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D. S. Falconer and Introduction to Quantitative Genetics

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Perspectives
Anecdotal, Historical and Critical Commentaries on Genetics
Edited by James F. Crow and William F. Dove

D. S. Falconer and Introduction to Quantitative Genetics

William G. Hill*,1 and Trudy F. C. Mackay†

*School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3JT, United Kingdom and
†
Department of Genetics, North Carolina State University, Raleigh, North Carolina 27695

D OUGLAS Falconer died on February 23, 2004, in

Edinburgh. In his career he made many important
contributions to quantitative genetics and to the genet-
in science, being both more able and older than his
contemporaries and, judging by his later years, a clearer
thinker and more organized. He took an honors degree
ics of the mouse. Undoubtedly, however, his greatest in zoology under D’Arcy Thompson (of Growth and Form
impact was through his textbook, Introduction to Quanti- fame), but did not study any genetics. Falconer (unpub-
tative Genetics (Falconer 1960a), which for many lished notes) wrote that Thompson presided over the
throughout the world has been both their lead into and final year course, “but did almost nothing. I asked him
their lifelong reference on the subject. The first edition at the beginning for recommendations as to what to
was published in 1960 when one of us (W.G.H.) was read and he said ‘Just browse, my boy, just browse.’ So
an undergraduate student in agriculture and becoming I worked away on my own . . . and at the end of the
interested in animal breeding; the book (albeit ad- year he came along to me and said ‘Well, Douglas, my
vanced for that naı̈ve student) gave a lead into the sci- boy, you’re a very good lad and I don’t think we need
ence behind the practice. The fourth edition in 1996 was give you an examination this year.’” But the Dean pre-
co-authored by T.F.C. Mackay (Falconer and Mackay vailed and Douglas had to take an exam, in which he
1996). We both have read it many times, consulted it, received First Class Honors. He was unfit for military
and taught from it. The book has been part of our lives. service and took his Ph.D. in 1943 under James Gray at
In this Perspectives we shall give some brief biographical Cambridge on “The behavior of wireworms in relation
information about Douglas, who was our teacher, col- to temperature and light,” which is less esoteric than it
league, and friend, and a summary of some of his most appears as the wireworm is a major crop pest and there
significant research. We shall concentrate, however, on was a wartime need to increase agricultural production;
the development and impact of Introduction to Quantita- but Douglas did not find it inspiring.
tive Genetics. While still a student he met and married Margaret
Douglas Falconer’s family, which came from Edin- Duke, a classicist, and they had two sons; all three survive
burgh, had no significant tradition of science. His father him. Among their shared interests was a love of music,
was a minister in the church when Douglas was born which they played together. Douglas played the flute
on March 10, 1913, in Old Meldrum, Aberdeenshire, and continued to do so until he was well into his eighties.
but the family soon moved back to Edinburgh, where In midlife Douglas developed diabetes, which he man-
Douglas went to school. There he developed an interest aged well, but he became increasingly blind in his last
in science, although no biology was taught. He was ac- years. To those who met him from at least the 1960s
cepted for university but his studies were delayed by 5 onward, he already appeared a frail individual, but his
years when he contracted tuberculosis. During those frailness belied his determination and toughness. He
long years of recuperation he read widely, including was still coming into work and writing long after his
Morgan’s Theory of the Gene. Finally, in 1936 he resumed retirement in 1980, not the least on further editions
his education at St. Andrews University, where he shone of his book, and he still kept an interest and enjoyed
scientific discussion until his death.
Subsequent to completing his Ph.D., Douglas had a
1
Corresponding author : Institute of Cell, Animal and Population Biol- temporary lectureship at Queen Mary College London,
ogy, School of Biological Sciences, University of Edinburgh, West
Mains Rd., Edinburgh EH9 3JT, United Kingdom. then based in Cambridge, where his teaching included
E-mail: [email protected] a course in genetics. At the end of the war, the Agricul-
Genetics 167: 1529–1536 ( August 2004)
1530 W. G. Hill and T. F. C. Mackay

