Natural selection

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Natural Selection ist eine von Valve unabhängige Modifikation des Computerspiels Half-Life. Das Spiel verbindet erstmals Spielelemente eines Ego-Shooters mit denen eines Echtzeit-Strategiespiels. Natural Selection (engl. für natürliche Auslese, kurz NS oder NaSe) ist eine von Valve unabhängige Modifikation des Computerspiels Half-Life. Das Spiel. Natural Selection 2 ist ein Computerspiel des amerikanischen Entwicklers Unknown Worlds Entertainment und die kostenpflichtige Fortsetzung der Mod Natural. Many translated example sentences containing "natural selection" – German-​English dictionary and search engine for German translations. natural selection Bedeutung, Definition natural selection: 1. the process that results in the continued existence of only the types of animals and.

natural selection

natural selection Bedeutung, Definition natural selection: 1. the process that results in the continued existence of only the types of animals and. Natural Selection ist eine von Valve unabhängige Modifikation des Computerspiels Half-Life. Das Spiel verbindet erstmals Spielelemente eines Ego-Shooters mit denen eines Echtzeit-Strategiespiels. Many translated example sentences containing "natural selection" – German-​English dictionary and search engine for German translations.

Natural Selection - Inhaltsverzeichnis

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Version 1. Neither notion, however, is meant to further delineate the circumstances in which selection occurs, or to narrow the scope of application of evolutionary theory for further discussion of these notions, see entry on units and levels of selection.

This is evident, for Hull at least, insofar as genes may be both replicators and interactors The view that evolutionary theory is a theory that applies to active germ-line replicators has come under fire from a multitude of directions.

Genes need not be germ-line to undergo selection, as it is at least arguable that the immune system exhibits selection processes Okasha Copying is beside the point, since only similarity across generations, rather than identity, is necessary for evolutionary change Godfrey-Smith , and entries on units and levels of selection and replication and reproduction.

For his part, Hull seems to agree with this last point, as he allows that organisms might well count as replicators, at least in cases in which they reproduce asexually Hull 28— Despite the bevy of attacks on replicator selectionism, replicator selectionists have not, to my knowledge at least, been criticized for being too permissive and allowing that systems that do not evolve count as undergoing selection.

But germ-line replicators may exert a causal influence on their probability of being copied without spreading in a natural population as a result, as in some cases of frequency-dependent selection of systems already at equilibrium.

In cases of frequency-dependent selection, variant genes cause their own reproduction, but the extent of influence on reproduction is a function of their frequency.

Suppose each type spreads when it is rarer. Because causing replication may not lead to differential replication in these and other cases, replicator selectionists do not effectively take evolution to be necessary for selection while Lewontin and those who follow his basic approach typically do do so.

One natural way to arbitrate the issue of whether systems that undergo selection must evolve is to attend to the point of statements of principles of natural selection, or statements of the requirements for selection.

Many theorists take it that the point of these principles is to set out the scope of a theory in the special sciences that deals with selection and evolution, evolutionary theory.

Lewontin claims that the theory of evolution by natural selection rests on his three principles Equally, Godfrey-Smith claims that statements of conditions for evolution by natural selection exhibit the coherence of evolutionary theory and capture some of its core principles For these writers, the or at least a point of the principles seems to be to capture the domain of application of the theory we have inherited from Darwin.

Darwin would have been surprised to hear that his theory of natural selection was circumscribed so as to apply only to evolving populations.

He himself constructed an explanation of a persistent polymorphism, heterostyly, using his own theory. Plants exhibiting heterostyly develop two, or sometimes three, different forms of flower whose reproductive organisms vary in a number of ways, principally length.

Some plants exhibit different forms of flower on the same plant, while some are dimorphic and trimorphic, with only one sort of flower per plant.

Darwin interpreted the flower variations as conducive to intercrossing, which he thought was beneficial, at least for many organisms.

Populations should not evolve directionally such that a single form of flower spreads throughout the population; instead, multiple variants should be retained, a polymorphism.

