Abstract
One of Darwin’s major contributions to our understanding of evolution, namely natural selection, seems a very simple idea. However natural selection is a very subtle concept and biologists and philosophers have been struggling for decades to make sense of it and justify its explanatory power. In this chapter, first I present the most general formulations of natural selection in terms of necessary conditions, and I argue that none of them capture all the aspects of the concept. Second, I question the explanatory status of selection, asking what exactly it is supposed to explain, and considering its relationship with stochastic factors (i.e. genetic drift). Second, I investigate its metaphysical status, asking whether it can be seen as a law, and to what extent it would deprive evolution of any contingency. The last section presents controversies about the units and levels of selection, and, after exposing the philosophical assumptions proper to various positions, sketches a pluralist conception.
This is a preview of subscription content, log in via an institution to check access.
Access this chapter
Tax calculation will be finalised at checkout
Purchases are for personal use only
Notes
- 1.
- 2.
Not “speculations”, but rather questions about ontological engagement and rules for validating scientific theories (for example, the debate on realism versus instrumentalism, the interpretation of probabilities, etc.).
- 3.
- 4.
- 5.
See Sect. 4 below.
- 6.
Sober’s distinction originally concerns selection for some traits and the selection of traits (or alleles, i.e. different versions of the same gene) correlated to precedents (and not for what they are in and of themselves). I mean here the relation between selection of organisms and selection for (or because of) these organisms’ traits, but clearly there is selection-of traits correlated to traits for which there is selection because the former are in the same organism as the latter.
- 7.
Beyond highlighting the probabilistic nature of selection and hence of fitness (Beatty and Mills 1979), which is a rather weak defense that would leave open the possibility that empirical validity of the selectionist explanation depends on the weakness of our cognitive ability (Michod 1999), there are other responses to this “argument”; for example, to point out that “tautology” is not in and of itself bad: mathematics are a great tautology, and are the basic structure of physics. By the same token, the principle of natural selection would support all population genetics, which are essentially a set of mathematical models, and in this sense the tautological nature is in no way a serious objection. On tautology, see Brandon (1990).
- 8.
See e.g. Fisher (1930).
- 9.
Whose elaboration would run its course over three decades, through Galton, Pearson, Fisher – See Gayon (1998) for this story.
- 10.
Gayon (1998) insists on this point that Darwin indeed offers a hypothesis, that afterwards Darwinians will construct a test and justification.
- 11.
- 12.
This does not hold true for all modern synthesis, see Mayr: “Evolution is not a change in gene frequencies, as is often stated, but the maintenance or improvement of the adaptation and the origin of diversity. Changes in gene frequencies are a result of this evolution, not its cause.” (Mayr 1998, 2093).
- 13.
- 14.
See Shoenauer’s, Chap. 28, this volume; and Holland (1995).
- 15.
Later, Endler (2006) recapitulates (by inverting C2 and C3): “Natural selection can be defined as a process in which: If a population has:
-
C1. variation among individuals in some attribute or trait: variation.
-
C2. a consistent relationship between that trait and mating ability, fertilizing ability, fertility, fecundity, and, or, survivorship: fitness differences.
-
C3. a consistent relationship, for that trait, between parents and their offspring, which is at least partially independent of common environmental effects: inheritance.” The formulation is clearer and I will refer back to it at times.
-
- 16.
Brandon and Mc Shea (2011) make a strong claim for drift (see below) being a cause of evolution and selection being very often stabilising.
- 17.
It is more explicit in Endler’s formulation.
- 18.
Genes are not required to be single determinants of a trait. It is only required that the fact of having a gene makes a difference to the value of the trait (see Waters 2005).
- 19.
Locus (plural: loci): the physical location of a gene on a chromosome.
- 20.
- 21.
Additive variance is variance caused by the contribution of alleles whose effects are presumed to be additive. In reality they are only rarely additive, but this is only a model, that can be made more complex and allows us to define h2. Recent findings on epigenetics call for a more sophisticated partition of h between genetic and epigenetic transmission variance (Danchin et al. 2011).
- 22.
Quantitative genetics takes selection experiments as its paradigm: one selects a group of individuals who have a required phenotypic value and breeds them. The result, and thus the “response” to the selection will be proportional both to the average of their phenotypic values and to the trait’s heritability. If this is 1, the following generation will have an average phenotypic value of the selected parents; if it is ½, the average phenotypic value will be half, etc.
- 23.
See the chapter “Variation” for theories of variation; here we are only assuming the fact of variation.
- 24.
- 25.
As to the role of chance variation related to selection, and especially the importance of the order of random mutations (as well as the fate of this notion by Darwin and by the Modern Synthesis biologists) see Beatty (2011).