Douglas Falconer in his office in 1987. (Photo-

graph by Antonia Reeve.)

tural Research Council (ARC) was planning an institute able pairs to mate, and I found out that he would not,
for the study of genetics in relation to animal breeding that he would find that many of his lines would become
extinct because there was not a suitable pair to continue
and was bringing a group together, initially near Lon- them. I thought I ought to tell him of this finding and
don, but later to move to Edinburgh. Seeing this pros- see what he thought about it, so I did one day at tea. He
pect, in 1945 Douglas took the opportunity to work with listened to what I said and then without uttering a word,
R. A. (Sir Ronald) Fisher at Cambridge (whose mouse he turned round and walked out of the room. The next
labs were in his own house). Douglas set to work on day he came to me and said, “I think you should arrange
to go to Edinburgh as soon as you can.”
mapping mutant genes and also to work on the inheri-
tance of milk yield in mice (recording the trait as the Douglas Falconer was appointed to the Genetics sec-
increase in litter weight after feeding, not with a me- tion of the ARC Animal Breeding and Genetics Research
chanical milking machine) but his time with Fisher was Organization in Edinburgh in 1947, under the direction
not entirely satisfactory. He recalls in an anecdote: of Professor C. H. Waddington, newly appointed to the
Fisher was engaged in a complicated experiment involv- Buchanan Chair of Animal Genetics at the University
ing a large number of mouse genes, and also involving of Edinburgh. A very illustrious group were brought
inbreeding, that is, brother by sister mating. Inbreeding together. In addition to Falconer, those in quantitative
is well known to reduce fertility, and I thought it would
be interesting to see how much the fertility would be genetics included Alan Robertson, Jim Rendel (who left
reduced in the next two or three generations, and see in 1951 for Australia), Eric Reeve, Forbes Robertson,
whether he would have sufficient animals to choose suit- and Ian Mason and, later, Crad Roberts and George
 Perspectives 1531