Darwin thinks it clear that heterostyly is an adaptation:. The benefit which heterostyled dimorphic plants derive from the existence of the two forms is sufficiently obvious [….

Darwin 30; thanks to Jim Lennox for this reference. Even though the population is not evolving, but instead remaining the same over time, it exhibits an adaptation that consists in this persistent lack of change, an adaptation that Darwin thought explicable using his theory.

These sorts of behaviors result from specific assignments of values for theoretical parameters in many of the very same models that are used to explain simple directional selection where a single variant spreads throughout a population, as in the wolf case discussed in the introduction.

The point is that systems seemingly governed by evolutionary theory exhibit a variety of different sorts of dynamics, and this variety includes both different sorts of evolution, including at least cyclical and directional, as well as a lack of evolution at all, as in cases of stabilizing selection.

Consider in particular how the difference between stabilizing and directional selection in the simplest deterministic models of diploid evolution lies in the value of a single parameter in the genotypic selection model, heterozygote fitness:.

If we hold evolution as a condition for selection, we will issue the curious ruling that a system governed by the first sort of model falls within the scope of evolutionary theory while a system governed by the second sort of model only does so up until it reaches a stable intermediate state but then no longer.

Moreover, populations exhibiting stable polymorphisms resulting from heterozygote superiority, or overdominance, are just one case among many different sorts of systems that equally exhibit stable polymorphisms.

The above models are deterministic, while the dynamics of natural systems are to some extent random. A system governed by both the deterministic equations and the binomial sampling equation is said to undergo drift; all natural systems do so.

For more on drift, effective population size, and randomness in evolutionary theory, see entry on genetic drift.

A system exhibiting heterozygote superiority whose dynamics are a function of the binomial sampling equation will not simply rest at its stable intermediate frequency but will hover around it, in some generations evolving toward it, more rarely evolving away, and in some generations exhibiting no evolution at all.

Which of these cases are cases in which the system undergoes natural selection in the capacious sense?

That is, which cases are cases in which the system falls within the purview of evolutionary theory? A natural answer is all of them.

To answer in this way, however, we must not make evolution necessary for natural selection. This last pattern of argument can be extended.

Indeed, given that every natural system undergoing selection also undergoes drift, evolutionary theory is arguably applicable also to systems that undergo drift even in the absence of selection in the focused sense.

Is the point at which the values equalize so momentous that it marks the point at which systems governed by the equations cease to fall within the purview of one theory and instead fall within the purview of another?

If Brandon is right, then conditions for the application of evolutionary theory must not even include conditions for selection in the focused sense, much less conditions for evolutionary change.

The point of stating conditions for evolution by natural selection need not be to state the conditions of deployment of a particular theory in the special sciences.

Godfrey-Smith mentions that the principles may be important to discussions of extensions of evolutionary principles to new domains.

Statements of the conditions for evolution by natural selection might have value for other reasons.

But evolutionary theory is, despite the name, at least arguably a theory that is applicable to more systems than just those that evolve, as the replicator selectionists would have it.

One of the two chief uses of the notion of natural selection is as an interpretation of one or another quantity in formal models of evolutionary processes; this is the focused sense distinguished above.

Two different quantities are called selection in different formal models widely discussed by philosophers.

This is standard textbook usage Rice ; Hedrick The recursive structure of these models is important.

They can be used to infer how a system will behave into the future though of course only if causes of the variables in the system do not change their values in dynamically-relevant ways that are not explicitly modeled in the recursive equations.

Writers working with type recursion models have developed explicit interpretations of their theoretical terms, including the fitness variables quantifying selection.

So, for instance, Beatty and Millstein defend the view that the fitness coefficients representing selection in type recursions should be understood as modeling a discriminate sampling process, while drift, controlled by effective population size, should be understood as indiscriminate sampling Beatty ; Millstein Philosophers have also contended that particular terms in models of systems featuring the formation of groups or collectives should be understood as quantifying the influence of selection at different levels.

Kerr and Godfrey-Smith discuss one such system of recursions; Jantzen defends an alternative parameterization of group selection as part of different system of equations.