- 26.
See especially Godfrey-Smith 2009.
- 27.
Godfrey-Smith (2000) demonstrates with a thought experiment that the concept of replication itself is not essential for selection.
- 28.
See Griesemer (2000) for an attempt at reinterpreting selection in general using the yardstick of works on evolutionary transitions.
- 29.
- 30.
See Glymour (2006) for a radical critique of the notion that population genetics provides a general dynamic of selection.
- 31.
Group of statements corresponding to what is explained or to be explained (singular: explanandum).
- 32.
“Adaptation” refers both to the result of selection – a trait – and the process that leads to it. Here, this second meaning is completely set aside.
- 33.
Variance: see footnote 10 in Christine Clavien’s, Chap. 34, this volume.
- 34.
See Philippe Grandcolas’s chapter on adaptation, Chap. 5, this volume.
- 35.
The word “strategy” of course does not mean that organisms consciously deliberate and plan their actions; it just means a kind of determinate behaviour in given circumstances, distinct from another determinate behaviour, so that all strategies constitute a “strategy set” (for example: fight a competitor/ flight in the face of a competitor, care for the offspring after hatching/don’t care for them and mate with other partners, etc.),
- 36.
Roughgarden (2006) goes as far as contesting the validity of the idea itself, in favour of what she calls “social selection”, i.e. the forming of teams to raise offspring, but her views are controversial.
- 37.
Fitness is measured traditionally in the number of offspring, adaptedness (in the sense of adjustment to the environment allowing for a longer survival) and traits maximizing access to females are two ways of optimizing this fitness; the traits that are ultimately selected often appear as trade-offs between these two pressures.
- 38.
Grafen (1990) proposed a mathematical model of the handicap principle, which made a very powerful and explanation of it using behavioral ecology.
- 39.
A population where all mating between individuals are random; all individuals are potential partners.
- 40.
Note that in the expression of these frequencies, AA, aa, Aa are the genotypes. W is fitness; the assumptions are unrealistic of course, but this is a model; the inclined planes, with no friction, etc. in classical mechanics are the same type of unrealistic models.
- 41.
Sewall Wright elaborated the idea of “adaptive landscape”, the surface defined by the frequencies of n possible alleles on n axes, and the average fitness of the population corresponding to the combination of these n frequencies on the final axis. Such a landscape clearly shows the local and global optima, and the question is: why don’t all populations remain most often on local optima. The “shifting balance theory” mentioned here aims to resolve this problem. Moreover, the peaks are not really stable, since a population that reaches the local optimum loses genetic diversity and thus becomes more vulnerable to environmental changes. However recently Gavrilets has shown that since real landscapes are high-dimensional their mathematical properties are different from three-dimensional intuitive landscapes and allow for n-dimensional shapes that make possible shift between peaks without loss of fitness (“neutral network”) (see Gavrilets 2011).
- 42.
For neutralists, it is not exactly an question of drift in the sense Wright uses it, since he would consider the alleles themselves whereas neutralists are more interested in the stochastic fluctuation of the nucleotide composition of alleles. In both cases, though, it is a matter of a selectively neutral stochastic alternative to natural selection.
- 43.
What is called the degeneration of the genetic code.
- 44.
If a child of a blue eyed couple has brown eyes, then his real father is someone else, because the gene for blue colour is recessive.
- 45.
The compared importance of drift and selection is a crucial topic for modern evolutionary biology. Recently, Lynch (2007) argued that drift has been a very important cause of the architecture of eukaryote genome, especially because since eukaryote are often large-sized organisms, their population tend to be small, therefore drift is powerful relative to selection.
- 46.
- 47.
It does happen that one can experimentally separate the two; see Millstein (2006) who studies Lamotte’s work on the evolution of snails.
- 48.
One can argue whether or not selection is deterministic, but here I am simply pointing out that the stochasticity in the theory of evolution comes out of genetic drift and not natural selection. This is less of an ontological argument than it is an observation concerning the mathematical modeling of these concepts (see Malaterre and Merlin, Chap. 17, this volume).
- 49.
Lewens (2010) proposes a subtle analysis of the difference between “force of selection” and “selection for”.
- 50.
As is often the case with Dawkins the metaphorical nature of formulations (“the blind watchmaker”, the “selfish gene” etc.) affects the precision of his remarks; and yet on this point we can certainly classify him together with Mayr or Gould, as well as many authors of the Modern Synthesis, as someone who insists on the “creative” sense of selection – with this precision that the essential thing (from the explanation’s point of view) is the complexity of traits generated by selection.
- 51.