Clayton. In related areas in the ARC or university during Biometrical Genetics, which dealt little with the kinds of
the late 1940s and 1950s were Alan Beatty, Charlotte randomly mating outbred populations and selection is-
Auerbach, Geoffrey Beale, Mick Callan, Toby Carter, sues encountered by animal and human geneticists; and
Ruth Clayton, Henrik Kacser, Mary Lyon, James Sang, Kempthorne’s (1957) An Introduction to Genetic Statistics.
and Barnet Woolf. In an earlier Perspectives, Falconer Perhaps the most comprehensive source was Lush’s
(1993) gives a much fuller account of the personnel (1948) notes, The Genetics of Populations, but these were
and activities of their groups. (We joined the group then available only as a mimeograph (posthumously
much later: W.G.H. was a student of Alan Robertson published; Lush 1994).
from 1963 and then on the university staff from 1965; Aimed at filling this void, and specifically to help
T.F.C.M was a student of Alan Robertson from 1976 to students in the Edinburgh courses, Douglas Falconer
1979 and then a staff member from 1980 to 1987.) wrote Introduction to Quantitative Genetics, the first edition
Waddington essentially allowed people to work with lit- of which was published in 1960 (Falconer 1960a). In
tle direction; in fact, the set up was much more like a the preface he states his objectives:
modern university department with independently
My aim in writing this book has been to provide an intro-
funded, self-directed groups, rather than the traditional ductory textbook of quantitative genetics, with the em-
hierarchical European university department. phasis on general principles rather than on practical ap-
plication, and one moreover that can be understood by
biologists of no more than ordinary mathematical ability.
INTRODUCTION TO QUANTITATIVE GENETICS, In pursuit of this latter aim I have set out the mathematics
FIRST EDITION in the form that I, being little of a mathematician, find
most comprehensible, hoping that the consequent lack
Quantitative genetics was then a little-known subject of rigour and elegance will be compensated for by a wider
and training was necessary. Douglas had spent time with accessibility. The reader is not, however, asked to accept
Fisher but had worked mostly on linkage. Alan Robert- conclusions without proof. . . . I have no particular class
of reader in mind, but have tried to make the book useful
son, however, spent a period in the United States with
to as wide a range of readers as possible (p. v).
Sewall Wright and Jay L. Lush prior to coming to Edin-
burgh; and there were important sabbatical visitors to In the preface, he also gave some significant specific
Edinburgh in the early days. Notable were Michael Ler- thanks:
ner in 1948, who wrote much of his Population Genetics
It is no exaggeration to say that without Dr. Alan Robert-
and Animal Improvement (Lerner 1950) while in Edin- son’s help this book could not have been written. Not only
burgh, and Wright in 1949, who gave a course of lec- has his reading of the manuscript led to the elimination
tures, parts of which appeared in a long methodological of many errors, but I have been greatly assisted in my
paper (Wright 1952), which were the basis of the first understanding of the subject, particularly its more mathe-
volume of his treatise (Wright 1968). Notwithstanding matical aspects, by frequent discussions with him (p. vi).
the contact Douglas and the Edinburgh group had with He also thanks Waddington for provision of facilities,
Wright and Lush, it was the analysis of variance structure Crad Roberts for reading the entire manuscript, and
and the methodology of Fisher (1918) rather than the the honors and diploma students for raising questions
path coefficient approach of Wright that Douglas used that led to the improvement of many topics.
in his book. The first five chapters dealt with the genetics of single
In the late 1940s there was a dearth of teaching of genes, perhaps more appropriately termed population
any branch of genetics in Britain, and from 1949, in genetics, namely Hardy-Weinberg, selection, inbreed-
addition to honors undergraduate courses, a postgradu- ing, and genetic drift. This is, however, important back-
ate Diploma in Genetics was started at the University of ground for the quantitative genetics that follow from
Edinburgh, taught by the university, ARC, and other chapter 6 on, and indeed was and remains a succinct and
staff. This included courses in quantitative genetics and clear introduction to the subject. It is again instructive to
its applications to livestock improvement, taught by show Falconer’s insight and clarity of writing by quoting
Douglas and colleagues. from the beginning of chapter 6, “Continuous Varia-
At that time no textbooks that covered all or indeed tion”:
much of the relevant material were available. C. C. Li’s
(1955) Population Genetics dealt mainly with the subject It will be obvious, to biologist and layman alike, that
the sort of variation discussed in the foregoing chapters
at the level of individual genes, and the books or long embraces but a small part of the naturally occurring varia-
papers by Fisher, Haldane, and Wright provided neither tion. One has only to consider one’s fellow men and
a comprehensive nor a simple coverage. Among the women to realize that they all differ in countless ways,
most relevant books were Lerner’s (1950) Population but that these differences are nearly all matters of degree
Genetics and Animal Improvement and The Genetic Basis of and seldom present clear-cut distinctions attributable to
the segregation of single genes. If, for example, we were
Selection (Lerner 1958), but their aims were more lim- to classify individuals according to their height, we could
ited; Lush’s (1945) Animal Breeding Plans, which con- not put them into groups labeled “tall” and “short,” be-
tained even less background theory; Mather’s (1949) cause there are all degrees of height, and a division into
1532 W. G. Hill and T. F. C. Mackay

