The Herd as a Means

Author(s): David L. Hull

Source: PSA: Proceedings of the Biennial Meeting of the Philosophy of Science Association,
Vol. 1980, Volume Two: Symposia and Invited Papers (1980), pp. 73-92
Published by: The University of Chicago Press on behalf of the Philosophy of Science Association
Stable URL: http://www.jstor.org/stable/192587 .
Accessed: 21/07/2014 16:44
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp

.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.

The University of Chicago Press and Philosophy of Science Association are collaborating with JSTOR to
digitize, preserve and extend access to PSA: Proceedings of the Biennial Meeting of the Philosophy of Science
Association.

http://www.jstor.org

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

The Herd as a Means

David L. Hull
University of Wisconsin-Milwaukee

In 1886 Friedrich Nietzsche noted the following basic error in the

philosophies of his day, .". ..to place the goal in the herd and not in
single individuals!
The herd is a means, no more! But now one is attempting to understand the herd as an individual and to ascribe to it a
higher rank than to the individual--profound misunderstanding!!!
Also
to characterize that which makes herdlike, sympathy, as the more valuable side to our nature!'
(Nietzsche 1901, p. 403).
The recent flap over sociobiology has stemmed largely from the fear
that biology is being used to justify a Nietzschean view of human societies, as if the chief good in human relations must be basically
selfish,
as if apparently altruistic
behavior is fundamentally hypocritical.
From Darwin to the present, biologists
have blithely attributed the
presence and persistence of all sorts of traits, including behavioral
traits, to the "good of the species".
Wynne-Edwards (1962) pushed this
view to such an extreme that finally biologists
were roused to inquire
whether the emperor really was wearing any clothes.
Beginning with Williams (1966), a whole series of biologists
have shown exactly how difficult it is for anything to be done for the good of the species if species evolve the way that we think they do. The biological
issues are:
(a) the nature of organization,
(b) the levels of organization which
actually exist in particular sorts of organisms, and (c) the evolutionary processes which can take place at each of these levels.
These are
among the questions with which the following papers by Sober (1981) and
Wimsatt (1981) deal.
The purpose of this paper is to explain the current state of biological
theory on these issues and its implications for
human societies.
I argue that the nature of biological
evolution and
biological
organization have important implications for human societies,
butnot the ones usually claimed.
In the first place, the fundamentals
of evolutionary theory are currently in a state of flux.
Now is not the
time to take a particular interpretation of biological
evolution and
apply it uncritically
to social evolution.
However, our understanding

PSA 1980, Volume 2, np. 73-92

Copyright (
1981 by the Philosophy of Science Association

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

74
of the complexities, problems and possible solutions in biology can
profitably be used to help us increase our understanding of societies
and how they can change. Conversely, there is no reason why our knowledge of social organization cannot be used to help understand biological evolution.
1.

Group Characteristics

and Group Selection

One fundamental question in theoretical biology today is the nature,

in the evolutionary proexistence and role of "group characteristics"
For example, Wynne-Edwardsnotes that in developing his ideas:
cess.
... it soon became apparent that the greatest benefits of sociality
arise from its capacity to override the advantage of the individual
members in the interests of the survival of the group as a whole.
as explained more
The kind of adaptations which make this possible,
fully here, belong to and characterize social groups as entities,
This in turn seems to enrather than their members individually.
tail that natural selection has occurred between social groups as
evolutionary units in their own right, favoring the more efficient
variants among social systems wherever they have appeared, and furthering their progressive development and adaptation. (1963, p. 623).
Wynne-Edwardsis claiming three things in the preceding quotation:
(b) that selection can
(a) that groups themselves can have adaptations,
operate on such groups, and (c) that the good of the group can override
the good of its members. Most discussions of group selection have
If something is a genuine group, then the
dealt with the final claim.
conditions under which it can be selected over and above its individual
If group selection occurs at all, it is hardly
members are very rare.
Much less attention
a major feature of evolution (but see Wade 1978).
The crucial distinction for our
has been paid to the other two claims.
purposes is between properties of single organisms and properties of
A single mammalcan possess mammaryglands.
more inclusive entities.
These glands not only do not aid this organism in its own survival, but
Hence, one might be tempted to explain the
are actually detrimental.
possession of mammaryglands by individual mammals in terms of the good
of some group--the family, the tribe or the species.
However, these are not the traits of greatest interest to Wynne-Edwards. Some properties seem to characterize groups as such and not
For example, most species have a 50-50 sex
their members severally.
may be analyzable entirely in
Although such a characteristic
ratio.
terms of the sex of the members of a species, it is a property of the
As Williams (1966, p. 108) puts
species and not the members severally.
it, the contrast is "between a population of adapted insects and an
Other examples of putative group charadapted population of insects."
are balanced polymorphisms and frequency dependent selecacteristics
These are the sorts of adaptations which Wynne-Edwardsis chieftion.
One element in
ly concerned to explain in terms of group selection.
As Nietzsche
Wynne-Edwards' argument, however, tends to get overlooked.
fears, Wynne-Edwardsis "attempting to understand the herd as an indi-

