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Natural Selection
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Stanford Encyclopedia Darwin's theory of evolution by natural selection provided the first, and
only, causal-mechanistic account of the existence of adaptations in nature.
of Philosophy As such, it provided the first, and only, scientific alternative to the
“argument from design”. That alone would account for its philosophical
significance. But the theory also raises other philosophical questions not
encountered in the study of the theories of physics. Unfortunately the
concept of natural selection is intimately intertwined with the other basic
concepts of evolutionary theory—such as the concepts of fitness and
Edward N. Zalta Uri Nodelman Colin Allen R. Lanier Anderson
adaptation—that are themselves philosophically controversial. Fortunately
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we can make considerable headway in getting clear on natural selection
Editorial Board
http://plato.stanford.edu/board.html without solving all of those outstanding problems.

Library of Congress Catalog Data 1. Natural Selection and Evolutionary Theory

ISSN: 1095-5054
2. Natural Selection and Fitness
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Center for the Study of Language and Information 1. Natural Selection and Evolutionary Theory
Stanford University, Stanford, CA 94305
Natural Selection The theory of evolution by natural selection forms a central part of modern
Copyright c 2014 by the author
Robert Brandon
evolutionary theory. There is some controversy among biologists as to just
how important natural selection is compared to other processes producing
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1
 Natural Selection Robert Brandon

evolutionary change, but there is no controversy over the proposition that from repeated iterations of this set-up. It leaves open the question of
natural selection is important. Some good might come of the efforts to whether this process is natural selection or drift (see below). It—the
produce a general selection theory that would include the natural selection causally neutral statement—does not suffice to state Darwin's causal
that occurs as a part of the evolutionary process as a special case (e.g. Hull theory. Darwin clearly recognized this (see, for example 1871) as did
2001), but here the focus will be solely on the evolutionary process. Lewontin (1978); although many contemporary commentators fail to see
this.
Biology starts when reproduction begins. Stars may be said to evolve, but
they do not reproduce and so biological theory does not apply to them. Why is it that some variants leave more offspring than others? In those
Biological evolution requires reproducing entities that form lineages. It is cases we label natural selection, it is because those variants are better
these lineages that evolve. Thus it is only within such lineages that we will adapted, or are fitter than their competitors. Thus we can define natural
properly apply the term natural selection. A kindergarten class may selection as follows: Natural selection is differential reproduction due to
certainly select among different colored candies, but since those candies differential fitness (or differential adaptedness) within a common selective
are not part of self-reproducing lineages, we should not confuse this environment (see next section). This definition makes the concept of
selection process with natural selection. natural selection dependent on that of fitness, which is unfortunate since
many philosophers find the concept of fitness deeply mysterious (see e.g.,
2. Natural Selection and Fitness Ariew and Lewontin 2004). But like it or not, that is the way the theory is
structured. And, fortunately, we can make considerable headway in
Natural selection is a causal process. Distinguishing it from other understanding natural selection without solving all of the philosophical
processes in evolution is one of major conceptual and empirical problems problems surrounding the concept of fitness.
of evolutionary biology. The bare bones of Darwin's theory of evolution
by natural selection are elegantly simple. Typically (but not necessarily) 3. Common Selective Environments
there is variation among organisms within a reproducing population.
Oftentimes (but not always) this variation is (to some degree) heritable. As a causal theory natural selection locates the causally relevant
When this variation is causally connected to differential ability to survive differences that lead to differential reproduction. These differences are
and reproduce, differential reproduction will probably ensue. This last differences in organisms' fitness to their environment. Or, more fully, they
claim is one way of stating the Principle of Natural Selection (from here are differences in various organismic capacities to survive and reproduce
on PNS). The PNS goes beyond the causally neutral statement that is in their environment. When these differences in capacities are heritable,
sometimes listed as the third of what are often called “Darwin's Three then evolution will (usually) ensue. This sort of case must be carefully
Conditions”, viz., different variants sometimes reproduce at different rates. distinguished from cases where the causes of differential reproduction are
That statement leaves open the question of whether or not the variation in not located in the organisms, but rather in their different environments. Let
question is causally responsible for the differential reproduction. It leaves us make this distinction more concrete. Imagine two genotypes of an
open the question of whether a qualitatively similar outcome would result annual plant that grows in dense patches. Sunlight is at a premium and