Kauffmann (1993) studies properties of Boolean networks in order to see the emergence of stable ordered patterns from iterated interactions between nodes.
- 52.
- 53.
Certain philosophers (Dretske 1977, or even Tooley and Armstrong) have argued that a law, before being a general statement concerning individuals, is a singular statement that links properties (for example, gravity is a single statement that links mass and distance). This position avoids well known pitfalls that appear when trying to specify seriously what separates an accidentally true universal judgment (“there is no mountain higher than 10,000 km”) and a nomothetically true universal judgment (“there is no liquid mountain”). The difficulty then boils down to understanding what constitutes an ‘genuine’ property (intuitively, “weighing 20 kilos” is an genuine property, “liking Brahms or having voted for Obama” is not; but finding the criterion that sets apart these two types of properties is tricky (see Shoemaker 1984)).
- 54.
On this concept, See Kim (1993).
- 55.
- 56.
That is, variance due to the addition of alleles’ contribution to the phenotypic value, ignoring the relationships that contradict this additivity: epistasis, dominance.
- 57.
The “hawk-dove” game was popularized by Maynard-Smith 1982 (See Clavien, Chap. 34, this volume). Hawks fight doves and the doves flee the fight; the hawks’ fitness is higher and so their fitness rises, but when there are too many hawks, it becomes more advantageous to be a dove (the hawks eliminate each other). In this sense, the mean fitness of the population does not rise, contrary to the theorem, since increasing the number of hawks increases mean fitness up to a point where hawks’ fitness becomes lower than doves’ fitness, and then population mean fitness decreases.
- 58.
- 59.
Following this line of argumentation leads easily to a semantic vision of the theory of evolution – and not a syntactic one, originally adapted for physical theories (See Thompson 1989). Since the 1960s philosophers have indeed distinguished between two conceptions: the traditional view, the syntactic one, for which sciences can be rendered axiomatically in language of first order logics, relying on semantic rules that allow for the construction of theoretical terms based on observations; and the recent alternative, the “semantic” view initiated by Bas Van Frassen, Patrick Suppes and Frederick Suppe, for which theories are structures defined in a formal language and satisfied by families of mathematical models. The most general statements under the first conception are laws of nature, whereas the second, insofar as it does not have the equivalent of “correspondence rules” between terms of observation and theory, gives no status to the idea of natural law (See Van Fraassen 1980).
- 60.
- 61.
See Delord, Chap. 25, this volume.
- 62.
Population genetics concerns microevolution in time periods that are not very long and with limited environmental variations; macroevolution, on a larger time scale, starts with speciation; and, with variations on an even larger scale (emergence and extinction of clades, etc.) one sometimes talk of megaevolution in the history of life.
- 63.
- 64.
To make it simpler we speak of the altruism allele. In reality the reasoning, like any selectionist reasoning, never implies genetic determinism, which is an absurdity. It is simply enough that possession of the allele A makes a difference for altruism with regard to allele S in a fixed given environment in order for selection to take its effect. One can thus speak of an “altruism gene”, but of course it’s just a way of speaking, not the claim that altruism (or selfishness) is the expression of a given allele.
- 65.
This calculation only works if A is rare in a population.
- 66.
This last expectation explains why the calculation above was only valid if A is rare. In fact, r is approached by kinship relations, but its true value is defined here, so that its measurement is sometimes rather complicated. Grafen (1984) proposes two measurement techniques, and Frank (2006, p. 352) gives a more formal definition. In certain cases the probability that a shares a gene with b is higher than the probability that it is shared with c, even when a and c are relatives in the ordinary sense instead of a and b. In particular when the kinship structures are not as simple as they are in most mammal populations the calculation becomes increasingly complex. The straightforward way of considering r by starting with kinship is sometimes enough, but the most complete definition comes in terms of probabilities; with such a definition many of the controversies surrounding kin selection disappear, as I discuss further on (see also West et al. 2010).
- 67.
In a diploid system of reproduction such as ours, some brothers have 50 % of genes in common, so the probability of having an identical gene to one of mine by randomly choosing one of my brother’s genes is ½. It is easier to understand the degree of genetic relatedness between individuals if it is defined by probability.
- 68.
This is only valid when there is only one queen and when she does not mate with many males; in other cases the explanations are more sophisticated.
- 69.
The monkeys’ warning screams could have many explanations, which differ according to the species and are not exclusive to one; Charnov and Krebs (1975) have demonstrated that the effect of disorder that the shrieks have on the group play to the crying monkey’s advantage, who is less easy to target by the predator thanks to the chaos. In this way the shrieking monkey’s individual fitness also rises.