See also Krupp for causal-graphical conceptualization of the notion of group selection. For much more on multi-level selection, see entry on units and levels of selection.

The other formal model of particular interest to philosophers is the Price Equation. The Price Equation represents the extent of evolution in a system with respect to a given trait across a single generation using statistical functions:.

In the Price Equation, selection is associated with the first right-hand side quantity, while the second represents transmission bias.

Identities among algebraic functions of statistical functions make possible the mathematical manipulation of the Price Equation such that it may feature a variety of different quantities.

As with type recursions, quantities in various transformations of the Price Equation are equated with selection at different levels for different systems; Okasha, following Price, treats the covariance of the fitness of collectives with the phenotype of collectives as collective-level selection, while the average of the within-collective covariances between particle character and particular fitness is identified with particle-level selection.

The Price Equation can equally be manipulated to yield distinct notions of inheritance; Bourrat distinguishes temporal, persistence, and generational heritabilities and argues for the temporal notion as appropriate for the purposes of stating conditions for evolution by natural selection Bourrat The distinction between type recursions and the Price Equation is important, because selection is interpreted differently in each.

The two formalisms will issue in different verdicts about whether, and the extent to which, focused selection operates within a single system.

To see this, consider how type recursions are structured such that inferences about dynamics over multiple generations may be made by means of them.

If fitness coefficients in these models quantify selection, and these take fixed values as they do in the genotypic selection model considered above and a great many others , then the extent of selection will remain the same over the time period governed by the model: the fitness variables remain at fixed values so selection remains an unchanging influence.

Consider, for instance, the extent to which the population evolves, according to the genotypic selection model above, when the following values are plugged into the model:.

If we understand selection as quantified by the fitness coefficients in this sort of set-up, then the whole time, selection operates in a constant fashion, since the fitness coefficients remain fixed.

In particular, the operation of selection is the same when the system is evolving toward its stable equilibrium as when it remains at that stable equilibrium.

By contrast, the covariance term in Price Equation model of the system will diminish in value until it reaches zero as the system evolves to its equilibrium state.

When selection is identified with the covariance between type and reproduction, the frequency of the different types matters to the extent of selection.

When selection is identified with fitness variables in type recursions, the frequency of different types has no influence on the extent of selection in the system.

Thus, the different interpretations of selection that correspond to different quantities in different formal models are actually incompatible.

We should expect, then, at least one of these interpretations of selection to fail, since focused selection cannot be two different things at once, at least if what counts as natural selection is non-arbitrary.

One way to reconcile these competing interpretations of selection is to make first right-hand side term in the Price Equation quantify the extent of the influence of selection in a system.

If we assume that focused selection accounts for whatever covariance exists between parental offspring number and phenotype, then we may treat the first right-hand side term of the Price Equation as a measure of the extent of the influence of focused selection, at least at a given type frequency see Okasha This approach puts the logical house in order, allowing for a univocal concept of selection, but it does so at the expense of other commitments.

To note just one, the Price Equation will no longer be causally interpretable, since its quantities may no longer be said to represent causes but instead measure the extents of their influences given further limiting assumptions.

There exists a sizable literature on which of multiple alternative manipulations of the Price Equation represents the actual causal structure of different sorts of system see Okasha and section 5 below for more on this issue.

A substantial debate has arisen over the question of whether what counts as selection is indeed non-arbitrary.

A related issue, discussed in the subsequent section, concerns the causal interpretability of the theory: Advocates of the non-arbitrary character of selection also typically treat selection and drift not only as non-arbitrary quantities, but also as causes, while those who allege that the distinction is arbitrary typically equally challenge the treatment of selection and drift as causes.

When biologically realistic scenarios are discussed, systems of equations for inferring how such systems behave are not made part of the discussion for more on population genetics, see entry on population genetics.

We consider next a case they discuss because it provides a way of contrasting how the contrast between selection and drift is made in type recursions and how it is made in the Price Equation.

There is a sort of arbitrariness here, but it emerges only from analysis of a hypothetical system using population genetics modeling techniques.