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 Natural Selection Robert Brandon

taller plants shade shorter plants thus garnering more energy for growth covariances, are employed to separate out the effects of genotype vs.
and reproduction. One genotype, G1, grows more quickly at germination environment.)
than the other, G2. Thus G1s are initially taller than G2s and this
difference persists throughout the growing season due to G1s increased Clearly the two cases sketched above are highly simplified, and it might
energy stores. The consequence of this is that G1s produce more flowers, be objected that nature is unlikely to contain many examples that either
more pollen and more seeds than G2s. This is natural selection at work. model exactly applies to. This objection has both epistemological and
ontological sides. The epistemological side of the objection has, I think,
In contrast, imagine two genotypes of the same species, G3 and G4 that do been met. First, the statistical techniques mentioned in the last paragraph
not differ in germination date, growth rate, resource allocation between mitigate the force of it. Even in messy cases biologists are fairly
growth and reproduction or any other factor relevant to reproduction when successful in separating out environmental causes from genetic, or
grown in a common environment. Now imagine that seeds of these two organismic, causes of differential reproduction. (It should at least be
genotypes are distributed randomly across a patchwork of two quite mentioned here that this sort of analysis also often yields genotype by
different soil types, call them E1 and E2. E1 and E2 are identical except environment interactions, G × E. This occurs when, in contrast to case 2,
that E1 contains high levels of lead whereas E2 does not. Lead the relative performance of different genotypes differs in different
dramatically and equally reduces growth and reproduction in both G3 and environments. Although common in nature, and very important, we can
G4. Finally imagine that this random distribution process results in a ignore that here.) Second, the experimental techniques and conceptual
correlation between genotype and environment—i.e., G3s are resources developed by Antonovics et al. (1988), Brandon (1990) and
disproportionately found in E1. In consequence of all of this, G4s produce Brandon and Antonovics (1996) allow for precise measurements of
more flowers, more pollen and more seeds than G3s. But natural selection environmental heterogeneity in real life populations.
is not occurring here. (Indeed this is a type of drift, see below.)
Ontologically the objection is this: when measured precisely enough each
In both cases differential reproduction occurs, and in both cases I have organism lives in a unique environment. Thus if natural selection requires
already sketched causal explanations of this. In neither case is differential multiple organisms competing in a common selective environment, then it
reproduction mysterious (although chance does play a role in the second never really occurs. That, if true, would be a serious objection to the
explanation, but not the first). But only the first case could result in picture of natural selection I am presenting here. But I think we have very
adaptive evolution. (See Brandon 1990, chapter 2 for fuller discussion.) good reason to believe it is not true. Unfortunately, here I can present only
Biologists have long recognized the importance of the difference the briefest sketch of it. What natural selection explanations require is
discussed above, thus the importance of so-called “common garden” consistent ordinal relations in fitness of the competing types—they do not
experiments in experimental evolutionary genetics. (In common garden require precise agreement of absolute fitnesses of the competing types.
experiments, the environments in which different genotypes are placed are Lots of empirical studies of natural selection in the wild have shown
controlled as far as possible. Furthermore, when environmental control is consistent ordinal fitness relations (see Endler 1986). Furthermore, the fact
imperfect, statistical techniques, in particular the analysis of variances and of adaptive evolution, the consistent and persistent increase of one type