- 70.
See Sect. 1.
- 71.
Dawkins gives selfish genetic elements as another argument; it is a matter of genic selection in which the organism has nothing to do, thus no controversy can exist. Genic selectionism is an argument concerning selection in general.
- 72.
See Heams, “Heredity”, Chap. 3, this volume.
- 73.
Keller and Ross (1998) first pointed out a “green-beard” effect in nature, with ants. Dawkins rejects the green-beard effect because he thinks it is vulnerable to cheaters who would have the beard without having the altruist gene; but Jansen and Van Baalen (2006) show that in theory, if there are several colored beards, the system remains stable.
- 74.
Price equation is one of the general mathematical formulas of natural selection. I did not include it in the review of principle statements of selection since, though it is no doubt less subject to counter-examples and more rigorous than Hull’s definition of Lewontin’s condition, the equation does assume that the entities in play present heritability and fitness, so the subsequent discussion would be the same as that of Lewontin’s conditions.
- 75.
That is, the variation of a trait between two generations is correlated to the probability of reproduction that the value of the trait confers to the organism who carries it, which is another way of stating the principle of natural selection articulated earlier – for example, the more the tallest ones have the tendency to have more offspring, the more height will rise in subsequent generations and size is under selection.
- 76.
The formal definition of the altruistic act A and selfish act S demands this: A has a cost for X and a benefit for something other than X, S has not cost to X but only a benefit. The cost can be absolute (when the act benefits another while costing the altruist) or relative – when the act benefits the group of n individuals including the altruist himself: she then gets a benefit b/n, but her benefit is smaller than that of the others (b/n-c instead of b/n). Obviously the costs are in fitness, and this altruism is not psychological altruism, (see Clavien, Chap. 34, this volume).
- 77.
This is the basis of Darwin’s explanation of moral sense, see Jérôme Ravat’s, Chap. 35, this volume.
- 78.
In a very close investigation of some of the diverse processes leading to cooperation, Frank (2006) distinguishes between actual kin selection, which explains self-sacrifice that operates in the casts of sterile workers in insects, for example, and the behavioral correlation, which explains cooperation within groups. Whereas there is selective advantage in benefiting from cooperative acts while others remain altruistic when one is in a group, in the second case cooperation benefits the group in general, including the focal individual. Independently of the issue of knowing if these two models perform the same process, Frank thus suggests that they are formally different contrary to Sloan Wilson and Sober’s thesis on the universality of multi-level selection. However, others will say that in all cases, what is causally relevant is the relatedness, which compensates in terms of indirect benefits the cost paid by the focal altruistic individual (West et al. 2010).
- 79.
Here we return again to Hamilton’s rule (West et al. 2007, 423). From this perspective, opposition between two selections at work is a rhetorical artifact, since there is only one single process at work, mathematically speaking.
- 80.
See Huneman (2010b) for an analysis of the involvement of genotypes and of organisms in the controversies over genic selection.
- 81.
Of course, Sober and Wilson are part of the “pluralist” camp in the sense of those that think there are several levels of selection; but “explanatory pluralist” here means believing that there are several possible explanatory frameworks for altruism or cooperation, which is clearly not their case, since they think that the only explanatory process is multi-level selection.
- 82.
- 83.
Some clades persist more than others; if we think that the number of species inside a clade, or its level of branching, or any other property the clade itself has as a clade, contributed to it lasting longer than another clade, then there is clade selection, that is, selection of clades in virtue of clades properties.
- 84.
References
Abrams, M. (2007). How do natural selection and random drift interact? Philosophy of Science, 74(5), 666–679.
Basalla, G. (1988). The evolution of technology. Cambridge: Cambridge University Press.
Beatty, J. (1995). The evolutionary contingency thesis. In G. Wolters & J. G. Lennox (Eds.), Concepts, theories, and rationality in the biological sciences. Pittsburgh: University of Pittsburgh Press.
Beatty, J. (2011). Reconsidering the importance of chance variation. In G. B. Müller & M. Pigliucci Evolution: The extended synthesis. Cambridge: MIT Press, pp. 21–40.
Beatty, J., & Mills, S. (1979). The propensity interpretation of fitness. Philosophy of Science, 46, 263–286.
Bedau, M., & Packard, N. (1998). A classification of long-term evolutionary dynamics. In C. Adami et al. (Eds.), Artificial life VI (pp. 189–198). Cambridge: MIT Press.
Bouchard, F., & Huneman, P. (2013). From groups to individuals. Evolution and emergent individuality. Cambridge: MIT Press.
Bouchard, F., & Rosenberg, A. (2004). Fitness, probability and the principles of natural selection. British Journal for Philosophy of Science, 55, 693–712.