classes would be purely arbitrary. Variation of this sort, The second edition was published in 1981, with the fol-
without natural discontinuities, is called continuous varia- lowing comments in the preface (Falconer 1981, p. ix):
tion, and characters that exhibit it are called quantitative
characters or metric characters, because their study depends In preparing this revised edition, my aims have been
on measurement instead of on counting. The genetic (1) to keep the character of the book, and its length,
principles underlying the inheritance of metric characters unchanged; (2) to include some account of all the main
are basically those outlined in the previous chapters, but developments in the last twenty years; and (3) to be less
since the segregation of the genes concerned cannot be neglectful of plants.
followed individually, new methods of study have had to
be developed and new concepts introduced. A branch The headings of the 20 chapters were unchanged,
of genetics has consequently grown up, concerned with except chapter 16, “Inbreeding and Crossbreeding: III.
metric characters, which is called variously population ge- The utilisation of heterosis,” became more broadly “…
netics, biometrical genetics or quantitative genetics. The impor- III. Applications.” Apart from adding new experimental
tance of this branch of genetics need hardly be stressed;
most of the characters of economic value to plant and data, including some from his own selection lines, the
animal breeders are metric characters, and most of the main changes were inclusions of new or expanded sec-
changes concerned in micro-evolution are changes of tions on the influence of assortative mating on correla-
metric characters. It is therefore in this branch that genet- tions among relatives, on the effects of selection on
ics has its most important application to practical prob- variance in the infinitesimal model (the “Bulmer ef-
lems and also its most direct bearing on evolutionary
theory (p. 104).
fect”), on variability among replicate selection lines, on
selection limits theory, and on interpretation of long-
After this introductory paragraph, in which only stylis- term response. Chapter 18 on threshold characters was
tic changes have been made through the fourth edition, substantially changed, not least to reflect his own impor-
Falconer goes on to explain how “the intrinsically dis- tant work. In response to a query from Cyril Clarke,
continuous variation caused by genetic segregation is Falconer (1965, 1967) had proposed and used a
translated into the continuous variation of metric char- method for analysis of human data on diseases with all-
acters.” or-none expression but complex inheritance, including,
for example, diabetes from which Douglas himself suf-
fered. In this he showed that the heritability of “liability”
LATER EDITIONS (a term he did not introduce until the second edition)
could be computed simply from two numbers, the inci-
The first edition of his book was a major success. It
dence of the disease in the population and in relatives
was reprinted five times by Oliver & Boyd and twice by
of affected individuals. The method uses the population
Longman. Douglas realized the increasing need for a
incidence to compute the base mean and selection in-
new edition, but other commitments prevented his giv-
tensity and the incidence in relatives to compute the
ing time to it.
“response,” thereby providing the realized heritability
He had been appointed Deputy Director of the ARC
of liability.
Unit of Animal Genetics under Waddington, but “Wad”
The third edition (Falconer 1989) contained less
became less interested in genetics and also went on a
revision. The main changes in the text were to incorpo-
long sabbatical. In 1968 Falconer was appointed to a
rate more on mutation in quantitative genetics and a
Personal Chair in Genetics at the University of Edin-
little on mixed model analysis, including REML and
burgh and also Director of the ARC Unit. The following
BLUP, and to incorporate his set of problems, previously
year he was appointed Head of the Department of Ge- published separately (see below).
netics with responsibilities for all the diverse groups, Recognizing that some sections were becoming out-
including other major ones in mutation research, epi- dated and that he was less up to date with the subject
genetics, and protozoan genetics, housed in several (he was in his eighties), Douglas asked T.F.C.M. to join
buildings (later known as the Genetics Village) on the as a co-author of the fourth edition, published in 1996
King’s Buildings campus. This was a heavy task, which (Falconer and Mackay 1996). The basic structure of
Douglas undertook in an even-handed manner, notwith- the book was unaltered, but the main changes are out-
standing the competing claims of many prima donnas, lined in the preface:
giving it all the commitment he gave to other work
including his research, which he continued throughout Quantitative genetics is now merging with molecular ge-
netics and this very active area of the subject needs more
this time. In 1977 he was able to give up the Headship consideration than it was given in the previous edition.
of the Department following John Fincham’s appoint- Accordingly, a new chapter has been added, on quantita-
ment to the Buchanan Chair. He could then devote tive trait loci (QTL)—the location and characterization
time to revising his book, both prior to his formal retire- of the genes causing quantitative variation. Chapter 20,
ment and consequent closure of the ARC Unit of Animal on natural selection, has been largely rewritten, with fuller
treatment of mutation and the maintenance of genetic
Genetics in 1980 and subsequently when he continued variation; we hope these additions will make the book
research and writing but ceased active experimental more useful to students of evolutionary quantitative ge-
work with the mouse. netics. In the earlier chapters, the treatment of polymor-
 Perspectives 1533

