Does selection intensity increase when populations decrease? Absolute fitness, relative fitness, and the opportunity for selection

The variance in relative fitness, commonly called the “opportunity for selection,” is a measure of the maximum amount of selection that can occur in a population. I review the relation between fitness variance and population growth, showing that fitness variance is higher during periods of population decline. This is true both for survival and for commonly used models for variable descendant number (Poisson, negative binomial, generalized Poisson). Empirical evidence suggests that not just the opportunity for selection but also the actual selection occurring is commonly greater during such periods of population reduction.     

Pollen selection — past,present and future

 A series of studies, recently reviewed, has established that approximately 60% of the structural genes which are expressed in the sporophytic portion of the angiosperm life cycle are also expressed and exposed to selection in the pollen. Given the haploidy and large population sizes of pollen grains, a substantial portion of the sporophytic genome could thus be periodically exposed to a bacterial type of mass screening. This extraordinary possibility is often subject to some skepticism which may, of course, be justified. However, recent attempts to apply models appear to be inappropriate in this context, in part because these attempts overlook an important source of genetic variation, and also because they assume fixed values for selection and fitness. More recently, studies of pollen/pollen interactions have suggested that what Linskens termed the ”programic phase” may represent an arena for important, and largely unexplored phenomena, some of which are discussed here. Received: 3 June 1996 / Revision accepted: 26 July 1996     

Variable selection on Eurosta's gall size,III: Can an evolutionary response to selection be detected?

Standard quantitative genetic theory predicts that when a trait is exposed to selection, the between-generation change in the phenotypic mean, Δz?_i, will be equal to the product of the trait's heritability and the selection differential, h²S. By extension, this theory implies that if a number of replicate populations are exposed to varying intensities of selection, the between-generation changes in means should covary with the selection differential applied. This relationship offers an opportunity for a statistical test to detect evolutionary change when selection is measured in replicate populations. If an evolutionary response to phenotypic selection occurs, the regression of over S_i, where i indicates population, will have a positive slope. This statistical test was applied to data on the insect Eurosta solidaginis (Diptera: Tephritidae). The larvae of this fly induce galls on the stems of the host plant, Solidago altissima (Asteraceae). Previous work has shown that gall size is a heritable trait of the insect. Further, size-dependent attack on Eurosta larvae by parasitoids selects for larger gall size (Weis and Abrahamson, 1986). Long-term data on phenotypic selection in 16 populations across 5 generations were analyzed for selection response. Apparent upward evolutionary responses were seen in 2 of the 4 between-generation transitions. However, no response was seen when the analysis was applied to the cumulative change in gall size. Examination of the data suggested that some of the change in mean gall size was a developmental response to spatial and temporal variation in the environment. Non-linear developmental effects of environment, when combined with non-linear fitness functions, can induce a spurious selection response; these non-linear relationships can account for the apparent evolutionary change gall size found in the by-generation analysis. Thus, there is no reliable evidence for evolutionary change in Eurosta's gall size over the generations studied. Stasis of gall size in the face of ongoing selection may be due to counterbalancing selection on the gallmaker imposed by host plant resistance.     

K-selection, α-selection, effectiveness, and tolerance in competition: Density-dependent selection revisited

In the Drosophila literature, selection for faster development and selection for adapting to high density are often confounded, leading, for example, to the expectation that selection for faster development should also lead to higher competitive ability. At the same time, results from experimental studies on evolution at high density do not agree with many of the predictions from classical density-dependent selection theory. We put together a number of theoretical and empirical results from the literature, and some new experimental results on Drosophila populations successfully subjected to selection for faster development, to argue for a broader interpretation of density-dependent selection. We show that incorporating notions of alpha-selection, and the division of competitive ability into effectiveness and tolerance components, into the concept of density-dependent selection yields a formulation that allows for a better understanding of the empirical results. We also use this broader formulation to predict that selection for faster development in Drosophila should, in fact, lead to the correlated evolution of decreased competitive ability, even though it does lead to the evolution of greater efficiency and higher population growth rates at high density when in monotypic culture.     

Cereal phytochromes: targets of selection, targets for manipulation?

Plants respond to shading through an adaptive syndrome termed shade avoidance. In high-density crop plantings, shade avoidance generally increases extension growth at the expense of yield and can be at odds with the agronomic performance of the crop as a whole. Studies in Arabidopsis are beginning to reveal the essential role phytochromes play in regulating this process and to identify genes underlying the response. In this article, we focus on how phytochrome signaling networks have been targeted in cereal breeding programs in the past and discuss the potential to alter these pathways through breeding and transgenic manipulation to develop crops that perform better under typical high density conditions.     

Phenotypic selection in natural populations: what limits directional selection?