Boyd, R., & Richerson, P. (Eds.). (1985). Culture and the evolutionary process. Chicago: Chicago University Press.
Brandon, R. (1988). The levels of selection: A hierarchy of interactors. In H. C. Plotkin (Ed.), The role of behavior in evolution (pp. 51–71). Cambridge, MA: MIT Press.
Brandon, R. (1990). Adaptation and environment. Princeton: Princeton University Press.
Brandon, R. (1996). Concepts and methods in evolutionary biology. Cambridge, MA: Cambridge University Press.
Brandon, R. (2008). Natural selection. In E. N. Zalta (Ed.), The Stanford encyclopedia of philosophy (Spring 2014 ed.). http://plato.stanford.edu/archives/spr2014/entries/natural-selection/
Brandon, R., & Mc, S. D. (2011). Biology’s first law. Chicago: University of Chicago Press.
Burian, R. M. (1983). Adaptation. In M. Grene (Ed.), Dimensions of Darwinism (pp. 287–314). New York: Cambridge University Press.
Burt, A., & Trivers, R. (2006). The biology of selfish genetics elements. New Haven: Harvard University Press.
Cavalli-Sforza, L., & Feldman, M. (1981). Cultural transmission and evolution. A quantitative approach. Princeton: Princeton University Press.
Charnov, J., & Krebs, E. (1975). The evolution of alarm calls: Altruism or manipulation? American Naturalist, 109(965), 107–112.
Coyne, R., Barton, E., & Turelli, M. (1997). Perspective: A critique of Sewall Wright’s shifting balance theory of evolution. Evolution, 51, 643–671.
Damuth, J., & Heisler, I. L. (1988). Alternative formulations of multi-level selection, selection in structured populations. Biology and Philosophy, 17(4), 477–517.
Danchin, É., Charmantier, A., Champagne, F. A., Mesoudi, A., Pujol, B., & Blanchet, S. (2011). Beyond DNA: Integrating inclusive inheritance into an extended theory of evolution. Nature Review Genetics, 12, 475–486.
Darden, L., & Cain, J. (1989). Selection type theories. Philosophy of Science, 56(1), 106–129.
Darwin, C. (1859). The origin of species. London: John Murray.
Darwin, C. (1999). La filiation de l’homme [1871]. Paris: Syllepse.
Dawkins, R. (1976). The selfish gene. Oxford: Oxford University Press.
Dawkins, R. (1982). The extended phenotype. Oxford: Oxford University Press.
Dennett, D. (1995). Darwin’s dangerous idea. New York: Simons & Shuster.
Doolittle, W. F., & Sapienza, C. (1980). Selfish genes, the phenotype paradigm and genome evolution. Nature, 284(5757), 601–603.
Dretske, F. (1977). Laws of nature. Philosophy of Science, 44, 248–268.
Edwards, A. W. F. (1994). The fundamental theorem of natural selection. Biological Reviews of the Cambridge Philosophical Society, 69(4), 443–474.
Eigen, M. (1983). Self replication and molecular evolution. In D. S. Bendall (Ed.), From molecules to man (pp. 105–130). Cambridge, MA: Cambridge University Press.
Endler, J. (1986). Natural selection in the wild. Princeton: Princeton University Press.
Falconer, D. (1960). Introduction to quantitative genetics. New York: McKay.
Fisher, R. (1930). The genetical theory of natural selection. Oxford: Clarendon.
Frank, S. A. (2006). Social selection. In C. W. Fox & J. B. Wolf (Eds.), Evolutionary genetics: Concepts and case studies (pp. 350–363). Oxford: Oxford University Press.
Frank, S. A., & Slatkin, M. (1992). Fisher’s fundamental theorem of natural selection. Trends in Ecology and Evolution, 7, 92–95.
Gardner, A. (2013). Adaptation of individuals and groups. In F. Bouchard & P. Huneman (Eds.), From groups to individuals. Evolution and emergent individuality (pp. 99–116). Cambridge: MIT Press.
Gavrilets, S. (2011). High dimensional fitness landscapes and speciation. In G. B. Muller & M. Pigliucci (Eds.), Evolution: The extended synthesis (pp. 45–80). Cambridge: MIT Press.
Gayon, J. (1993). La biologie entre loi et histoire. Philosophie, 38, 30–37.
Gayon, J. (1998). Darwinism’s struggle for survival: Heredity and the hypothesis of natural selection (M. Cobb, Trans.). Cambridge, MA: Cambridge University Press.
Gillespie, J. (2004). Population genetics. New York: Oxford University Press.