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

75
vidual."
If too many adaptations "belong to and characterize social
groups as entities,"
these groups cease to be "groups" and become individuals in their own right.
One might be tempted to treat an organism
as a group of cells.
Typically we do not because of the numerous organizational
properties which serve to integrate these cells into a
single system (Hull 1976, 1978).
The same line of reasoning should
apply as readily to entities more inclusive than single organisms.
Organization is what counts.
Thus, two different sorts of group selection must be distinguished:
(i) selection of groups as well-integrated wholes, and (ii) selection
of groups because of extrinsic constraints.
Most discussions concern
the possibility
and relative frequency of the second sort of group selection, the selection of aggregates of organisms which are selected
together merely because they all happen to live in the same pool of
water or on the same host (Wilson 1980).
Much less attention has been
In this
paid to the selection of groups which are really not "groups".
second sense, a group can function as a unit of selection only if it is
The trouble is that
characterized by enough organizational
properties.
such highly organized groups are no longer properly interpreted as
groups, but as individuals.
Anything which is sufficiently well-organized to be selected must be sufficiently well organized to count as an
individual.
Williams (1966) reasons along much the same lines as Wynne-Edwards
He argues that the fitbut comes to somewhat different conclusions.
ness of a group can be treated as a simple summation of the fitnesses
of its constituent organisms.
The organisms are the entities with the
For example, the fleetness of a herd of deer is totally a
adaptations.
function of the fleetness of individual deer in the herd. Only if the
herd were a well-organized whole could it have adaptations of its own.
"Such individual specialization
in a collective
function would justify
recognizing the herd as an adaptively organized entity.
Unlike indivldual fleetness,
such group-related adaptation would require something
more than the natural selection of alternative alleles
as an explanation."
(Williams 1966, p. 17).
One point on which Williams and Wynne-Edwardsdisagree is the actual
status of such things as herds of ungulates and schools of fish.
WTynne-Edwards
thinks they are organized wholes; Williams thinks they
are not. Williams views himself as an "individual selectionist"
because he believes that "adaptation need almost never be recognized at
any level above that of a pair of parents and associated
offspring."
(Williams 1966, p. 19).
Thus, he would agree with Wilson (1971) that
certain sorts of colonies can exhibit adaptations.
For example, the
organisms which comprise a hive in certain eusocial insects exhibit division of labor, functional and structural differentiation,
characteristic distributions
in the hive both spatially at any one time and temporally during the "life cycle" of the hive, and so on. If such hives
are not supraorganismic individuals,
nothing is.
For this reason,

Wilson (1971)

thinks that selection

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

can take place

76
Williams (1966) might agree that such hives are
at the level of hives.
individuals and exhibit adaptations of their own, but he would draw the
This conclusion
line at their functioning as units of selection.
should come as no surprise because Williams does not think that even
"One necessary condiorganisms can function as units of selection.
tion" for an entity to be selected "is that the selected entity must
have a high degree of permanence and a low rate of endogenous change,
relative to the degree of bias (differences in selection coefficients).?
Dawkins (1976, 1978) has made himself extreme(Williams 1966, p. 23).
first, by stripping the emperor of yet
ly unpopular among biologists,
another layer of clothes and, second, by doing so in a popular format.
can be found saying such things
Time and again, population biologists
Although
as "evolution is nothing but changes in gene frequencies.?"
models in population biology need not be limited to the relative freThe claim is,
at a single locus, most are.2
quencies of two alleles
however, that the fitness of an organism can be treated as a simple
Although Dawkins has
summation of the fitnesses of its separate loci.
taken the heat for emphasizing this position, it can be found explicitAccording to Williams:
ly expressed in Williams.
to believe that a gene actually exObviously it is unrealistic
ists in its own world with no complications other than abstract
The unity of the genoselection coefficients and mutation rates.
type and the functional subordination of the individual genes to
each other and to their surroundings would seem, at first sight, to
Actually
invalidate the one-locus model of natural selection.
these considerations do not bear on the basic postulates of the
theory. No matter how functionally dependent a gene may be, and no
matter how complicated its interactions with other genes and environmental factors, it must always be true that a given gene substitution will have an arithmetic mean effect on fitness in any
can always be regarded as having a certain
One allele
population.
selection coefficient relative to another at the same locus at any
given point in time. Such coefficients are numbers that can be
and conclusions inferred for one locus can
treated algebraically,
be iterated over all loci.
Adaptation can thus be attributed to
the effect of selection acting independently at each locus.
(1966,
pp. 56-57).
The preceding conviction is at the heart of the levels of selection
Some of the objections raised
controversy and from it to sociobiology.
to sociobiology have concerned any attempt to explain human social
but others are raised to the attempt to
characteristics
biologically,
model. These critics argue that this
do so on the "gene selectionist"
traits.
It
overly simple model will not do for ordinary biological
surely will prove inadequate for social traits.
Although Williams acknowledges the existence of adaptations at levels more inclusive than single genes and even organisms, he is a gene
because he thinks that all these adaptations can be exselectionist
For
plained entirely in terms of selection acting on particular genes.
example, the adaptations exhibited by hives are to be explained by

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

77
not selection acting on kinship groups.
means of kin selection,
Even
though Williams (1966, pp. 159, 96-97) acknowledges that the "central
biological
problem is not survival as such, but design for survival,"
he maintains that an organic adaptation is a "mechanism designed to
promote the success of an individual organism, as measured by the extent to which it contributes genes to later generations of the population of which it is a member." Once again, if Williams thinks that selection does not act on entities composed of parts which all have the
same genetic makeup--such as organisms, he certainly cannot acknowledge
as units of selection entities composed of parts with
different genetic makeups--such as beehives.
2.

Replication

and Interaction

One minor source of confusion in the group selection controversy is

equivocation over what actually counts as "individual" versus "group"
selection.
Gene selectionists
term themselves "individual
selectionists" because genes are individuals.
Organism selectionists
also feel
that they have the right to term themselves "individual selectionists"
because organisms are also individuals.
Finally, even such classic
group selectionists
as Wynne-Edwardshave some right to be termed "individual selectionists"
because they argue that many so-called "groups"
are really individuals!
A more serious source of confusion has been a
systematic equivocation over two different senses of "selection"
and
"unit of selection."
Williams (1966, p. 25) proposes to redefine "gend'
in evolutionary contexts as "any hereditary information for which there
is a favorable or unfavorable selection bias equal to several or many
times its rate of endogenous change."
Dawkins (1978, p. 67) suggests
replacing the term "gene" in such contexts with the more general term
"replicator",
which he defines as "any entity in the universe which interacts with its world, including other replicators,
in such a way that
copies of itself are made. A corollary of the definition is that at
least some of these copies, in their turn, serve as replicators...
.'"
I find Dawkins' notion of a replicator an important Improvement in
the conceptual foundations of evolutionary theory, but in his definition he runs two sorts of interaction together, the sort of interaction
necessary for a replicator
to replicate
itself and the sort which produces differential
replication.
That these are two different processes
can be seen in the fact that they are usually carried on by different
entities at different levels of organization.
Not only that, but because these functions are so different, the entities which perform them
tend to be characterized by different sorts of general properties.
The
only "adaptations" which a replicator needs are those to promote replication.
All that an entity need be able to do to function as a replicator is to replicate
itself,
the more directly the better.
If all
that goes on is replication,
evolution of sorts might result, but not
evolution through selection.
In order for selection to occur, an additional process is necessary.
Either the replicator itself or else
some more inclusive entity produced by the replicator must interact
with its environment in such a way that replication
is differential.
These latter entities are the entities which have "adaptations"
in the

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

78
usual biological

sense of this term.