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over others in evolutionary history, requires these consistent ordinal period the first may have gone extinct, while the second persists in virtue
relations, or what I have termed “common selective neighborhoods”. Thus of simply surviving without reproduction. Isn't this radical view just as
I think the ontological objection has been met as well. defensible as the more mainstream view? If so, then differential
persistence would count as natural selection, and differential reproduction
4. Does Natural Selection Require Differential would not be necessary for natural selection to occur.
Reproduction?
Interesting as this view is, I reject it because, with Dawkins (1982), I
The short answer to this question is “Yes”. I have already offered believe that reproduction, in particular going through a single cell bottle
arguments in favor of that answer. However the longer more complete neck from whence the developmental process is started anew, is necessary
answer is “No, no, but yes”. I will explain. for the evolutionary process of adaptation. Only by restarting the
developmental process each generation can fundamental alterations of that
There are two reasonable arguments that suggest a negative answer to our process be achieved. Consider a grove of aspens that grows vertically (in
question. I will review them both. In the beginning of this essay I stated what we intuitively think of as trees) and horizontally by underground
that considerable progress could be made in articulating a clear and runners that produce more “trees”. The phenotype of the whole grove can
adequate conception of natural selection without solving all of the change through time—growing vigorously here, not growing there. But
philosophical problems associated with the notion of fitness. However, in consider a genetic mutation that would fundamentally alter Aspen
this section I will have to make certain assumptions about fitness in order phenotype. It may occur in a multicellular runner. If so, it will be
to address the issues to be raised. The assumptions are quite plausible, but incorporated in the resultant cell lineage. But that cell lineage will be one
not defensible in a short essay of this sort. of many in the next “tree” produced, since the next tree grows not from a
single cell, but from a multi-cell runner. Thus the resultant “tree” will be a
One negative answer is based on a radical rethinking of the problem nature chimera, and not fundamentally different from the others in the grove. In
presents to evolving entities. Mainstream evolutionary biology measures contrast, were the runners single-celled at any cross-section, then a
fitness in terms of reproductive success. (Exactly how it defines fitness somatic mutation would be incorporated in the whole of the downstream
will not be addressed here.) Survival is important exactly insofar as it “tree”. This, according to Harper (1977) and Dawkins (1982), would count
contributes to reproduction. Evolutionary success is reproductive success. as reproduction, not growth, because this could fundamentally rearrange
But suppose we turn the relation between reproduction and survival on its Aspen development. (But that is not the way Aspens grow.) Accordingly,
head and think of reproduction as important only insofar as it contributes the evolution of complex adaptations, fundamental rearrangements of
to lineage survival. The fundamental evolutionary problem is persistence, development, requires differential reproduction.
and reproduction is but one means of achieving that (Bouchard 2004).
Compare two lineages, one composed of short-lived organisms that A second reason to answer our question in the negative comes from a
reproduce every year, the other composed of organisms that live for (quite appropriate) focus on the ecological process of selection. For
1,000+ years, but reproduce only rarely. At the end of a thousand year practical reasons, many studies of natural selection in the wild focus on

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only a small part of the life cycle. For instance, in the most famous such action. I, and the vast majority of evolutionary biologists, would agree
study H. B. D. Kettlewell (1955, 1956) marked different morphs of the with this conclusion.
peppered moth, Biston betularia, released them (into polluted woods in
one treatment, non-polluted woods in another) and then recaptured them In contrast, imagine a second example. It is just like the first except that in
three days later. The difference in the proportion of dark to light forms in the larval stage the Lights out survive the Darks by a two to one margin.
the recaptured vs. the released groups was taken to measure natural This difference in survival in the larval stage exactly counterbalances the
selection in action. As Rudge (1999) points out, Kettlewell had preformed difference in survival in the adult stage, leading to no difference in
numerous auxiliary studies to justify this inference, but for present reproductive success between the two morphs. Were this going on we
concerns what is crucial is that Kettlewell looked at only a small portion couldn't explain the match between frequencies observed in different
of the entire life cycle, and in particular a portion that did not involve woods and the selective processes occurring in the adult stage. But that is
reproduction. It is beyond the scope of this essay to critically examine not crucial to our question. The question for us is: Are the two processes
Kettlewell's actual work, but let us use it as a basis to construct two observed in the mark-release-recapture studies in our hypothetical cases
hypothetical examples. both examples of natural selection?