Glymour, B. (2006). Wayward modeling: Population genetics and natural selection. Philosophy of Science, 2006(73), 369–389.
Godfrey-Smith, P. (2000). The replicator in retrospect. Biology and Philosophy, 15, 403–423.
Godfrey-Smith, P. (2007). Conditions for evolution by natural selection. Journal of Philosophy, 104, 489–516.
Godfrey-Smith, P. (2009). Natural selection and Darwinian populations. New York: Oxford University Press.
Gould, S. J. (1989). Wonderful life. The Burgess shale and the nature of history. New York: Norton.
Gould, S. J., & Lewontin, R. (1979). The spandrels of san Marco and the panglossian paradigm: A critique of the adaptationist programme. Proceedings of the Royal Society of London, B205, 581–598.
Grafen, A. (1984). Natural selection, kin selection and group selection. In J. R. Krebs & N. B. Davies (Eds.), Behavioural ecology (pp. 62–84). Oxford: Blackwell.
Grafen, A. (1990). Biological signals as handicaps. Journal of Theoretical Biology, 144, 517–546.
Grafen, A. (2002). A first formal link between the price equation and an optimization program. Journal of Theoretical Biology, 217, 75–91.
Grafen, A. (2006). Optimisation of inclusive fitness. Journal of Theoretical Biology, 238, 541–563.
Grafen, A. (2009). Formalizing Darwinism and inclusive fitness theory. Philosophical Transactions of the Royal Society B, 364, 3135–3141.
Griesemer, J. (2000). The units of evolutionary transitions. Selection, 1, 67–80.
Haig, D. (2000). The kinship theory of genomic imprinting. Annual Review of Ecology and Systematics, 31, 9–32.
Hamilton, W. D. (1963). The evolution of altruistic behavior. American Naturalist, 97, 354–356.
Holland, J. (1995). Hidden order. How adaptation builds complexity. Readings: Helix.
Hull, D. (1980). Individuality and selection. Annual Review of Ecology System, 11, 311–332.
Huneman, P. (2009). L’individualité biologique et la mort. Philosophie, 1, 63–90.
Huneman, P. (2010). Topological explanations and robustness in biological sciences. Synthese, 177, 213–245.
Huneman, P. (2012). Natural selection: A case for the counterfactual approach. Erkenntnis, 76(2), 171–194.
Huneman, P. (2013). Assessing statistical views of natural selection: Is there a room for non local causation? Studies in History and Philosophy of Biological and Biomedical Sciences, 44, 604–612.
Jansen, A., & Van Baalen, M. (2006). Altruism through beard chromodynamics. Nature, 440, 663–666.
Kauffman, S. (1993). Origins of order: Self-organization and selection in evolution. New York: Oxford University Press.
Keller, E. F. (2001). The century of the gene. Cambridge, MA: Harvard University Press.
Keller, L., & Ross, K. G. (1998). Selfish genes: A green beard in the red fire ant. Nature, 394, 573–575.
Kerr, B., & Godfrey-Smith, P. (2002). Individualist and multi-level perspectives on selection in structured populations. Biology and Philosophy, 17(4), 477–517.
Kerr, B., Godfrey-Smith, P., & Feldman, M. W. (2004). What is altruism? Trends in Ecology and Evolution, 19, 135–140.
Kettlewell, H. B. D. (1955). Selection experiments on industrial melanism in Lepidoptera. Heredity, 9(3):323–342.
Kim, J. (1993). Supervenience and mind. Cambridge: Cambridge University Press.
Kimura, M. (1983). The neutral theory of molecular evolution. Cambridge: Cambridge University Press.
Laeng, B., Mathisen, R., & Johnsen, J. (2007). Why do blue-eyed men prefer women with the same eye color? Behavioral Ecology and Sociobiology, 61(3), 371–384.
Lange, M. (2007). Natural law in scientific practice. New York: Oxford University Press.
Lehmann, L., & Keller, L. (2006). The evolution of cooperation and altruism. A general framework and a classification of models. Journal of Evolutionary Biology, 19, 1365–1376.
Lehmann, L., Ravigne, V., & Keller, L. (2008). Population viscosity can promote the evolution of altruistic sterile helpers and eusociality. Proceedings of the Royal Society of London Series B, 275, 1887–1895.
Lenski, R., & Trevisano, M. (1994). Dynamics of adaptation and diversification: A 10,000-generation experiment with bacterial populations. Proceedings of the National Academy of Sciences of the United States of America, 91, 6808–6814.
Lewens, T. (2010). The natures of selection. British Journal for Philosophy of Science, 61(2), 313–333.