phism has been expanded, and some sections in the chap- first sex-linked gene in the mouse, Tabby (Falconer
ters on inbreeding have been shortened (p. ix). 1952a).
The total length of the book was increased by little more The impact of Douglas’s work can best be appreciated
than that of the additional chapter on QTL, which, by considering what was understood about the genetics
together with chapter 20, was written mainly by T.F.C.M. of quantitative traits in the late 1940s and early 1950s.
There were many unanswered theoretical and empirical
questions to be addressed. How effective is artificial
PROBLEMS ON QUANTITATIVE GENETICS selection in changing the mean of a trait? How long
The first and second editions did not include sets of does response to selection continue? How closely do
problems, but Falconer knew that these would improve observed responses to selection match theoretical pre-
the utility of the book for students. He had already dictions? What deductions about the nature of genetic
prepared a large number as part of his teaching, but variation can be made from results of selection experi-
set out to provide a more complete set using, as far as ments? How important is recombination and linkage in
possible, real rather than synthetic data. These were first patterning natural variation for quantitative traits and
published separately as Problems on Quantitative Genetics governing response to selection? The latter questions
(Falconer 1983), but then incorporated into the third arise from the puzzle of abundant genetic variation for
and fourth editions of Introduction to Quantitative Genet- quantitative traits, yet relatively stable mean values in
ics. The problems vary in difficulty and do not necessar- most populations. Could this be explained by the action
ily, as is the case for most real data, have simple right of the population genetic processes of mutation, natural
or wrong answers, as the aim was to get students to selection, migration, and genetic drift on genes affect-
think. Here is one in that category from chapter 10, ing quantitative traits? Or are genes affecting such traits
“Heritability,” raising the sort of problem frequently qualitatively different from those affecting Mendelian
encountered in analysis of metric data (the interested variation? The latter hypothesis was espoused by
reader can consult the book for Falconer’s answer; p. Mather (1941, 1949), who proposed that natural varia-
183): tion for quantitative traits was caused by multiple poly-
genes with individually undetectable but similar and
A study of morphological variation in a population of supplementary, largely additive, effects, which are orga-
Geospiza fortis, one of Darwin’s finches in the Galapagos,
provides the following data on the depth of the bill. How nized in balanced polygenic systems of alleles increasing
would you interpret these data? and decreasing the trait value (i.e., genes in linkage
disequilibrium).
Regressions, ⫾ s.e. Strong predictions regarding the response of quanti-
Offspring-midparent 0.82 ⫾ 0.15 tative traits to artificial selection arise from extrapolating
Offspring-father 0.47 ⫾ 0.17
Offspring-mother 0.48 ⫾ 0.13 population genetic models of response of single genes
Correlations, ⫾ s.e. to selection. Assuming that many genes affect the trait
Full sibs 0.71 ⫾ 0.12 and that allele frequencies are not correlated with the
Father-mother 0.33 magnitude of their effects, a symmetrical response to
divergent selection for increasing and decreasing values
Data are from Boag, P. T. & Grant, P. R. (1978) Nature,
274, 793–4. of the trait is expected, the rate of which should gradu-
ally decline as the frequencies of genes affecting the
While we were delighted to have the problems avail- trait in each direction approach fixation. Ultimately,
able, some of us teaching the course in Edinburgh limits to selection will be reached at which all favorable
would rather that he had not published his answers, as alleles are fixed and no genetic variation remains. Doug-
we had to create another batch of problems to test the las’s results of artificial selection on mice and those of
students. his colleagues with Drosophila (e.g., Robertson and
Reeve 1952; Clayton et al. 1957; Clayton and Robert-
son 1957) rather surprisingly revealed that these predic-
RESEARCH ACHIEVEMENTS
tions generally did not hold well in practice!
Falconer’s major contributions in quantitative genet- For example, after 30 generations of selection for
ics were on the response to artificial selection in mice, increased and 24 generations of selection for decreased
the concept of the cross-environment genetic correla- 6-week body weight in mice, Douglas observed that the
tion, and, as noted above, development of the theory absolute magnitude of response appeared equal in both
for understanding the genetics of complex human dis- directions (Falconer 1955). (This work was reported
eases in terms of an underlying continuous liability. This at the 1955 Cold Spring Harbor Symposium, a major
research also provided much illustrative material for the event in the history of quantitative genetics. Figure 2.)
book. Particularly in his early years of mouse genetic However, Douglas proposed that one should describe
research, Falconer also worked on identification and the selection response in a manner that takes account
mapping of individual genes and notably identified the of the amount of selection applied. He invented the
1534 W. G. Hill and T. F. C. Mackay

