Sewall Wright

Lived: December 16, 1889 - March 3, 1988

Population genetics
Visualization of a fitness landscape. The X
and Y axes represent continuous
phenotypic traits, and the height at each
point represents the corresponding
organism's fitness. The arrows represent
various mutational paths that the
population could follow while evolving on
the fitness landscape.
His papers on inbreeding, mating systems,
and genetic drift make him a principal
founder of theoretical population genetics,
along with R. A. Fisherand J. B. S. Haldane.
Their theoretical work is the origin of
the modern evolutionary synthesis or
neodarwinian synthesis. Wright was the
inventor/discoverer of the inbreeding
coefficient and F-statistics, standard tools
in population genetics. He was the chief
developer of the mathematical theory
of genetic drift, which is sometimes known
as the Sewall Wright effect, cumulative

stochastic changes in gene

frequencies that arise from random births,
deaths, and Mendelian segregations in
reproduction. In this work he also
introduced the concept of effective
population size. Wright was convinced that
the interaction of genetic drift and the
other evolutionary forces was important in
the process of adaptation. He described
the relationship between genotype or
phenotype and fitness as fitness
surfaces or evolutionary landscapes. On
these landscapes mean population fitness
was the height, plotted against horizontal
axes representing the allele frequencies or
the average phenotypes of the
population. Natural selection would lead to
a population climbing the nearest peak,
while genetic drift would cause random
wandering.
Evolutionary theory[edit]
Wright's explanation for stasis was that
organisms come to occupy adaptive peaks.

In order to evolve to another, higher peak,

the species would first have to pass
through a valley of maladaptive
intermediate stages. This could happen
by genetic drift if the population is small
enough. If a species was divided into small
populations, some could find higher peaks.
If there was some gene flow between the
populations, these adaptations could
spread to the rest of the species. This was
Wright's shifting balance theory of
evolution. There has been much skepticism
among evolutionary biologists as to
whether these rather delicate conditions
hold often in natural populations. Wright
had a long standing and bitter debate
about this with R. A. Fisher, who felt that
most populations in nature were too large
for these effects of genetic drift to be
important.
Path analysis[edit]
Wright's statistical method of path
analysis, which he invented in 1921 and

which was one of the first methods using

agraphical model, is still widely used in
social science. He was a hugely influential
reviewer of manuscripts, as one of the
most frequent reviewers for Genetics. Such
was his reputation that he was often
credited with reviews that he did not write.
Plant and animal breeding[edit]
Wright strongly influenced Jay Lush, who
was the most influential figure in
introducing quantitative
genetics into animaland plant breeding.
From 1915 to 1925 Wright was employed
by the Animal Husbandry Division of the
U.S. Bureau of Animal Husbandry. His main
project was to investigate the inbreeding
that had occurred in the artificial selection
that resulted in the leading breeds of
livestock used in American beef
production. He also performed experiments
with 80,000 guinea pigs in the study of
physiological genetics. Further more he
analyzed characters of some 40,000 guinea

pigs in 23 strains of brother-sister matings

against a random-bred stock. (Wright
1922a-c). The concentrated study of these
two groups of mammals eventually led to
the Shifting Balance Theory and the
concept of "surfaces of selective value" in
1932. (Wright 1988 Pg 122 American
Naturalist)
He did major work on the genetics
of guinea pigs, and many of his students
became influential in the development of
mammalian genetics. He appreciated as
early as 1917 that genes acted by
controlling enzymes. An anecdote about
Wright, disclaimed by Wright himself,
describes a lecture during which Wright
tucked an unruly guinea pig under his
armpit, where he usually held a chalkboard
eraser: according to the anecdote, at the
conclusion of the lecture, Wright absentmindedly began to erase the blackboard
using the guinea pig.
Wright and philosophy[edit]

 Wright was one of the few geneticists of

his time to venture into philosophy. He
found a union of concept in Charles
Hartshorne, who became a lifelong friend
and philosophical collaborator. Wright
endorsed a form of panpsychism. He
believed that the birth of the
consciousness was not due to a mysterious
property of increasing complexity, but
rather an inherent property, therefore
implying these properties were in the most
elementary particles.[11]
Ronald Fisher:
17 February 1890 - 29 July 1962
He worked at Rothamsted Research for 14
years[3] from 1919, where he developed
the analysis of variance(ANOVA) to analyse
its immense data from crop experiments
since the 1840s, and established his
reputation there in the following years as
a biostatistician. He is known as one of the
three principal founders ofpopulation
genetics. He outlined Fisher's principle as

well as the Fisherian runaway and sexy son

hypothesistheories of sexual selection. He
also made important contributions to
statistics, including the maximum
likelihood, fiducial inference, the derivation
of various sampling distributions among
many others.
Anders Hald called him "a genius who
almost single-handedly created the
foundations for modern statistical
science",[4] while Richard Dawkins named
him "the greatest biologist since Darwin.
Not only was he the most original and
constructive of the architects of the neoDarwinian synthesis. Fisher also was the
father of modern statistics and
experimental design. He therefore could be
said to have provided researchers in
biology and medicine with their most
important research tools, as well as with
the modern version of biology's central
theorem."[5] and Geoffrey Miller said of him