Lewens, T. (2013). Cultural evolution. In E. N. Zalta (Ed.), The Stanford encyclopedia of philosophy (Spring 2013 ed.). http://plato.stanford.edu/archives/spr2013/entries/evolution-cultural/
Lewontin, R. C. (1970). Units of Selection. Annual Review of Ecology and Systematics, 1, 1–18.
Limoges, C. (1977). La sélection naturelle. Paris: Vrin.
Lloyd, E. (2001). Units and levels of selection. An anatomy of the levels of selection debate. In R. Singh, K. Krimbas, D. Paul, & J. Beatty (Eds.), Thinking about evolution: Historical, philosophical and political perspectives (pp. 267–291). Cambridge: Cambridge University Press.
Lumsden, C., & Wilson, E. O. (1981). Genes, minds and culture. Cambridge: Harvard University Press.
Lynch, M. (2007). The origins of the genome architecture. London: Sunderland.
Matthen, M., & Ariew, A. (2002). Two ways of thinking about natural selection. Journal of Philosophy, 49(2), 55–83.
Maynard, S. J. (1978). The evolution of sex. Cambridge: Cambridge University Press.
Maynard, S. J. (1982). Evolution and the theory of games. Cambridge: Cambridge University Press.
Maynard, S. J., & Szathmary, E. (1995). The major transitions in evolution. New York: Freeman.
Mayr, E. (1959a). Typological versus population thinking. In E. Mayr (Ed.), Evolution and the diversity of life (pp. 26–29). Cambridge: Harvard University Press, 1976.
Mayr, E. (1959b). The emergence of evolutionary novelties. In E. Mayr (Ed.), Evolution and the diversity of life (pp. 88–113). Cambridge: Harvard University Press, 1976.
Mayr, E. (1961). Cause and effect in biology. Science, 134, 1501–1506.
Mayr, E. (1965a). Sexual selection and natural selection. In E. Mayr (Ed.), Evolution and the diversity of life (pp. 73–87). Cambridge: Harvard University Press, 1976.
Mayr, E. (1965b). Selection and directional evolution. In E. Mayr (Ed.), Evolution and the diversity of life (pp. 44–52). Cambridge, Harvard University Press, 1976.
Mayr, E. (1984). What is Darwinism today? Proceedings of the Biennial Meeting of the Philosophy of Science Association, 1984, 145–156.
Mayr, E. (1998). The evolutionary synthesis. Cambridge: Harvard University Press.
Mayr, E., & Provine, W. (1980). The evolutionary synthesis. Perspectives on the unification of biology. Cambridge: Harvard University Press.
Medawar, P. (1957). The uniqueness of the individual. London: Methuen.
Michod, R. (1999). Darwinian dynamics. Evolutionary transitions in fitness and individuality. Princeton: Princeton University Press.
Millstein, R. (2006). Natural selection as a population-level causal process. British Journal for the Philosophy of Science, 57, 627–653.
Müller, G., & Pigliucci, M. (2011). Evolution: The extended synthesis. Cambridge: MIT Press.
Neander, K. (1995). Pruning the tree of life. British Journal for Philosophy of Science, 46, 59–80.
Nowak, M. A. (2006). Five rules for the evolution of cooperation. Science, 314, 1560–1563.
Okasha, S. (2006). Evolution and the levels of selection. Oxford: Oxford University Press.
Pál, C., Balázs, P., & Martin, J. (2006). An integrated view of protein evolution. Nature Reviews Genetics, 7, 337–348.
Price, G. R. (1972). Extension of covariance selection mathematics. Annals of Human Genetics, 35(4), 485–490.
Reisman, K., & Forber, P. (2005). Manipulation and the causes of evolution. Philosophy of Science, 72, 1113–1123.
Rice, S. H. (1995). A genetical theory of species selection. Journal of Theoretical Biology, 177, 237–245.
Rosenberg, A. (1995). Instrumental biology, or the disunity of science. Chicago: University of Chicago Press.
Rosenberg, A. (2001). How is biological explanation possible. British Journal for Philosophy of Science, 52, 735–760.
Roughgarden, J. (2006). Reproductive social behavior: Cooperative games to replace sexual selection. Science, 17, 965.
Shoemaker, S. (1984). Causality and properties. In S. Shoemaker (Ed.), Identity, cause and mind: Philosophical essays (pp. 206–233). Oxford: Oxford University Press.
Sober, E. (1980). Evolution, population thinking and essentialism. Philosophy of Science, 3(47), 350–383.
Sober, E. (1984). The nature of selection. Cambridge: MIT Press.
Sober, E., & Lewontin, R. (1982). Artifact, cause and genic selection. Philosophy of Science, 44, 157–180.