Studies of phenotypic selection document directional selection in many natural populations. What factors reduce total directional selection and the cumulative evolutionary responses to selection? We combine two data sets for phenotypic selection, representing more than 4,600 distinct estimates of selection from 143 studies, to evaluate the potential roles of fitness trade-offs, indirect (correlated) selection, temporally varying selection, and stabilizing selection for reducing net directional selection and cumulative responses to selection. We detected little evidence that trade-offs among different fitness components reduced total directional selection in most study systems. Comparisons of selection gradients and selection differentials suggest that correlated selection frequently reduced total selection on size but not on other types of traits. The direction of selection on a trait often changes over time in many temporally replicated studies, but these fluctuations have limited impact in reducing cumulative directional selection in most study systems. Analyses of quadratic selection gradients indicated stabilizing selection on body size in at least some studies but provided little evidence that stabilizing selection is more common than disruptive selection for most traits or study systems. Our analyses provide little evidence that fitness trade-offs, correlated selection, or stabilizing selection strongly constrains the directional selection reported for most quantitative traits.     

Trait selection in flowering plants: how does sexual selection contribute?

By highlighting and merging the frameworks of sexual selectionenvisioned by Arnold (1994) and Murphy (1998), we discuss howsexual selection can occur in plants even though individualsdo not directly interact. We review studies on traits that influencepollen export and receipt in a variety of hermaphroditic andgynodioecious plants with the underlying premise that pollinationdynamics influences mate acquisition. Most of the studies reviewedfound that phenotypes that enhance pollen export are in harmonywith those that enhance pollen receipt suggesting that in manycases pollinator visitation rates limit both male and femalefunction. In contrast, fewer traits were under opposing selection;but when they were, the traits most often were associated withenhancing the specific aspects of a given sex function. Ourreview helps clarify and illustrate why sexual selection canbe a component of trait evolution in hermaphrodite plants.     

What is hierarchical selection?

It has been proposed that natural selection occurs on a hierarchy of levels, of which the organismic level is neither the top nor the bottom. This hypothesis leads to the following practical problem: in general, how does one tell if a given phenomenon is a result of selection on level X or level Y. How does one tell what the units of selection actually are? It is convenient to assume that a unit of selection may be defined as a type of entity for which there exists, among all entities on the same “level” as that entity, an additive component of variance for some specific component F of fitness which does not appear as an additive component of variance in any decomposition of this F among entities at any lower level. But such a definition implicitly assumes that if f(x, y) depends nonadditively on its arguments, there must be interaction between the quantities which x and y represent. This assumption is incorrect. And one cannot avoid this error by speaking of “transformability to additivity” instead of merely “additivity”. A general mathematical formulation of the concepts of interaction and non-interaction is proposed, followed by a correspondingly modified approach to the definition of a unit of selection. The practical difficulty of verifying the presence of hierarchical selection is discussed.     

Paradoxes of tumour complexity: somatic selection,vulnerability by design,or infectious aetiology?

The aetiology of cancer involves intricate cellular and molecular mechanisms that apparently emerge on the short timescale of a single lifetime. Some of these traits are remarkable not only for their complexity, but also because it is hard to conceive selection pressures that would favour their evolution within the local competitive microenvironment of the tumour. Examples include ‘niche construction’ (re‐programming of tumour‐specific target sites) to create permissive conditions for distant metastases; long‐range feedback loops of tumour growth; and remarkably ‘plastic’ phenotypes (e.g. density‐dependent dispersal) associated with metastatic cancer. These traits, which we term ‘paradoxical tumour traits’, facilitate the long‐range spread or long‐term persistence of the tumours, but offer no apparent benefit, and might even incur costs in the competition of clones within the tumour. We discuss three possible scenarios for the origin of these characters: somatic selection driven by specific selection regimes; non‐adaptive emergence due to inherent vulnerabilities in the organism; and manipulation by putative transmissible agents that contribute to and benefit from these traits. Our work highlights a lack of understanding of some aspects of tumour development, and offers alternative hypotheses that might guide further research.     

Teleological reasoning,not acceptance of evolution,impacts students’ ability to learn natural selection

Background

How acceptance of evolution relates to understanding of evolution remains controversial despite decades of research. It even remains unclear whether cultural/attitudinal factors or cognitive factors have a greater impact on student ability to learn evolutionary biology. This study examined the influence of cultural/attitudinal factors (religiosity, acceptance of evolution, and parents’ attitudes towards evolution) and cognitive factors (teleological reasoning and prior understanding of natural selection) on students’ learning of natural selection over a semester-long undergraduate course in evolutionary medicine.

Method

Pre-post course surveys measured cognitive factors, including teleological reasoning and prior understanding of natural selection, and also cultural/attitudinal factors, including acceptance of evolution, parent attitudes towards evolution, and religiosity. We analyzed how these measures influenced increased understanding of natural selection over the semester.