"To biologists, he was an architect of the

'modern synthesis' that used mathematical
models to integrate Mendelian genetics
with Darwin's selection theories. To
psychologists, Fisher was the inventor of
various statistical tests that are still
supposed to be used whenever possible in
psychology journals. To farmers, Fisher
was the founder of experimental
agricultural research, saving millions from
starvation through rational crop breeding
programs."[6]
Fisher was elected to the Royal Society in
1929. He was made a Knight Bachelor by
Queen Elizabeth II in 1952 and awarded
the Linnean Society of London Darwin
Wallace Medal in 1958.
In 1950, Maurice Wilkes and David
Wheeler used the Electronic Delay Storage
Automatic Calculator to solve a differential
equation relating to gene frequencies in a
paper by Ronald Fisher.[74] This represents

the first use of a computer for a problem in

the field of biology. The Kent
distribution (also known as the Fisher
Bingham distribution) was named after him
and Christopher Bingham in 1982
while Fisher kernel was named after Fisher
in 1998.[75]
The R. A. Fisher Lectureship is a prize of a
lecture given yearly in North America that was
established in 1963. On April 28, 1998 a minor
planet, 21451 Fisher, was named after him.[76]

Herbert Spencer

27 April 1820 8 December 1903

Spencer first articulated his evolutionary
perspective in his essay, 'Progress: Its Law
and Cause', published in
Chapman's Westminster Review in 1857,
and which later formed the basis of
the First Principles of a New System of
Philosophy (1862). In it he expounded a
theory of evolution which combined

insights from Samuel Taylor Coleridge's

essay 'The Theory of Life' itself derivative
from Friedrich von
Schelling's Naturphilosophie with a
generalisation of von Baer's law of
embryological development. Spencer
posited that all structures in the universe
develop from a simple, undifferentiated,
homogeneity to a complex, differentiated,
heterogeneity, while being accompanied by
a process of greater integration of the
differentiated parts. This evolutionary
process could be found at work, Spencer
believed, throughout the cosmos. It was a
universal law, that was applying to the
stars and the galaxies as much as to
biological organisms, and to human social
organisation as much as to the human
mind. It differed from other scientific laws
only by its greater generality, and the laws
of the special sciences could be shown to
be illustrations of this principle.

 However, as Bertrand Russell stated in a

letter to Beatrice Webb in 1923, this
formulation has problems: 'I don't know
whether [Spencer] was ever made to
realise the implications of the second law
of thermodynamics; if so, he may well be
upset. The law says that everything tends
to uniformity and a dead level, diminishing
(not increasing) heterogeneity'.[16] As an
objection to evolution, this case is still
regularly made by anti-evolutionists but
does not apply to Darwinian approaches. It
is applicable to matter and energy, not
complex social systems.
Spencer's attempt to explain the evolution
of complexity was radically different from
that to be found in Darwin's Origin of
Species which was published two years
later. Spencer is often, quite erroneously,
believed to have merely appropriated and
generalised Darwin's work on natural
selection. But although after reading

Darwin's work he coined the phrase

'survival of the fittest' as his own term for
Darwin's concept,[6] and is often
misrepresented as a thinker who merely
applied the Darwinian theory to society, he
only grudgingly incorporated natural
selection into his preexisting overall
system. The primary mechanism of species
transformation that he recognised
was Lamarckian use-inheritance which
posited that organs are developed or are
diminished by use or disuse and that the
resulting changes may be transmitted to
future generations. Spencer believed that
this evolutionary mechanism was also
necessary to explain 'higher' evolution,
especially the social development of
humanity. Moreover, in contrast to Darwin,
he held that evolution had a direction and
an end-point, the attainment of a final
state of equilibrium. He tried to apply the
theory of biological evolution to sociology.
He proposed that society was the product

of change from lower to higher forms, just

as in the theory of biological evolution, the
lowest forms of life are said to be evolving
into higher forms. Spencer claimed that
man's mind had evolved in the same way
from the simple automatic responses of
lower animals to the process of reasoning
in the thinking man. Spencer believed in
two kinds of knowledge: knowledge gained
by the individual and knowledge gained by
the race. Intuition, or knowledge learned
unconsciously, was the inherited
experience of the race.
Spencer in his book Principles of
Biology (1864), proposed
a pangenesis theory that involved
"physiological units". These hypothetical
hereditary units were similar to
Darwin's gemmules.[17]
Stephen Gould
September 10, 1941 May 20, 2002

 Gould made significant contributions

to evolutionary developmental biology,
[32]

especially in his work Ontogeny and

Phylogeny.[22] In this book he emphasized

the process of heterochrony, which
encompasses two distinct
processes:neoteny and terminal additions.
Neoteny is the process where ontogeny is
slowed down and the organism does not
reach the end of its development. Terminal
addition is the process by which an
organism adds to its development by
speeding and shortening earlier stages in
the developmental process. Gould's
influence in the field of evolutionary
developmental biology continues to be
seen in such areas as the study of
evolution of feathers.[33]
Gould favored the argument that evolution
has no inherent drive towards long-term
"progress". Uncritical commentaries often
portray evolution as a ladder of progress,

leading towards bigger, faster, and smarter

organisms, the assumption being that
evolution is somehow driving organisms to
get more complex and ultimately more like
humankind. Gould argued that evolution's
drive was not towards complexity, but
towards diversification. Because life is
constrained to begin with a simple starting
point (like bacteria), any diversity resulting
from this start, by random walk, will have a
skewed distribution and therefore be
perceived to move in the direction of
higher complexity. But life, Gould argued,
can also easily adapt towards
simplification, as is often the case
withparasites.[43]
In a review of Full House, Richard
Dawkins approved of Gould's general
argument, but suggested that he saw
evidence of a "tendency for lineages to
improve cumulatively their adaptive fit to
their particular way of life, by increasing

the numbers of features which combine

together in adaptive complexes. ... By this
definition, adaptive evolution is not just
incidentally progressive, it is deeply, dyedin-the-wool, indispensably progressive."[44]