected to the
National Academy of Sciences in 1973, where he chose to
affiliate with the genetics section and later with the section
on population biology, evolution, and ecology after it was
formed, rather than with the agricultural sections. This choice
illustrates that his first love was genetics, despite a lifelong
devotion to agriculture.
More than any other worker, Bob Allard is responsible
for laying the rigorous experimental foundations for plant
ROBERT WAYNE ALLARD 17
population genetics, and he played a major role in melding
the union of ecology and genetics that emerged as ecological
genetics. Perhaps his most enduring scientific legacy
was the series on population studies in predominantly selfpollinated
species. This series illustrated one of Bob’s greatest
strengths. He was first and foremost an empiricist who found
innovative ways to test theory and to expand our empirical
understanding of genetic systems. His belief in interaction
ran counter to the dogma of his time and often led to
intense arguments, but he never modified his views. He was
passionate about his scientific views and at times the strength
of his convictions seemed to overwhelm the available evidence.
In retrospect, his intuition was excellent and his
views have been largely vindicated.
Bob was a prolific teacher and mentor of graduate and
postdoctoral students. Altogether he trained 56 Ph.D.
students, and he hosted numerous visiting scientists and
postdoctoral students; he also trained a host of M.S. students.
He had a large number of international students,
and many have become prominent figures in countries
around the world. I recall students from all continents, and
as a consequence he left a global intellectual legacy. He
taught throughout his long tenure at UC Davis in both the
Department of Agronomy and the Department of Genetics.
He wrote a wonderful set of lecture notes on population
genetics that were used in the introductory genetics course
at Davis. During the late 1960s, I encountered his lecture
notes as an undergraduate and immediately decided I wanted
to study population genetics. He also taught in the introductory
genetics course for many years as well as a graduate
course in quantitative genetics and an undergraduate course
in population genetics. He was not a classroom performer,
but his lectures, like his writing, were clear and carefully
organized to make a central point.
18 BIOGRAPHICAL MEMOIRS
Around 1967 Bob became chair of the Department of
Genetics at UC Davis, where he served with energy and
skill. He played a major role in bringing Th. Dobzhansky
and F. J. Ayala to Davis in the early 1970s and helped catapult
the Department of Genetics to international preeminence.
At its peak the department included among its faculty
five members of the National Academy of Sciences. He
also served on virtually every major committee of the university
and for a period chaired the Davis division of the
UC Academic Senate. Bob was an active member of the
National Academy Sciences, where he chaired Section 27
for three years. He served on a number of National Research
Council committees, including a committee that produced
several volumes on managing global genetic resources.
He was unstintingly generous with his time, and he served
the university and the academy he loved with great devotion.
Bob retired in 1986 but remained very active. He published
a remarkable number of research papers during his
retirement, along with the new edition of his classic plantbreeding
book. During this period, Bob and Ann Allard
spent much of their time at their home at Bodega Bay on
the northern California coast. He loved walking on the seaside
cliffs examining plants, especially Avena barbata, and
speculating about their unique adaptations to the California
environment. He was always eager to entertain friends
and colleagues; evenings with the Allards at Bodega Bay
were very special events. Bob loved wine and was an accomplished
student and collector of fine wines, so any dinner
was resplendent with excellent wine. Bob finally had to leave
his Bodega Bay home about two years before his death,
owing to the onset of Alzheimer’s disease and circulatory
difficulties. He died on March 25, 2003, in Davis at the age
of 83.
ROBERT WAYNE ALLARD 19
SELECTED BIBLIOGRAPHY
1946
With H. R. DeRose and R. J. Weaver. Some effects of plant growth
regulators on seed germination and seedling development. Bot.
Gaz. 107:575-583.
1954
With R. G. Shands. The inheritance of resistance to stem rust and
powdery mildew in cytologically stable wheats derived from Triticum
timopheevi. Phytopathology 44:266-274
1956
The analysis of genic-environmental interactions by means of diallele
crosses. Genetics 41:305-318.
1960
Principles of Plant Breeding. New York: Wiley.
With S. K. Jain. 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. 46:1371-1377.
1963
With P. L. Workman. Population studies in predominantly self-pollinated
species. III. A matrix model for mixed selfing and random
outcrossing. Proc. Natl. Acad. Sci. U. S. A. 48:1318-1325.
1964
With J. Weil. The mating system and genetic variability in natural
populations of Collinsia heterophylla. Evolution 18:515-525.
1965
With C. Wehrhahn. The detection and measurement of the effects
of individual genes involved in the inheritance of a quantitative
character in wheat. Genetics 51:109-119.
20 BIOGRAPHICAL MEMOIRS
1966
With J. Harding and D. G. Smeltzer. Population studies in predominantly
self-pollinated species. IX. Frequency dependent selection
in Phaseolus lunatus. Proc. Natl. Acad. Sci. U. S. A. 56:99-104.
1967
With S. K. Jain and P. L. Workman. The genetics of inbreeding
populations. Adv. Genet. 14:55-131.
1970
With A. H. D. Brown. Estimation of the mating system in open
pollinated maize populations using isozyme polymorphisms. Genetics
66:133-145.
1972
With J. L. Hamrick. Microgeographical variation in allozyme frequencies
in Avena barbata. Proc. Natl. Acad. Sci. U. S. A. 69:2000-
2004.
With B. S. Weir and A. L. Kahler. Analysis of complex allozyme
polymorphisms in a barley population. Genetics 72:505-523.
1973
With M. T. Clegg. Viability versus fecundity selection in the slender
wild oat Avena barbata L. Science 181:667-668.
1977
With W. T. Adams. The effect of polyploidy on phosphoglucose
isomerase diversity in Festuca microstachys. Proc. Natl. Acad. Sci.
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1981
With D. V. Shaw and A. L. Kahler. A multilocus estimator of mating
system parameters in plant populations. Proc. Natl. Acad. Sci. U.
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1982
With O. Muona and R. K. Webster. Evolution of resistance to
Rhynchosporium secalis (Oud.) Davis in barley Composite Cross
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ROBERT WAYNE ALLARD 21
1984
With M. A. Saghai Maroof, R. A. Jorgensen, and K. Soliman. Ribosomal
DNA (rDNA) spacer-length (sl) variation in barley: Mendelian
inheritance, chromosomal location, and population dynamics.
Proc. Natl. Acad. Sci. U. S. A. 81:8014-1018.
1987
With D. B. Wagner, G. K. Furnier, M. A. Saghai Maroof, S. M.
Williams, and B. P. Dancik. Chloroplast DNA polymorphisms in
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