Elsewhere I (Hull 1980, p. 318) have termed the entities which

function in this second process "interactors" and defined these two
technical terms as follows:
on its structure directly

in

replicator:

an entity that passes

replication.

interactor:

an entity that directly interacts as a cohesive whole

is
with its environment in such a way that replication
dif ferential.

With the aid of these two technical

self can be defined as follows:
selection:

terms, the selection

process

it-

a process in which the differential extinction and proliferation of interactors cause the differential perpetuation of the replicators which produced them.

In the
replicate themselves.
In selection processes, replicators
probably also functioned as the only
beginning, the first replicators
However, as evolution proceeded, these two funcinteractors as well.
Replicators began to produce evermore intions became differentiated.
clusive interactors to cope with evermore inclusive and complex environments. The result is the part-whole hierarchies which are so charof the living world. Some entities are extremely simple.
acteristic
They
As organisms they are hardly more than encapsulated replicators.
No higher levels of
reproduce themselves asexually and that is that.
Some organisms are themselves highly complex,
organization are present.
systems. Some organisms form colonies
highly stratified hierarchical
Sexual organisms, at least, form
and other sorts of kinship groups.
(Contrary to commonusage, just as not all organisms form
species.
The question now bekinship groups, not all organisms form species.)
comes, for any particular sort of organism, at what level or levels is
taking place, at what level or levels is interaction taking
replication
occurs at the lowest levels of organizaTypically replication
place?
tion, primarily at the level of the hereditary material, while interaction occurs both at these levels and at increasingly more inclusive
The point I wish to stress is that not only are both
levels as well.
but also both are important. Neiprocesses necessary for selection,
ther can be omitted, and neither takes precedence over the other.
With this distinction in mind, one disagreement between gene and orWhengene secan be shown to be only apparent.
ganism selectionists
lectionists
argue that selection occurs only at the level of the genetin mind. Rates of endogenous change
ic material, they have replication
Whenorganism selectionnot interaction.
are relevant to replication,
ists claim that selection occurs primarily at the organismic level,
they have interaction in mind. Genes may be the entities which replicate themselves most directly, but they tend to interact with their
Although organevermore inclusive environments evermore indirectly.

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

79

isms interact directly with their environments, they tend to replicate

themselves only indirectly via their genes.
Regardless of how it might
appear, the distinction between replication
and interaction is not
"merely semantic".
Entities more inclusive than single genes can function as replicators.
They do not thereby become genes.
A general term
like "replicator"
is necessary if confusion is to be avoided.
Similarly, entities both less inclusive and more inclusive than organisms can
If populations can function as interactors,
function as interactors.
then they are interactors, not "superorganisms".
3.

Linkage Disequilibrium

and the Unity of the Genotype

Once the preceding disagreement between the gene and organism selectionists has been shown to be apparent, an important difference nevertheless remains. Organisms are well-integrated cohesive wholes.
That
is why they can function so well as interactors.
Each organism is produced by its genome in interaction with successive environments. An
extremely complex system of feedback loops becomes established between
the developing organism, its environments and the genome which is producing it.
But all of this is relevant only to interaction.
How about
replication?
Biologists
such as Mayr (1963, 1975) have emphasized the
role of the unity of the genotype in evolution.
Although there is a
one-to-one correspondence between genomes and organisms, many organisms
can possess the same genotype, e.g., clones.
Conversely, many different genotypes can produce, for all intents and purposes, the same phenotype, i.e.,
phenocopies.
The issue of the unity of the genotype refers to the role of genotypes themselves as cohesive systems. Genotypes are functionally and structurally organized systems. In selection processes can they be treated as if they were not?
Biologists
can be found arrayed on both sides of this issue.
As the
earlier quotations indicate, Williams (1966) maintains that one-locus
models should be adequate for characterizing the evolutionary process.
"iNo matter how functionally dependent a gene may be, and no matter how
complicated its interactions with other genes and environmental factors,
it must always be true that a given gene substitution will have an
arithmetic mean effect on fitness in any population."
(Williams 1966,
p. 57).
The issue is the extent of linkage disequilibrium.
The coefficient of linkage disequilibrium
is a measure of the statistical
dependence between two loci.
To some extent the term is misleading in
that this dependence need not have anything to do with linkage of loci
on the same chromosome. (Roughgarden 1979, p. 113).
If genotypes are as unitary as Mayr (1963, 1975) claims, one would
expect to discover very high linkage disequilibrium coefficients when
genes in the same functional complex are studied.
So far the evidence
is not as unequivocal as one might wish.
(See Lewontin 1974 and Roughgarden 1979 for reviews.)
No matter how this issue is decided, there
are gains and losses on both sides.
If genomes are highly organized
functional systems, then evolutionary models which take only one or two
loci into account at once are liable to be inadequate.
The recognition

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

80
is purchased at the
of higher levels of organization in replicators
price of complicating studies of evolutionary processes significantly,
If one-locus models are good enough, then the
possibly prohibitively.
is much simpler, but he is left
task of the evolutionary biologists
with explaining why the highly complicated internal organization of
genomes can be ignored.
Both Sober (1981) and Wimsatt (1981) argue against reductionism in
Two sorts of reductionism are involved in the controversy
biology.
over the unity of the genotype. WhenWilliams (1966) argues that a
of organisms, he is claiming that
herd is nothing but a collection
herds are not sufficiently well-organized to be counted entities in
Certainly a herd of ungulates is composed of nothing
their own right.
Any relations which might justify viewing a
but individual ungulates.
herd as a higher-level entity would be relations between these individual organisms. However, when Williams (1966) argues that replication
can be treated as if only the genetic material can function as replicators and that individual genes can be treated as if they were functionally independent of each other, he is presenting an even more reThey
Organisms are not composed just of genes.
ductionist position.
Only a small percentage
are made up of numerous other parts as well.
of an organism's mass consists of DNA. On the gene selectionists'
is concerned, organisms are nothing but
view, as far as replication
of genes.
collections
4.