course of the experiment; indeed, the phenotypic vari-

ance of the small line actually increased. Douglas also
proposed that the relationship of body size to fitness
could at least be partially assessed by comparing the
expected and realized selection differentials. These were
nearly equal in the large line, but the realized selection
differentials were much lower than the expected differ-
entials in the small line as selection proceeded, indicat-
ing that natural selection was countering artificial selec-
tion for reduced body size.
To assess whether patterns of response to artificial
selection differ between traits, Douglas initiated long-
term divergent selection for litter size (also an important
character in farm animals). An immediate and unex-
pected result was that the response was opposite to the
direction of selection for the first two generations, al-
though continued selection yielded an average realized
h2 of 0.17 (Falconer 1955). This anomalous result was
attributed to a negative nongenetic maternal effect
whereby females reared in large litters were smaller and
Douglas Falconer (left) with Mel Green at the Cold Spring
Harbor Symposium in 1955 (photograph courtesy of Cold consequently had smaller litters. Again, the selection
Spring Harbor Laboratory). response was asymmetrical, with realized h2 of 0.08 in
the high line and 0.23 in the low line (Falconer 1963).
In this case, Douglas demonstrated quite elegantly that
concept of “realized heritability” (h2), obtained by re- the asymmetry was because the “trait” selected—“litter
gressing the cumulated selection response on the cumu- size”—is actually a composite of ovulation rate and em-
lated selection differential, the latter weighted by the bryonic survival rate. Increased litter size was due to an
number of progeny measured. (Douglas credited the increase in the ovulation rate. Ovulation rate was not
origin of this concept to his colleague Barnet “Woggy” changed in the line selected for decreased litter size; a
Wolfe.) When described in this manner, it was apparent decrease in embryo survival accounted for this response.
that the resulting realized heritabilities were markedly An explanation that fits these observations is that genes
different in the high (h2 ⫽ 0.18) and the low (h2 ⫽ 0.52) affecting embryonic survival would be deleterious reces-
lines. This important result indicates that predictions sives and rare in the initial population, and hence selec-
of response to selection from heritabilities estimated tion for reduced litter size would increase their fre-
from correlations among relatives in the base popula- quency and thereby yield a greater response than would
tion could be misleading. Further, asymmetrical re- reducing the frequency further in the high line. Segre-
sponses to selection de facto imply that one or more of gation of rare recessive alleles affecting embryo survival
the assumptions underlying the simple prediction must could lead to a limit to selection for high litter size at
be false. For example, alleles increasing size may have which some genetic variance remains. If so, inbreeding
been more frequent than those decreasing size in the with continued selection, followed by crossing the newly
base population; there might be directional dominance, derived inbred lines, could break the selection limit by
and inbreeding depression could accelerate response purging some of the deleterious alleles. This was exactly
in one direction and hinder it in the other. Douglas what was observed when this experiment was conducted
had the biological insight to realize that in this case the (Falconer 1971).
asymmetry was most likely attributable to a maternal Douglas also made a major contribution to the practi-
effect. He speculated that 6-week body weight could cal problem of deciding in what environment artificial
be partitioned into weaning weight at 3 weeks, largely selection should be applied, if the selected individuals
determined by the mother, and growth between 3 and are to be reared in a wide range of environments. Should
6 weeks, largely a property of the individual. Remark- selection be conducted in a good environment, giving
ably, the asymmetrical response was due to weaning maximal expression to the desired character, as Ham-
weight alone. For some reason, selection for reduced mond (1947) had argued, or should it be carried out
body size was accompanied by a correlated response in under the conditions in which the organisms will eventu-
decreased mothering ability, but there was not a concomi- ally live?
tant increase in mothering ability in the lines selected for Douglas showed that the answer depends on the ex-
increased weight. In addition to the unexpected asymmet- tent to which the trait exhibits genotype by environment
rical responses, neither realized heritability nor pheno- interaction (G ⫻ E). If the rank order and relative mag-
typic variance tended to decline as predicted over the nitudes of phenotypic expression for genotypes affect-
 Perspectives 1535