Sober, E., & Wilson, D. S. (1998). Unto others: The evolution of altruism. Cambridge: Harvard University Press.
Sterelny, K., & Kitcher, P. (1988). The return of the gene. Journal of Philosophy, 85, 339–360.
Strassmann, J., & Queller, D. (2007). Insect societies as divided organisms : The complexities of purpose and cross-purpose. Proceedings of the National Academy of Sciences USA, 104, 8619–8626.
Taylor, P. D. (1992). Altruism in viscous population: An inclusive fitness model. Evolutionary Ecology, 6, 352–356.
Taylor, T. (2004). Niche construction and the evolution of complexity. In J. Pollack, M. Bedau, P. Husbands, T. Ikegami, & R. Watson (Eds.), Artificial life IX (Proceedings of the Ninth international conference on the simulation and synthesis of living systems) (pp. 375–380). Cambridge: MIT Press.
Tendero, P.-E. (2006). Épistémologie critique de la notion de gène. In F. Athané, É. Machery, & M. Silberstein (dir.), Matière première. Revue d’épistémologie et d’études matérialistes (n° 1/2006) : Nature et naturalisations, Paris: Syllepse.
Thompson, P. (1989). The structure of biological theories. New York: SUNY Press.
Traulsen, A., & Nowak, M. A. (2006). Evolution of cooperation by multilevel selection. Proceedings of the National Academy of Sciences USA, 103, 10952–10955.
Trivers, R. (1971). The evolution of reciprocal altruism. The Quarterly Review of Biology, 46, 35–57.
van Fraassen, B. C. (1980). The scientific image. Oxford: Oxford University Press.
voight, B. F., Kudaravalli, S., Wen, X., & Pritchard, J. K. (2006). A map of recent positive selection in the human genome. PLoS Biology, 4(3), e72.
Wade, M. (1977). An experimental study of group selection. Evolution, 131, 134–153.
Wade, M., & Kalisz, S. (1990). The causes of natural selection. Evolution, 44(8), 1947–1955.
Walsh, D. (2003). Fit and diversity: Explaining adaptive evolution. Philosophy of Science, 70, 280–301.
Walsh, D. (2007). The pomp of superfluous causes: The interpretation of evolutionary theory. Philosophy of Science, 74, 281–303.
Walsh, D., Lewens, T., & Ariew, A. (2002). Trials of life: Natural selection and random drift. Philosophy of Science, 69, 452–473.
Waters, C. K. (1991). Tempered realism about forces of selection. Philosophy of Science, 58, 553–573.
Waters, C. K. (2005). Why genic and multilevel selection are here to stay. Philosophy of Science, 72, 311–333.
West, S. A., Griffin, A. S., & Gardner, A. (2007). Social semantics: Altruism, cooperation, mutualism, strong reciprocity and group selection. Journal of Evolutionary Biology, 20, 415–432.
West, S., Wild, G., & Gardner, A. (2010). The theory of genetic kin selection. Journal of Evolutionary Biology, 24, 1020–1043.
Williams, G. C. (1966). Adaptation and natural selection. Princeton: Princeton University Press.
Williams, G. C. (1975). Sex and evolution. Princeton: Princeton University Press.
Williams, G. C. (1992). Natural selection: Domains, levels and challenges. New York: Oxford University Press.
Wilson, D. S., & Dugatkin, A. L. (1997). Group selection and assortative interactions. The American Naturalist, 149, 336–351.
Wright, S. (1932). The roles of mutation, inbreeding, crossbreeding and selection in evolution. In Proceedings of the sixth annual congress of genetics, 1: (pp. 356–366).
Wynne-Edwards, V. C. (1962). Animal dispersion in relation to social behaviour. Edinburgh: Oliver & Boyd.
Zahavi, A., & Zahavi, A. (1997). The handicap principle. New York: Oxford University Press.
Author information
Authors and Affiliations
Corresponding author
Editor information
Editors and Affiliations
Rights and permissions
Copyright information
© 2015 Springer Science+Business Media Dordrecht
About this chapter
Cite this chapter
Huneman, P. (2015). Selection. In: Heams, T., Huneman, P., Lecointre, G., Silberstein, M. (eds) Handbook of Evolutionary Thinking in the Sciences. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-9014-7_4
Download citation
DOI: https://doi.org/10.1007/978-94-017-9014-7_4
Published:
Publisher Name: Springer, Dordrecht
Print ISBN: 978-94-017-9013-0
Online ISBN: 978-94-017-9014-7
eBook Packages: Biomedical and Life SciencesBiomedical and Life Sciences (R0)