Genetic Diversity

and the Unity of the Genotype

One of Mayr's main goals in his Systematics and the Origin of Species (1942) was to counter the typological species concept by emphasizing the amount of genetic diversity present in natural populations and
According to the typological species concept, spespecies at large.
by means of fixed sets of essencies are natural kinds characterizable
All members of a particular species must possess all the
tial traits.
essential
traits of its species, and no other species can be characterAll variation, whether at
ized by precisely this same set of traits.
Mayr (1942) showed
any one time or through time, is purely accidental.
that if one follows a species through its range, one discovers considAn allele which is commonin one populaerable geographic variation.
tion is rare in another, and so on. Populations at the termini of
if anything, in common. They may not
these clines may have little,
According to current best estimates,
even be able to mate successfully.
sexually reproducing species of animals are polymorphic for a third of
their genes, and at an average polymorphic locus, a quarter of the in(Roughgarden 1979, p. 87).
dividuals in a species are heterozygous.
This means that at a third of the loci in a species two or more alleles
can be found, and that at these loci, a quarter of the organisms are
If phyletic evolution
possess different alleles.
heterozygous, i.e.,
only
one should expect to discover that this variability
is possible,
increases if one follows a species through time. Other sorts of polymorphism also exist, e.g., trophic polymorphisms (Turner and Grosse
1980).

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

81

Because of the extensive genetic diversity which exists in species,

Mayr (1969, p. 369) argues that the notion of a "typical" memberof a
species makes no sense.
"Species consist of variable populations, and
No single specimen
no single specimen can represent this variability.
sense."
can be typical in the Aristotelian
Mayr, in his second synoptic work on the evolutionary process, Animal Species and Evolution
of species.
As var(1963), emphasizes just the opposite characteristic
iable as species are, each species possesses its own basic genotype:
The basic gene complex of the species (with all the species-specific canalizations
and feedbacks) functions optimally in the area for
which it had evolved by selection,
usually somewhere near the center.
Here it is in balance with the environment and here it can
afford much super-imposed genetic variation and experimentation in
niche invasion.
Toward the periphery this basic genotype of the
species is less and less appropriate and the leeway of genetic variation that it permits is increasingly narrowed down until much
uniformity is reached.
(Mayr 1963, p. 527).
Because the "unity of the genotype places well-defined limits on the
potential for variation"
(Mayr 1963, p. 176), a "genetic revolution" is
usually needed for new species to arise.
Instead of speciation occurring through numerous generations by the gradual accumulation of minor
changes in gene frequencies, it usually occurs by means of the isolation of a small population at the periphery of the species.
Most such
peripheral isolates
go extinct, but every once in a while, one of them
becomes established as a new species with its own characteristic
genotype.
(See also Eldredge and Gould (1972).)
Three issues are involved here:
(a) the unity of particular genotypes as discussed earlier,
(b) the prevalence of a single basic genotype throughout a species in spite of considerable genetic heterogeneity, and (c) the role of the genotype in promoting the cohesion of the
gene pool.
Mayr is concerned to argue that neither genomes nor species
are aggregates.
Both are organized wholes.
One explanation for the
cohesiveness of the gene pool is that all organisms belonging to the
same gene pool have basically
the same genetic makeup. However, this
is not the only explanation for or mechanism which can serve to enhance
the cohesiveness of the gene pool.
Mayr (1963, p. 542) remarks that
throughout his book, he has "stressed the tremendously cohesive effect
of gene flow. Yet, when one tries to calculate
the time it takes for
genes to percolate from one end of the range of a widespread species to
the other, one arrives at rather astronomical figures."
He goes on,
however, "Without wanting to depreciate the importance of gene flow, I
advance the thesis that the cohesion of the species is also due to the
fact that all those of its populations that have not undergone a genetic revolution share the same homeostatic systems and that these systems
give great stability."
(See also Eldredge and Gould (1972) .)
Thus, a
species is cohesive, both because gene flow promotes cohesiveness and
because all organisms in the species possess the same basic genotype.
The problem is how to reconcile

the unity of the genotype with the

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

82
One possisimultaneous existence of extensive genetic heterogeneity.
At
is limited to certain loci.
ble solution is that this variability
All organisms within a single speexists.
other loci, no variability
The question
the same genes at these loci.
cies possess identically
then arises if these are also the genes which distinguish this species
If so, then species have an essence in an Arfrom all other species.
Or it may be the case that a species shares its consense.
istotelian
stant loci with several other species and is distinguished from them
If so, then species lack
only by differences at its variable loci.
Or it may be the case that
sense.
essential genes in the Aristotelian
the structure of the genotype supplies its unity. Although a variety
can exist at any one locus, the overall arrangeof different alleles
ment of loci remains the same. This structure is then its "essence."
species seem capable of remaining homeostatic sysSomehowbiological
Furthermore, if all
tems in spite of considerable internal diversity.
the organisms which belong to the same species possess, in some sense,
in using this comthe same genotype, then there is some justification
If so, then asexual organisms
monality in defining particular species.
form species as surely as do sexual organisms.3
As Dawkins (1979) documents, the recent revolution in our thinking
about the evolutionary process has its origin in W. D. Hamilton's 1964
In these papers Hamilton
papers in the Journal of Theoretical Biolog.
introduces his notion of inclusive fitness--the contribution which an
individual makes to the gene pool of the next generation, both directly
of its own genes and indirectly via duplicates in relatvia replicates
It is in this paper that the mathematics or altruistic
ed organisms.
behavior receives its first extensive treatment. Hamilton concludes
bethe first of these papers by stating that, in order for altruistic
havior to evolve:
...the benefit to a sib must average at least twice the loss to the
the benefit to a half-sib must be at least four times
individual,
the loss, to a cousin eight times and so on. To express the matter
more vividly, in the world of our model organisms, whose behavior
is determined strictly by genotype, we expect to find that no one
is prepared to sacrifice his life for a single person but that evit when he can thereby save more than two
eryone will sacrifice
. (1964,
brothers, or four half-brothers, or eight first cousins...
p. 16).
erAs Nisbett and Ross (1980, p. 45) note, one of the most coammon
included, is to be
rors in reasoning made by human beings, scientists
This is certainly the
unduly affected by the vividness of information.
becase with Hamilton's discussion of inclusive fitness and altruistic
In most cases, it is his vivid description of the consequences
havior.
Although
of his argument which gets quoted (e.g., Wilson 1975, p. 415).
too ofmade in the technical literature,
the necessary distinctionsare
Time and aten they tend to be neglected in more popular expositions.
gain we are told that the investment which one organism makes to another should covary with the number of genes the two have in common. Energy flow should go with gene flow. For example, in the commonestform