In a recent paper, Walsh, Ariew, and Matthen put forward a case of temporally variable selection and claim that it could be treated as a case either of selection or of drift The case is of a discrete-generation system with yearly reproduction in which each of two types of organisms produce different numbers of offspring depending on whether the year is warm or cold, with each type of year being equally probable: the H types produce 6 offspring in warm years while the T types produce 4, and the reverse holds for cold years.

The scenario is illuminating because it involves randomness that cannot be quantified by effective population size in a type recursion but can be quantified as such by the drift parameter in Price Equation.

When deploying type recursions, we must treat cases of temporally variable selection as cases of selection, but we are under no similar constraint when it comes to the Price Equation.

When fitnesses are equal, the frequency of each type is determined solely by the binomial sampling equation above since post-selection frequency, the input to the sampling equation, is just pre-selection frequency.

Such a determination makes next-generation frequency a normal, bell-shaped distribution whose mean is the initial frequency of the types in the system.

The above point generalizes: what is quantifiable as selection and drift in type recursions is made definite by how fitness variables and effective population size function in those models.

The story is different, however, with the Price Equation, owing to how randomness is handled in that formalism.

A version of the Price Equation in which both selection and drift are represented is this Okasha 32 :.

Here, the second term quantifies change due to drift Okasha Nothing about the Price Equation formalism constrains such determinations.

If, for instance, she is ignorant of how the weather changes year to year, she may treat the weather as drift; if she can predict it, she may not do so.

Deployment of the Price Equation is compatible with both treating the weather as contributing to expected fitness and treating it as causing deviation from expectation.

The result is that a theorist deploying the Price Equation may treat as drift that is, quantify as deviation from expectation what a theorist deploying type recursions must treat as selection quantify by fitness variables.

It is possible to make assumptions using the Price Equation such that the drift term quantifies what is quantified by the drift term in type recursions, but nothing about the Price Equation proper forces one to do this, and indeed proponents of the Price Equation, such as Grafen , tout how the drift term in the Price Equation may quantify all sorts of randomness, explicitly including randomness that is not quantifiable as drift in type recursions.

As noted earlier, selection and drift are construed in logically distinct fashions in type recursions and the Price Equation.

Ultimately, the conflict between the two modeling approaches with respect to what counts as selection may be resolvable in at least a couple of ways.

Perhaps one modeling approach is simply wrong about what selection is. Alternatively, something having to do with selection is arbitrary here.

According to this second way of resolving the conflict, the choice between the two modeling approaches is not dictated by nature, and is thus at least metaphysically if not pragmatically arbitrary.

As noted above, evolutionary theory cannot do its job unless it has an explanatory structure. Philosophers have contended selection explains a variety of things in a variety of ways.

There exists a longstanding debate among both scientists and philosophers over exactly what natural selection can explain, one begun by Sober and Neander who were concerned with what natural selection can explain Sober , ; Neander , On the non-creative view, natural selection merely eliminates traits while doing nothing to create new ones; the latter phenomenon is the result of mutation.

Proponents of the creative view see natural selection as a creative force that makes probable combinations of mutations that are necessary for the development of at least some traits.

While Razeto-Barry and Frick grant that natural selection cannot explain the origin of traits that arise by a single mutation, they argue that it can explain the occurrence of sequences of phenotypic changes that would otherwise be wildly unlikely to occur without selection operating to cause the spread of the changes prior to the final one in the sequence.

Additionally, it is disputed whether natural selection can explain why an arbitrary individual has the traits it has Walsh ; Pust ; Birch On the positive view, selection affects the identity of individual organisms, and it is part of the identity of an individual to have been produced by the parents that produced it, so natural selection explains why individuals have the traits they do.

On the negative view, the explanatory scope of natural selection is limited to population level properties. Razeto-Barry and Frick further consider the question of whether natural selection can explain the existence of individuals, ultimately arguing against it.

The capacity for natural selection to explain has come under fire from several directions. Another attack, or set of attacks, on the ability of natural selection to explain have to do with the threat that selectionist explanations are circular.

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