ing the trait are the same across a range of environ- in the adult animals could be detected. Falconer and
ments, then there is no G ⫻ E and it does not matter colleagues found that body weight was linearly related
in which environment the selection is conducted. How- to the mean cell proportions, which accounted for most
ever, if the expression of the trait changes rank or mag- or all of the chimeric variance of body weight, and that
nitude among the different genotypes, there is G ⫻ E no single organ was found to have a predominant effect
and it might be best to select in the environment in on growth. Hence they concluded that control of growth
which the organisms will ultimately be reared. To ana- must be systemic and not under the control of any single
lyze this, Douglas made a major contribution to quanti- organ or tissue. It will be interesting to see how these
tative genetic theory by showing that the magnitude of results come to be explained in the modern era of
G ⫻ E could be quantified by the cross-environment functional genomics.
genetic correlation, rGE, in which the same character
measured in two environments is considered to be two
LOOKING BACK
different characters. The magnitude of G ⫻ E declines
as rGE approaches unity and increases as rGE approaches One or more editions of Introduction to Quantitative
zero. Thus the answer to the question regarding the Genetics have been translated into at least nine lan-
appropriate environment in which to select comes from guages, sometimes eccentrically. For example, Carlos
evaluating the relative magnitude of the correlated to Lopéz-Fanjul explained to us that in the first Spanish
the direct response to selection (Falconer 1952b). translation by F. M. Sanchez (Introducción a la Genética
Here the correlated response (CR Y) is the response of Cuantitativa, CECSA, Mexico, 1970), Drosophila “bristle
the trait in the environment (Y) in which it is expected number” was translated as “número de setas.” While in
to perform, given selection in a different environment archaic Spanish “seta” means “hair,” in current Spanish
(X), while the direct response (R Y) is for selection in lexicon “número de setas” translates to “number of
the environment in which the organisms will ultimately mushrooms.”
be reared. Assuming equal selection intensities in the What is remarkable is that Introduction to Quantitative
two environments, CR Y ⬎ R Y if rGEhX ⬎ h Y, where h X and Genetics has lasted over 40 years, with only evolution and
h Y are, respectively, the square roots of the heritabili- not revolution of content, and is still used for courses.
ties of the trait in the environment in which selection There have been many predictions of the death of quan-
is made and in the environment in which the individuals titative genetics as a subject, as new techniques have
are expected to ultimately perform. If the genetic corre- been introduced. This has not happened. When the
lation is low, selection should be conducted in the envi- book first appeared, the most obvious market was for
ronment in which the strain is expected to perform, students and researchers in animal and, to a lesser ex-
as was demonstrated by Douglas’s classic experiment tent, plant breeding. Subsequently, as the numbers in-
describing direct and correlated responses of growth volved in such industries have declined, there has been
weight of mice reared on high and low “planes” of nutri- a heightened interest in quantitative genetics in those
tion (Falconer 1960b). studying natural populations and in conservation genet-
We outline only some highlights of Douglas’s subse- ics. The complex nature of the inheritance of much
quent research, which continued to 1990 when he came human disease has been realized, such that quantitative
back to theoretical issues of selection in different envi- genetics of humans is now a fashionable and heavily
ronments (Falconer 1990) and which included the researched subject.
demonstration of genetic variability in susceptibility to The expansion of the subject is illustrated by Genetics
tumors obtained by selection of mice for the number and Analysis of Quantitative Traits by Lynch and Walsh
induced by urethane exposure (Falconer and Bloom (1998), which is over twice the length of Falconer’s
1964). In his later selection experiments Douglas sought Introduction to Quantitative Genetics and is purportedly
to find out more about the genetic basis of the responses only the first of two volumes. (We look forward to the
to selection obtained. First he appreciated the need to second, but have stopped holding our breath.) Other
replicate experiments because one selection line is just books deal with particular parts and their titles illustrate
a single sample, subject to sampling by genetic drift, the breadth of the subject, for each relies heavily on
and so he initiated selection for high and low 6-week quantitative genetic methods, including Roff’s (1997)
body weight with six each of high-selected, low-selected, Evolutionary Quantitative Genetics, Frankham et al.’s
or unselected controls (Falconer 1973). He then (2002) Introduction to Conservation Genetics, and several
sought to find what had contributed to the genetic books covering aspects of animal and plant breeding.
changes. Perhaps his most elegant experiment was to That of Kearsey and Pooni (1996) focuses more on
make aggregation chimeras of embryos from high, con- the approach of the Birmingham school, but brings the
trol, and low lines (Falconer et al. 1981). Such chimeras notation and coverage of Mather and Jinks’ (1971)
vary in the contribution that they get from each “par- Biometrical Genetics closer to that of texts such as Falcon-
ent,” both overall and between organs and tissues. The er’s. Aspects of the theory of quantitative genetics have
lines were genetically marked so that the origin of tissues been covered by Crow and Kimura (1970) in An Intro-
1536 W. G. Hill and T. F. C. Mackay