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

83
of inheritance, siblings should share half of their genes, grandchildren a quarter, greatgrandchildren an eighth, and so on. Similar calculations can be made for nieces and nephews, cousin, etc.
The trouble with this way of putting it is that it conflicts with
the actual facts of the case.
Even though species are genetically
quite heterogeneous, most alleles
are present in very low frequencies.
Hence, any two organisms picked at random are likely to possess exactly
the same alleles
at the vast majority of their loci, say, 90%. The 507,
those loci at which the two organ25%, etc. progression concerns o
isms happen to differ.
Thus, any two organisms picked at random are
liable to have the same alleles
at 90% of their loci, while siblings
under these same circumstances should share 95% of their genes.
As a
result, although nepotism should exist, it should not be as difficult
to overcome as one might expect.
A second issue in which the preceding difficulty arises is the cost
of meiosis.
In many cases, organisms are "outbreeders".
They tend to
mate with organisms that are not especially
closely related to them.
They are members of the same species, but that is all.
In most sorts
At meiosis,
of sexual reproduction, each organism loses half its genes.
reduction division occurs.
Thus, sexual reproduction has a 50% cost.
If the name of the evolutionary game is to pass on one's genes, sexual
reproduction must be extremely advantageous since it has to make up a
50% loss.
Once again, the fact that most organisms have exactly the same alleles at the vast majority of their loci is being overlooked.
Two organisms which mate are likely to differ at, say, 10% of their loci.
Hence,
the cost of meiosis is reduced to half of the 10%, or 5%. Although a
5% cost is not negligible,
it does not pose quite the problem that 50%
does.
Barash (1976) raised precisely this objection to the supposed
50% cost of meiosis, only to have it dismissed curtly by Maynard Smith
and Williams (1976) as a total misunderstanding of the problem. A misunderstanding it surely is, but a misunderstanding which is perfectly
understandable given the early literature on the subject.
The relevant distinction
is between the percentage of genes which
two organisms share and the likelihood that a particular gene will be
passed on. In most sorts of sexual reproduction, a particular offspring will always get 50% of each genes from one parent and 50% from
the other.
Similarly, siblings will share 50% of their genes.
In the
case of the next generation, the 25% figure is an average.
On the average a grandchild will receive 25% of its genes from each grandparent,
although it is possible for it to receive no genes from one grandparent
(maternal or paternal) and 50% from the other.
In all cases these are
genes which are identical by descent.
The second set of figures refers
to likelihoods
of transmission.
Given a particular gene in a parent,
what is the likelihood
that it will be passed on to a particular of fspring and not its allele?
Given a particular autosomal gene in an
offspring, what is the likelihood
that this gene was obtained from one
parent rather than the other? The answer in both cases is 50%. Be-

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

84
cause the numbers come out the same, it is easy to overlook the fact
that we are dealing with percentages, average percentages, and likelihoods.
A second important distinction is between genes with exactly the
same structure and genes which have the same structure because they are
immediate replicates
of the same ancestral gene. The latter are genes
identical by descent.
Whenwe say that each offspring receives 50% of
its genes from each of its parents, we are referring to genes as material entities.
Nothing is implied necessarily about similarity in
In all natural populations, more than 50% of the offsprings'
structure.
genes will be similar in structure.
How many depends on the percentage
of loci at which its parents possess alleles
with the same structure.
Although these alleles
will also be identical by descent, their common
If
ancestral gene may have existed numerous generations in the past.
one counts only those genes which are similar because of iimmediate descent, 50% of the genes of each offspring will be identical by descent
to the genes of each of its parent.
However, if more distant descent
is allowed, this figure begins to approach 100% as the percentage of
loci at which the parents have alleles
with the same structure approaches 100%.
As Stampe and Metcalf (1980, p. 613) point out, "Disagreement between predictions of several theories can be traced to differences in
the interpretation of the meaning of coefficient of relationship
(r).
Genetic models suggest that r is best defined as the probability that
a certain gene is shared with a relative through commondescent, rather than as the proportion of genes shared between relatives
through
commondescent."
Both kin selection and the cost of meiosis are best
expressed with r defined in the first way. The fact that many authors
define it in the second way (e.g., Wilson 1975) explains why controversies on these issues are so common. When the criterion of identity
through descent is ignored and all genes with the same structure are
If one
considered genes of the same sort, confusion is only increased.
is not concerned with selection processes, genes can be considered to
belong to the same natural kind solely on the basis of structural identity or similarity, just as organisms can be considered to belong to
the same species if they have sufficiently similar genomes (Caplan
1980).
However, if genes are the things which are being selected and
species are the things which are evolving, descent takes priority to
similarity.
Only entities which are identical
(or similar) by descent
belong to the same reference class (Hull 1976, 1978, 1980).
6.

Implications

for Human Societies

Present-day human societies

are biologically
peculiar in several retended to be
spects.
Throughout most of our history, human societies
of the tribal sort--50 to 150 individuals,
most of whomwere relatives.
Present-day societies
are much larger and much more heterogeneous in
all respects, including genetic heterogeneity.
Tribal wars in which
one tribe successively
annihilated its neighboring tribes could affect
Wars between
the genetic makeup of the human gene pool in that area.