Results

After controlling for other related variables, parent attitude towards evolution and religiosity predicted students’ acceptance of evolution, but did not predict students’ learning gains of natural selection over the semester. Conversely, lower levels of teleological reasoning predicted learning gains in understanding natural selection over the course, but did not predict students’ acceptance of evolution.

Conclusions

Acceptance of evolution did not predict students’ ability to learn natural selection over a semester in an evolutionary medicine course. However, teleological reasoning did impact students’ ability to learn natural selection.

     

Kin selection in disguise?

In a recent article E.O. Wilson and B. H?lldobler (2005) describe an heuristic model for the evolution of eusociality. They present their model as an alternative to the standard model of kin selection, and describe the evolution of eusociality in terms of changes in frequency to an hypothetical eusocial allele. Here I build on sentiments of Foster et al. (2006) to suggest that the proposed model is not a clear alternative to the standard model, but appears to represent a special case of kin selection involving preferential interactions among individuals sharing the same altruistic gene. The model proposed by Wilson and H?lldobler is consistent with the ‘greenbeard’ model of kin selection, first proposed by W.D. Hamilton. Received 23 May 2006; revised 27 June 2006; accepted 5 July 2006.     

Why don't zebras have machine guns? Adaptation, selection, and constraints in evolutionary theory

In an influential paper, Stephen Jay Gould and Richard Lewontin (1979) contrasted selection-driven adaptation with phylogenetic, architectural, and developmental constraints as distinct causes of phenotypic evolution. In subsequent publications Gould (e.g., 1997a,b, 2002) has elaborated this distinction into one between a narrow "Darwinian Fundamentalist" emphasis on "external functionalist" processes, and a more inclusive "pluralist" emphasis on "internal structuralist" principles. Although theoretical integration of functionalist and structuralist explanations is the ultimate aim, natural selection and internal constraints are treated as distinct causes of evolutionary change. This distinction is now routinely taken for granted in the literature in evolutionary biology. I argue that this distinction is problematic because the effects attributed to non-selective constraints are more parsimoniously explained as the ordinary effects of selection itself. Although it may still be a useful shorthand to speak of phylogenetic, architectural, and developmental constraints on phenotypic evolution, it is important to understand that such "constraints" do not constitute an alternative set of causes of evolutionary change. The result of this analysis is a clearer understanding of the relationship between adaptation, selection and constraints as explanatory concepts in evolutionary theory.     

Species richness among birds: body size,life history,sexual selection or ecology?

Why do some avian families contain so many more species than other families? We use comparisons between sister taxa to test predictions arising from six explanations to this puzzle: that differences between families are due to chance, body size, life history, sexual selection, intrinsic ecological factors or extrinsic abiotic factors, respectively. In agreement with previous analyses, we find no support for the idea that differences in species richness are simply due to chance. However, contrary to most previous work, we also find no support for the hypotheses that high species richness is correlated with small body size and fast life history. Rather, high species diversity is strongly associated with pronounced plumage dichromatism, generalist feeding habits and good dispersal capabilities as well as large and fragmented geographical ranges. In addition, all of these relationships are robust to the removal of the two most speciose avian lineages, the Ciconiiformes and the Passeriformes. The supposed relationships between species richness and both body size and life history are, however, due to phylogenetic non-independence. Together with previous work showing that differences between avian lineages in extinction risk are associated with variation in body size and life history, these results indicate that extinction rates and speciation rates are not necessarily determined by the same factors. Hence, high extinction rates are not inevitably associated with low speciation rates. Extinction-prone lineages may, in fact, have a high rate of speciation. In such lineages a high proportion of ''vulnerable'' species would be a natural, ongoing phenomenon.     

Species richness in agamid lizards: chance,body size,sexual selection or ecology?

Why does species richness vary so greatly across lineages? Traditionally, variation in species richness has been attributed to deterministic processes, although it is equally plausible that it may result from purely stochastic processes. We show that, based on the best available phylogenetic hypothesis, the pattern of cladogenesis among agamid lizards is not consistent with a random model, with some lineages having more species, and others fewer species, than expected by chance. We then use phylogenetic comparative methods to test six types of deterministic explanation for variation in species richness: body size, life history, sexual selection, ecological generalism, range size and latitude. Of eight variables we tested, only sexual size dimorphism and sexual dichromatism predicted species richness. Increases in species richness are associated with increases in sexual dichromatism but reductions in sexual size dimorphism. Consistent with recent comparative studies, we find no evidence that species richness is associated with small body size or high fecundity. Equally, we find no evidence that species richness covaries with ecological generalism, latitude or range size.     

How strong is natural selection?

The strength of selection in nature has long been a controversial subject, partly because there were few quantitative measurements of phenotypic selection available until recently. In a new paper, Kingsolver and colleagues reviewed 63 studies and found that the median standardized directional selection gradient (a measure of the strength of phenotypic selection) was 0.16. Whether this means selection in nature is strong or weak depends both on one's point of view and on the error in selection estimates.