The first example mirrors the situation with which Kettlewell was actually If you think, as has been suggested, that natural selection requires
dealing. Let us suppose that there are two genetically distinct morphs in a differential reproduction, then you must say that the second example is not
population of moths, call them Light and Dark. In survivorship through one of natural selection. But one might object to this conclusion as
the larval stage both forms are identical. They are also identical in fertility follows. The ecological process of birds preying on moths based on their
and fecundity. They differ only in survivorship during the adult stage prior relative conspicuousness is exactly the same in the two examples. That
to mating. This difference is due to a difference in their conspicuousness ecological process was identified as natural selection in the first example;
to birds as they rest on the trunks of trees. On dark trees the Darks are less therefore it must be natural selection in the second example as well.
conspicuous than Lights, and vice versa on light trees. In a polluted wood,
Although quite compelling I think the above should be resisted in the
most of the tree trunks are dark. A biologist marks equal numbers of Darks
following way. Fitness attaches to the whole life cycle, not some subpart
and Lights, releases them into the polluted woods and recaptures them
of it. Why? Again the short answer relies on a commitment to the
three days later. She recaptures twice as many Darks as Lights. Based on
fundamentality of reproduction. Reproduction is a reproduction of the
her knowledge of when pollution was introduced into the woods, the
entire life cycle. It is true that fitness is often measured, as in Kettlewell's
frequency of the two morphs in woods that have not been polluted, the
case or in our hypothetical examples, by looking at only a part of the life
underlying genetics of the two morphs, and her observed selection
cycle. But from an evolutionary point of view we are interested in this
differential between them, she builds a population genetic model of the
only when relative performance in this part of the life cycle actually
situation that predicts a relative frequency of the two morphs for the
influences relative reproductive success. Thus the first, but not the second,
present. That prediction fits the observed frequencies. She concludes that
is a case of natural selection. (Consider how Kettlewell's studies would
her mark-release-recapture experiment has captured natural selection in

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have been received by the evolutionary community if they had mirrored A value of 1 represents a perfect correlation between offspring and
our second hypothetical example rather than the first.) parental deviations from their respective means, while a value of 0 means
there is no correlation, so that, for example, the offspring of tall parents
And so natural selection does require differential reproduction. would not differ statistically from those of short.

5. Does Natural Selection Require Heritable Unfortunately many people think that some sort of genetic definition has
Variation? supplanted this basic concept of heritability. In particular, many would say
that the Galtonian notion corresponds to the “narrow sense” of heritability
In the preceding section we had to draw some fairly subtle distinctions, (h2), h2 = VA/VT , where VA is the additive genetic variance and VT is the
but our conclusion is one with which the vast majority of evolutionary total variance. (VT is usually decomposed into VA, VD—the variance due
biologists would agree. There is no such answer to the question of this to allelic dominance, and Ve—the environmental variance. The latter is in
section. Many biologists define natural selection as differential fact a statistical catch-all, so it includes not just variation due to
reproduction of heritable variation (see, e.g., Endler 1986). Many other environmental differences, but everything else.) Even though this equation
biologists follow the tradition of quantitative genetics and draw a sharp is very important and we will revert to it shortly, we cannot accept this as
distinction between the ecological process of selection and the a definition of heritability for at least two reasons: (1) Even for diploid
evolutionary response to selection (see e.g., Falconer 1981). There are a sexually reproducing organisms that equation holds only under certain
large number of advantages to the second approach, and I will follow it genetic conditions (Roughgarten 1979); and (2) It certainly is not
here. Thus we will arrive at a negative answer to the question of this applicable to pre-genetic or non-genetic systems. But that would mean that
section. the theory of evolution by natural selection would not be applicable to the
early stages of life on this planet, nor to epigenetic inheritance, nor to
First we must get clear on the relevant sense of heritability. Recall § 2 cultural transmission, nor to life elsewhere in the universe. Surely such
above where we stated “Darwin's Three Conditions”. They are: variation; consequences are unacceptable and entirely unnecessary. Thus the
heredity; and differential reproduction. The notion of heritability relevant Galtonian notion of heritability is fundamental.
here is the purely phenotypic, purely statistical notion developed by
Francis Galton (1869). That notion identifies heritability with the The motivation for saying that natural selection requires heritable
regression of the offspring phenotype on the parental (or biparental mean variation is clear. Only selection acting on heritable variation will have
in the case of sexual reproduction), where both phenotypes are presented evolutionary consequences. And since we are interested in the concept of
as z-scores (i.e., set to mean = 0 and standard deviation = 1). Intuitively natural selection from a purely evolutionary point of view (recall the
the heritability is a measure of how closely offspring deviation from the introduction to this piece), we don't count selection acting on non-heritable
(offspring) mean phenotypic value matches that of the parental deviation variation as natural selection. (This seems analogous to the argument in
(from the parental mean). That is, it measures the degree to which, for the last section saying that selection acting in a part of the life cycle that is
example, taller than average parents produce taller than average offspring. exactly counterbalanced later in the life cycle should not count as natural