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

85
of present-day nation states are not as likely to have such
alliances
an effect.
Although it presents precious little
comfort, the effects
of atomic warfare are liable to be genetically indiscriminate, having
as random an effect on human evolution as mutations.
The major problem posed by the size and genetic heterogeneity of
human societies,
however, concerns the efficacy of possible biological
mechanisms for promoting cohesiveness of these societies.
If one looks
just at the number of loci at which any two people picked at random
from a society are likely to have the same alleles,
one should expect
people to cooperate quite extensively.
Although human societies
are
genetically quite heterogeneous, any two members of a society are likely to be genetically quite similar.
The problems which have been
raised by sociobiologists
to cooperation among human beings arise only
if one concentrates on alternative alleles
If loci
at a single locus.
are selectively
independent of one another, then genes at different loci neither cooperate nor compete with each other, only different alleles at the same locus.
In this very particularized
context, it is
difficult to see how an "altruistic
allele"
could come to replace a
"selfish allele".
However, the entities referred to here are different
not different genes, and certainly not different organisms.
alleles,
The inferences from selfish alleles
to selfish genes, and from selfish
genes to selfish organisms are extremely problematic.
They may be justified, evolutionary biologists
are certainly warranted in continued
attempts to justify them, but they currently are less than crystal
clear.
After all, it should be remembered that, on exactly this same
line of reasoning, sexual reproduction should be rare, and according to
most workers, it is extremely prevalent.
(But see Hull 1980).
If we limit ourselves to the single locus interpretation,
it is hard
to see how behaviors contributing to social cohesiveness in human beings can have much of a genetic basis.
At most, such behavior can be
"misfirings" of previously adaptive behavior which has yet to be eliminated
(Dawkins 1976, p. 109).
For example, parental investment is
genetically quite advantageous as long as it is directed at biological
offspring.
Adoption of unrelated offspring is not. The desire by human beings to adopt children can be explained either as such a misfiring or as an extremely cynical form of exploitation.
Argumentsagainst
the efficacy of the simple application of kin selection models to human
beings cannot rest solely on unusually high rates of adoptions in certain societies,
such as Eskimos. It must also be shown that adopted
children are not by and large turned into reproductive neuters.
The
pleasing side of this same coin is that the effects of genes being identical by descent drop off very rapidly as genealogical
relationship
becomes more distant.
Either very low differences in inclusive fitness
can make a difference or else distant relatives should treat each other
no differently from how they treat non-relatives
(West Eberhard 1975).
However, it should be kept in mind that considerable disagreement
exists among biologists
about the precise nature of the strictly biological mechanisms which promote the cohesiveness of gene pools as such,
and that some biologists
doubt the very existence of such cohesiveness.

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

86
One should be both very careful and very tentative in reasoning from
such problematic biological
I do think, however, that exprinciples.
amining such issues as the unity of the genotype and the cohesiveness
of gene pools can be helpful in understanding human societies.
By
this I do not mean that one should reason from the nature of the human
genotype and structure of the human gene pool to the nature of human
societies,
although such inferences might well be warranted. Rather,
I think that all three might profitably be viewed as instances of the
same sort of phenomenon--highly organized systems which, nevertheless,
can undergo change.
How much can a genome be modified and still remain functional?
Can
a genome be modified successively
in time, one or two loci at a time,
until it forms a distinctly new genotype; or are genetic revolutions
necessary?
Currently the answers to these questions remain elusive,
but surely biologists
have investigated them in greater detail and have
discovered a greater variety of problems and suggested solutions for
genotypes and gene pools than sociologists
have for societies.
Human
societies,
like gene pools, are quite heterogeneous.
They are also cohesive, so it seems. How can such cohesiveness be maintained in the
face of such heterogeneity?
Calls for human freedom if answered produce more heterogeneous societies.
How "free" can the people in a society become without the society ceasing to be a society?
Too much homogeneity also has its costs.
If the analogy is appropriate, one is
in claiming that neither heterogeneity nor homogeneity is an
justified
unalloyed good. It all depends on the intensity and nature of the selection pressures.
Permitting conscientious objection in peacetime or
during limited wars might well be a beneficial escape valve.
During
all-out wars, it might prove detrimental.
Societies appear to be entities in their own right, with their o.m
From a biological
characteristics.
point of view, it is difficult to
treat societies
in this wray. Organisms, kinship groups and human societies differ from each other in being increasingly genetically heterogeneous.
The more heterogeneous they are, the less likely they are to
be able to function as replicators.
From this it does not follow that
they cannot function as interactors.
Just as cells do not interact
with their environments in isolation,
people do not interact with their
environments in isolation.
Human societies
pose problems for a purely
biological
theory of evolution, not because of any peculiarly human
but because of their strictly biological
characteristics
characteristics.
It is important to distinguish those problems which arise from
human societies
being systems from those which arise because of any peof human social systems. Too often critics of the "bioloculiarities
gizing" of the social sciences leap over substantial biological
problems to dwell exclusively on the sociological
problems.
No one mentions social evolution without emphasizing its partial independence of biological
evolution.
Culture flow does not always coincide with gene flow. In human beings, cross-lineage
borrowing is possible (Campbell 1972, p. 33).
One can teach one's own offspring, but
one can also teach the offspring of others.
The ease with which

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

87
human beings can teach non-relatives must, on the gene selectionist
model, be interpreted either as another "misfiring" or as an instance
Whenever one teaches
of "reciprocal altruism" (Trivers 1971, p. 46).
the offspring of others, social and biological
or "indoctrinates"
evolution can come into conflict.
Teaching one's own offspring the virtue
teaching
of patriotic self-sacrifice
is genetically quite altruistic;
others such a virtue is genetically selfish.
Organisms which must cooperate with their sexual competitors should be ambivalent in their
the ability to indoctrinate arelationships.
Add to this situation,
As Campbell (1972,
cross lineages, and the ambivalence only increases.
in human beings is a
p. 23) emphasizes, a self-sacrificial
disposition
"product of social indoctrination,
which is counter to rather than supported by genetically transmitted behavioral dispositions."
A second point which needs emphasizing is that currently we have no
detailed, well-developed theory of social evolution (Alexander 1979,
It is all well and good to mention possible differences
Blute 1979).
between biological
and social evolution, but until someone actually
produces a theory of social evolution comparable to current theories of
biological
evolution, such discussions must remain highly tentative.
of a
Recall all the really excellent arguments against the possibility
do what
genetic code (Commoner1961).
Time and again the Philistines
Humansocieties
their intellectual
may
superiors know is impossible.
represent just another level of organization, presenting no new problems, or it may represent an insurmountable barrier to the literal
extension of a strictly biological
theory of evolution.
Conceptual evolution represents yet another level in the levels of
selection controversy.
In order for biological
theories of evolution
to be adequate for conceptual evolution, scientific
ideas would have to
be transmitted by the genetic material.
It is plausible
that certain
general features of human societies
are to some extent influenced by
our genes.
It is also plausible
that the curiosity so necessary for
science is in human beings genetically based.
But it is very unlikely
that calculus or quantum mechanics is in any sense "programmedinto our
genes".
However, one feature of this controversy which I find curious
is that the very same scientists
who argue for biological
influences on
social evolution draw back at a parallel argument one level up--social
For example, Wilson (1975) argues
influences on conceptual evolution.
for a biological
basis for certain social features of human societies.
However, when his critics
(Allen, et. al. 1976) argued for a sociological basis for certain features of his conceptual system, Wilson obThese issues, obviously, need more careful investigation.
jected.
However, the implication for human beings from evolution which I
find most fascinating concerns the existence of "human nature".
For
centuries philosophers, scientists,
theologians and the general public
have argued over the particulars
of humannature while assuming that it
exists.
Numerous traits have been suggested for the "essence" of Homo
sapiens--rationality,
language use, intensionality,
the plantigrade
For some reason, this question has seemed extremely imporfoot, etc.
If bees have a language or computers can think, then we are in
tant.