duction to Population Genetics Theory, by Bulmer (1980) Longmans Green/John Wiley & Sons, Harlow, Essex, UK/New
York.
in The Mathematical Theory of Quantitative Genetics, and Falconer, D. S., 1990 Selection in different environments: effects
recently by Bürger (2000) in The Mathematical Theory on environmental sensitivity (reaction norm) and on mean per-
of Selection, Recombination, and Mutation. Modern meth- formance. Genet. Res. 56: 57–70.
Falconer, D. S., 1993 Quantitative genetics in Edinburgh: 1947–
ods of statistical analysis of quantitative genetic data are 1980. Genetics 133: 137–142.
presented by Sorensen and Gianola (2002). Falconer, D. S., and J. L. Bloom, 1964 Changes in susceptibility
Undoubtedly Douglas Falconer has had an enormous to urethane-induced lung tumours produced by selective breed-
ing in mice. Brit. J. Cancer 18: 322–332.
and lasting impact on quantitative genetics; indeed, he Falconer, D. S., and T. F. C. Mackay, 1996 Introduction to Quantita-
essentially defined the subject. He will be greatly missed. tive Genetics, Ed 4. Longmans Green, Harlow, Essex, UK.
Falconer, D. S., I. K. Gauld, R. C. Roberts and D. A. Williams,
1981 The control of body size in mouse chimaeras. Genet. Res.
38: 25–46.
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Bürger, R., 2000 The Mathematical Theory of Selection, Recombination, to Conservation Genetics. Cambridge University Press, Cambridge,
and Mutation. John Wiley & Sons, New York. UK/London/New York.
Clayton, G. A., and A. Robertson, 1957 An experimental check Hammond, J., 1947 Animal breeding in relation to nutrition and
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Crow, J. F., and M. Kimura, 1970 An Introduction to Population Genet- Lerner, I. M., 1950 Population Genetics and Animal Improvement. Cam-
ics Theory. Harper & Row, New York. bridge University Press, Cambridge, UK/London/New York.
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