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selection.) derived and not general; it is useful for simple genetic models like the one
just described. In this example, once the stable equilibrium is reached, the
One response to this line of reasoning is to insist on the importance of heritability, h2, is zero. This is true because the additive genetic variation
drawing a sharp distinction between the ecological process of selection is zero. (To apply the Galtonian notion of heritability to this example, we
and the evolutionary process of response to selection. The ecological would need to complicate it by specifying a genotype-phenotype mapping,
process of selection, the domain of ecological genetics, is complicated and since that notion relates phenotypes. We could do this, and provided Ve is
difficult to study. The evolutionary response to selection is the domain of not zero the Galtonian value would be zero. But the point presently under
population genetics. It too is complicated and difficult. From a purely discussion would not be made stronger by this extra complication.) So
strategic point of view it would seem wise not to conflate these two here we have a case where there is phenotypic variation caused by
complex processes into one. genotypic variation, but the heritability of this variation is zero because of
the non-linear relationship between genotype and fitness. How do we
I think most would agree with this bit of wisdom, but some would counter
categorize the differential reproduction that occurs in this example? Those
by saying that it does not require a definition of natural selection that is
who think that natural selection requires the differential reproduction of
neutral on the heritability of the variation in question. We can define
heritable variation would say there is no natural selection here. Those
‘natural selection’ one way or another and still agree on all of the facts.
following the quantitative genetics tradition would say that natural
The argument for one definition over the other will have to be made in
selection is occurring, but that since h2 is zero the response to selection is
terms of simplicity, elegance, or some other non-empirical virtue of
zero. (In accordance with the fundamental formula of quantitative
theoretical constructs.
genetics, R = h2S, where R is the response to selection and S is the
Let us illustrate the last point with a simple example, an example that has selection differential.) But, it seems, there is no empirical difference
been commonly used in the philosophical literature on units of selection. between these two points of view, so we need to decide between them on
The example is of heterozygote superiority, though any example where the the basis of some non-empirical reasons.
allelic effects on phenotype are non-additive would do. (As would any
The seeming empirical equivalence between the two accounts offered of
number of cases involving non-additive interactions among multiple
our example is, in fact, illusory. Godfrey-Smith and Lewontin (1993) have
genes.) There are two alleles at a locus, A and a, and thus three genotypes,
pointed out that there is an empirical difference between an account of this
AA, Aa, and aa. For simplicity suppose that the two homozygotes are
case that says there is selection vs. one that says there is no selection.
lethal, i.e., they die before reproducing. Thus all the matings are Aa x Aa.
(They make this point in the context of discussing genic selectionism,
Mendelism results in offspring of genotypes AA, Aa and aa, in the ratio of
which is committed to saying that there is no selection in this case since
1:2:1. In a single generation the allele frequencies equilibrate at 50:50 and
the allelic fitnesses are the same. For further discussion of this see
stay there unless the fates of homozygotes changes.
Brandon and Nijhout 2006.) The empirical difference is that the first, but
Although we criticized the genetic “definition” of heritability as being not the second, can account for the within generation change in genotype
frequencies. At the start of each generation the three genotypes are