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

88
danger of having to include them in our species.
However understandable these inferences are, given our common-sense ways of viewing biological species, they are not justified
on the basis of at least certain versions of evolutionary theory.
In the earlier discussion of inclusive fitness and the cost of meiosis, the importance of genes which are identical through descent as
distinct from being just similar in structure was emphasized.
If species are the things which evolve as the result of selection processes
occurring at lower levels,
then organisms are included in the same
species because of gene transmission, not similarity.
Being part of
the same genealogical
nexus is what counts (Ghiselin 1974).
As biologists have emphasized, organisms which are phenotypically quite disparate can belong to the same species while organisms which are all but
phenotypically indistinguishable
can belong to different species.
The
sort of variation which occurs in biological
species is such that the
notion of a "typical" organism makes no sense.
Perhaps at any one locus, there will be one allele which is most prevalent, but it is possible that no one organism has ever possessed all the commonest alleles.
In fact, they might be developmentally
incompatible.
What does this imply about Homo sapiens?
It implies that we are all
part of the same species in virtue of descent and mating. We are not
all members of the same species in virtue of possessing its essential
traits, or even enough of its most important traits.
Domesticated animals may be part of human societies,
but they are not part of the human
species.
Humanreproductive neuters, regardless of the mechanism, remain part of the human nexus, albeit at termini. The failure of human
neuters to contribute directly to human biological
evolution does not
mean that they cannot contribute indirectly through kin selection or by
means of social influences.
On this perspective, people with mental ano higher than those of apes nevertheless remain part of the
bilities
human species.
The same can be said for all other traits which have
for human nature.
As important as the
been suggested as "essential"
emergence of the apposable thumbwas in the evolution of Homo sapiens,
No fraudulent
people born without thumbs remain no less human beings.
references to "potentiality"
is needed, as if people without the genetic instructions necessary to develop a thumbnevertheless potentially
possess a thumb. In this sense, pandas and porpoises also potentially
possess apposable thumbs. Although individuals lacking one or more
"essential"
traits may be less than "human" in a variety of senses of
Althis term, they are no less a biological
part of !Homos
E
abthough human homosexuals may be immoral, sinful, psychologically
normal and even criminal, at least they need not be considered any more
biologically
abnormal than worker bees or soldier ants.
The message of the preceding discussion is that particular species
need have no essences on certain versions of evolutionary theory; they
Even so, the species category itcannot have them on other versions.
All species might
self might well be a natural kind with an essence.
Similar
have some essential
feature or features in common(Hull 1980).
if societies
observations hold for societies
evolve in anything like

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

89
then
If phyletic evolution is possible,
the same way that species do.
can change indefinitely through time until later stages of
societies
which characterized earlier
the society have none of the peculiarities
to it, the
Hence, nothing about American society is essential
stages.
turn out to be homeoEven if societies
Constitution notwithstanding.
static systems which require revolutions to change, they are likely to
None of this entails
be marked by considerable internal diversity.
as such have nothing in common. The class of all sothat societies
cieties might well form a natural kind characterized by certain essential traits.
So the story goes, Linnaeus has been designated the "type specimen"
for Homo sapiens.
The peculiarity of claiming that an organism (as
is typical of its species
distinct from a character state or an allele)
can be seen by asking in what sense a Caucasion, male, Swede is a
The point is not that Linnaeus is the wrong
"typical" human being.
person to choose as a type specimen but that no one organism could possignificant
sibly be "typical" of its species in an evolutionarily
Gould (1980, p. 116) remarks that human history "remains so resense.
ideographic because it is the story of single species--it
calcitrantly
of an individual (Ghiselin 1974) of unparrepresents the vicissitudes
It
W4hatgeneral theory could encompass it?"
flexibility.
alleled
had no nature,
would probably be misleading to say that H
ascribed
but species do not have "natures" in the sense traditionally
The implications of this feature of evolutionary
to natural kinds.
theories which are limited solely and necessarily
theory for scientific
are fundamental and far-reachto a single species such as Homo si
ing (Rosenberg 1980).
Notes

'The research for this paper was supported by a Guggenheim Fellowship

for 1980-1981.
I wish to thank Elliott
Sober and William Wimsatt for
commenting on an early draft of this paper.
remain reasonably
2T,o-locus models and models for multiple alleles
tractable, but they rapidly become prohibitively complex as they are
extended and combined
(Roughgarden 1979).
As Wimsatt (1980) argues,
no general solution is possible for multi-locus models.
3Mayr is not unaware of the conflicts and unresolved problems concerning issues on which he has worked for over forty years; see for
example the introduction to the 1964 edition of Mayr (1942).

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

90

References
Darwinismnand HumanAffairs.
Alexander, R. D. (1979).
University of Washington Press.
Allen,

Biological

"Sociobiology--Another
(1976).
L., et. al.
minism." Bioscience 26: 133-149.

WA:
Deter-

The American Nat-

"What Does Sex Really Cost?"

Barash, D. P. (1976).
uralist 110: 894-896.
(1979).
Blute, Marion.
ory." Behavioral

Seattle,

An Untried The-

Evolutionism:
"Sociocultural
Science 24: 46-59.

"On the Genetics of Altruism and the Counter(1972).

Campbell, D. T.
Journal of Social Issues
Hedonic Components in HumanCulture."
28: 21-35.
In PSA 1980, Vol"Have Species Become Declasse?"
Caplan, A. (1980).
East Lansing, MI:
ume 1. Edited by P. D. Asquith and R. N. Giere.
Pages 71-82.
Philosophy of Science Association.
Commoner,B.
Dawkins, R.
Press.

(1961).
(1976).

"In Defense of Biology."

The Selfish

Gene.

Science

New York:

1.33: 1745-174a

Oxford University

"Replicator Selection and the Extended Phenotypea"

(1978).
?-----.
Zeitschrift fiur Tierpsychologie 47: 61-76.
?-----.

(1979).

"Defining Sociobiology."

Nature 280: 427-428.

An Al"Punctuated Equilibria:
Eldredge, N. and Gould, S. J. (1972).
In Models in Paleobiology.
ternative to Phyletic Gradualism."
Freeman, Cooper &
Edited by T. J. M. Schopf. San Francisco:
Company. Pages 82-115.
"A Radical Solution
Ghiselin, M. (1974).
Systematic Zoology 23: 536-544.

to the Species

Problem."

"The Promise of Paleobiology as a Nomothetic,

Gould, S. J. (1980).
Paleobiology 6: 96-118.
Evolutionary Discipline."
"The Genetical
(1964).
Hamilton, W. D.
Journal of Theoretical Biology 7:
"Are Species
(1976).
Hull, D. L.
ology 25: 174-191.

Really

Evolution
1-52.

of Social

Individuals?"

"A Matter of Individuality."

(1978).
------.
ence 45: 335-360.

Behavior."

Systematic Zo-

Philosophy of Sci-

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

91

(1980).
and Selection."
"Individuality
Ecology and Systematics 11: 311-332.

?.------.

Lewontin, R. C.
New York:

The Genetic Basis

(1974).
Columbia University Press.

Maynard Smith, J. and Williams, G. C.

American Naturalist 110: 897.

Annual Review of

of Evolutionary

(1976).

"Reply to Barash."

(1942).
Mayr, E.
Systematics and the Origin of Species.
Columbia University Press; 1964 edition, New York:
cations, Inc.
(1963).
Animal Species
Harvard University Press.

?.------.

(1969).
McGraw-Hill.

?_____--.

Principles

Change.

and Evolution.

(1975).
"The Unity of the Genotype."
tralblatt 94: 377-388.

New York:
Dover Publi-

Cambridge, MA:

of Systematic Zoology.

----------.

The

New York:

Biologisches

Zen-

Nietzsche, F.
(1901).
Der Wille zur Macht. In Nachgelassene Werke,
Vol. 15. Leipzig:
C.G. Naumann. (As Reprinted in The Will to
Power. (1967).
Walter Kaufmann and R.J. Hollingdale.
(trans.)
New York: Random House.)
Nisbett, R. and Ross, L.
(1980). Human Inference Strategies and Shortcomings of Social Judgment. Englewood Cliffs, N.J.:
Prentice Hall.
Rosenberg, A. (1980).
Sociobiology and the Preemption of Social
ence.
Baltimore, MD: Johns Hopkins University Press.
Roughgarden, J.
Macmillan.

(1979).

Theory of Population

Genetics.

Sci-

New York:

Sober, E.
(1981).
"Holism, Individualism, and the Units of Selection."
In PSA 1980, Volume 2. Edited by P.D. Asquith and R. N. Giere.
East Lansing, MI: Philosophy of Science Association.
Pages 93-121.
Stampe, J. A. and Metcalf, R. A. (1980).
"Parent-Offspring Conflict."
In Sociobiology:
Beyond Nature/Nurture? Edited by G. W. Barlow
and J. Silverberg.
Boulder, CO: Westview Press.
Pages 589-618.
Trivers, R. L. (1971).
"The Evolution of Reciprocal
Quarterly Review of Biology 46: 35-57.

Altruism."

The

Turner, B. J. and Grosse, D. J. (1980).

"Trophic Differentiation
in
Ilyodon, A Genus of Stream-dwelling Goodeid Fishes:
Speciation
versus Ecological
Polymorphism." Evolution 34: 259-270.

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

92
Wade, M. J.
tion."

"A Critical Review of the Models of Group Selec(1978).

The Quarterly Review of Biology 53: 101-114.

"The Evolution of Social Behavior by Kin

West Eberhard, M. J. (1975).
The Quarterly Review of Biology 50: 1-33.
Selection."
Williams, G. C. (1966).
Adaptation and Natural Selection.
NJ: Princeton University Press.

Princeton,

The Natural Selection of Populations

(1980).
Wilson, D. L.
Menlo Park, CA: Benjamin Cummings.
munities.

and Com-

The Insect
Wilson, E. 0.
(1971).
University Press.
(1975).
Sociobiology:
Harvard University Press.

----------.

Societies.

Cambridge, MA: Harvard

The New Synthesis.

Cambridge, MA:

"Reductionistic Research Strategies

Wimsatt, W. (1980).
and their Bases in the Units of Selection Controversy."
Edited by
Case Studies.
In Scientific Discovery:
D. Reidel.
Pages 213-259.
T. Nickles.
Dordrecht-Holland:
"Units of Selection and the Structure of the Mul(1981).
ti-level Genome." In PSA 1980, Volume 2. Edited by P. D. AsEast Lansing, MI: Philosophy of Science
quith and R. N. Giere.
Pages 122-183.
Association.

?__--_--.

Animal Dispersion in Relation

(1962).
Wynne-Edwards, V. C.
Behavior.
Edinburgh and London: Oliver and Boyd.
----------------.

Social

(1963).
"Intergroup Selection
Systems." Nature 200: 623-626.

to Social

in the Evolution

This content downloaded from 142.51.1.212 on Mon, 21 Jul 2014 16:44:42 PM

All use subject to JSTOR Terms and Conditions

of