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represented in the 1:2:1 ratio discussed above. Later in the generation the process, namely a probabilistic sampling process (Brandon and Carson
homozygotes, both AA and aa, die, so that 100% of the population is 1996, Matthen and Ariew 2002, Walsh et al. 2002). Thus, although it is of
composed of Aas. However, this difference is not an evolutionary crucial importance to separate selection and drift, one cannot do so on the
difference, since evolution is trans-generational change. Our focus is on basis of process alone (contra Millstein 2002), one must do so on the basis
evolution. And it turns out that the two models are not empirically on outcomes (Brandon 2005). Why is this? If we think of fitness as a
equivalent with respect to evolutionary predictions. The standard account, probabilistic propensity, then, as we have seen, differential fitness is a
which says that selection occurs each generation against the homozygotes, necessary condition for natural selection. Thought of this way, natural
predicts a stable equilibrium—one actively maintained by strong selection. selection is a probabilistic sampling process. We can characterize a
But the account that says that there is no selection in this case has no basis continuum of all possible fitness differences starting with maximal fitness
to predict the same sort of stable equilibrium. This is not apparent if one differences at one end of the continuum (i.e. where all fitness equal either
only considers selection, because the genic selectionist recognizes the 1 or 0, with at least some of each value), and minimum fitness differences
existence of selection once one perturbs the population from its on the other (i.e., an equiprobable distribution). The two endpoints are
equilibrium and so, it would seem, both models predict the same stable exceptional with respect to the relation of selection and drift. At the
equilibrium. However, once one brings in considerations of the interaction maximal fitness difference end, one unlikely to occur in nature, drift is
of drift and selection (see next section) it has been shown that the models impossible and selection is necessary. (See Figure 1.) At the finite set of
do not give empirically equivalent predictions (Brandon and Nijhout minimal fitness differences at the other end of the continuum,
2006). This is because small perturbations from equilibrium will have very corresponding to absolute neutrality of the traits under consideration,
different likelihoods of drift, because they will experience quantitatively selection is impossible, because there are no fitness differences. And drift
quite different regions within which drift vs. selection is likely to is maximally likely, but not necessary. Why not? Because the sample may
dominate. be in accord with the probabilistic expectations and thus no drift occur.
(Imagine a population with two alleles, A and a, absolutely selectively
This example has a number of consequences (again see Brandon and neutral with respect to each other, and at a 50:50 ratio. The next
Nijhout 2006 and Brandon 2006). But for us the consequence is this. To generation may also contain the two alleles at the same ratio, if so, no drift
say that selection requires heritable variation is factually wrong. When has occurred.
applied to a broad class of cases it makes the wrong evolutionary
predictions. And so we must reject that view, and conclude that natural
selection does not require heritable variation.

6. Natural Selection and Drift

Why should an entry on natural selection contain a section on drift? One
good reason is that natural selection and drift are co-products of the same

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 Natural Selection Robert Brandon

gone in accord with the probabilistic expectations, or not. To the extent

that it has not, drift has occurred. Drift simply is the deviation from
probabilistic expectation. And since selection itself is a probabilistic
process there can be no purely process-derived distinction between
selection and drift—attractive as that idea may be.

A more fundamental reason for a discussion of drift here in an entry on

natural selection is the view that drift is the zero-force background against
which evolutionary forces, including natural selection, act (Brandon
2006). Just as in Newtonian mechanics one could not properly understand
the notion of gravitational force, without understanding Newton's 1st Law,
Figure 1. The heavy horizontal line, with dotted center section, similarly one cannot understand natural selection in evolutionary biology
represents the infinite number of possible fitness distributions from without understanding the background against which it operates. Put in
Maximal Probability Differences (MPD—all fitness = 0 or = 1, other terms, drift provides the appropriate null hypothesis against which to
with some of both) on the left to the Equiprobable Distribution (EP test any selection hypothesis. Unfortunately this is not always well
—all fitness the same) on the right. The arrows emanating from the understood. Indeed this way of thinking about evolution stands some
different descriptions of the modalities of selection and drift canonical versions of evolutionary theory on their heads. For instance,
indicate the areas of the distribution falling under these those who would take the Hardy-Weinberg Law as a Zero-Force Law of
descriptions. evolution (see, e.g., Ruse 1973, and Sober 1984, but also many standard
textbooks in evolution) view stasis as the default state of evolutionary
But these two endpoints of this continuum are exceptional. More common systems, with some evolutionary force needed to move (i.e. evolve) them.
is surely the vast middle area where drift and selection are both possible. Without a net force, no net change. But all modern methodology in
In this middle ground we cannot say until after the fact whether or not drift molecular evolution is predicated of the truth of just the opposite idea,
occurs, nor quantify the degree to which it occurs. Prior to the fact we can namely that left alone evolutionary systems drift. Drift is the default state.
only quantify the relative likelihoods of selection and drift—they vary So that, for instance, neutral alleles in different populations differentiate
according to the crucial quantity 4Ns. (Where N is the effective population from each other. But this molecular truth is iterated throughout the
size and s is the selection coefficient, i.e. the strength of selection. For biological hierarchy. Once speciation occurs, species differentiate (drift
discussion of 4Ns see any standard textbook on population genetics, e.g., apart) as a null expectation. Which is not to say that natural selection
Roughgarden 1979, see also Brandon and Nijhout, 2006.) The larger that cannot produce evolutionary change. Of course it can. But if we are to
quantity is, the larger N and the larger s, the greater is the likelihood that properly recognize it, we must be able to recognize the signature of
selection will dominate drift. And vice versa for small N and small s. Only selection and differentiate it from drift's signature (see, e.g., Bamshad and
after the fact can one tell whether or not the probabilistic sampling has Wooding 2003). Change in evolution is a heterogeneous category.

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 Natural Selection Robert Brandon

Stasis, on the other hand, is largely homogeneous. Long-term stasis can Philosophy of Science 73: 277–297.
only occur by natural selection. Darwin, C., 1871, The Descent of Man, London: John Murray.
Dawkins, R., 1982, The Extended Phenotype, Oxford: Freeman.
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variation and environmental variation: Expectations and York: Springer-Verlag.
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Bouchard, F., 2004, Evolution, Fitness and the Struggle for University Press.
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reply to Millstein,” Biology and Philosophy 20: 153–170. Lewontin, R. C., 1978, “Adaptation,” Scientific American 239 (9):
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Journal of Philosophy 103(7): 319–335. Matthen, M. and Ariew, A. 2002, “Two ways of thinking about
Brandon, R. N., and Antonovics, J., 1996, “The coevolution of natural selection,” Journal of Philosophy 49(2): 55–83.
organism and environment,” in Concepts and Methods in Millstein R. L. 2002, “Are Random Drift and Natural Selection
Evolutionary Biology, R. N. Brandon, pp. 161–178. Cambridge: Conceptually Distinct?” Biology and Philosophy 17: 33–53.
Cambridge University Press. Roughgarden, J. 1979, Theory of Population Genetics and
Brandon, R. N., and Carson, S. 1996, “The indeterministic character Evolutionary Ecology: An Introduction. New York: Macmillan
of evolutionary theory: no ‘no hidden variable proof’ but no room for Publishing Company, (Reprinted 1987 by Macmillan, and in 1996 by
determinism either,” Philosophy of Science 63: 315–337. Prentice Hall).
Brandon, R. N. , and Nijhout, H. F. 2006, “The empirical non- Rudge, D. W., 1999, “Taking the peppered moth with a grain of salt,”
equivalence of genic and genotypic models of selection: a (decisive) Biology and Philosophy 14: 9–37.
refutation of genic selectionism and pluralistic genic selectionism, ” Ruse, M., 1973, The Philosophy of Biology, London: Hutchinson

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 Natural Selection

Publishing Group, (Reprinted 1998 by Prometheus Books).

Sober, E., 2004, The Nature of Selection: Evolutionary Theory in
Philosophical Focus, Cambridge: MIT Press.
Walsh, D., Lewens, T. and Ariew, A., 2002, “Trials of life: natural
selection and random drift,” Philosophy of Science 69: 452–473

Other Internet Resources

Evolution 101: How It Works, at the Understanding Evolution
website, maintained at the University of California/Berkeley.
Natural Selection: How Evolution Works, at ActionBioscience.org,
maintained by the American Institute of Biological Science.

Related Entries
adaptationism | biology: philosophy of | causation: probabilistic |
character/trait | creationism | Darwinism | fitness | genetics: and genomics |
genetics: evolutionary | genetics: genotype/phenotype distinction |
genetics: population | heritability | laws of nature | life | natural selection |
natural selection: units and levels of | probability, interpretations of |
scientific explanation | species | teleology: teleological notions in biology

Copyright © 2014 by the author

Robert Brandon

20 Stanford Encyclopedia of Philosophy