The importance of evolutionary constraints in ecological time scales

Summary The importance of constraints, defined as factors that retard or prevent a population from reaching its immediate adaptive peak on an ecological time scale is analysed. This is done by means of simple quantitative genetic models, which if anything underestimate the importance of constraints. The results show that even in the simplest case the response to selection will not generally be in the same direction as the selection vector, i.e. the direction to the nearest optimum. Adding complexity identifies cases where selection may lead the population in suboptimal directions. It is concluded that information about univariate genetic variances is not sufficient to predict evolutionary responses and may even be misleading. However, genetic covariances are not always acting as constraints, but can under certain circumstances promote evolution towards the nearest optimum. This can be understood by a spectral decomposition of the genetic variance—covariance matrix, where it is shown that the eigenvector associated with the largest amount of variance will to various degrees determine the outcome of selection. A literature survey of the pattern of character covariation in morphological characters in natural populations shows a wide variety of correlation patterns, but quite often shows a high level of covariance between traits. This suggests that constraints to short-term evolution may be more common than generally appreciated.     

THE EVOLUTION OF RESISTANCE TO HERBIVORY IN IPOMOEA PURPUREA. I. ATTEMPTS TO DETECT SELECTION

In this study, we looked for evidence of directional or stabilizing/disruptive selection on plant size and on the level of damage (resistance) caused by four types of herbivores in the annual morning glory Ipomoea purpurea. Selection was estimated by standard phenotypic regression analysis and by regression on breeding values. The phenotypic regression analysis revealed directional selection for all five characters (i.e., plant size and resistance to four types of herbivores) and indicated that plant size and resistance to corn-earworm damage were subject to stabilizing selection. By contrast, the analysis using breeding values revealed directional selection only for plant size and resistance to corn earworms, while none of the characters examined indicated stabilizing or disruptive selection. These results suggest that intermediate levels of damage in I. purpurea are, in general, not maintained by stabilizing selection. Rather, they may reflect either 1) a transient state that exists while directional selection pushes the population toward complete resistance (or, in one case, total absence of resistance) or 2) the evolution of susceptibility to damage by genetic drift.     

August Weismann's concept of germ plasma as the basic reason for the inadequacy of neo-Darwinism

Neo-Darwinism is a result of synthesis of Darwinian concept of natural selection with Weismannian concept of germ plasma. The concept of germ plasma is based on a hypothesis that phenotypic traits are completely determined by genes. Hence, neo-Darwinism describes evolution as a process of alternation of gene frequencies under the effect of natural selection. This is an inadequate approach to the study of evolution. In the course of evolution, genes change their functions, whereas phenotypic characters change their corresponding genes. As a result, every step of evolutionary transformation changes the structure of phenotype-to-genotype correspondence. Therefore, phenotypic evolution cannot be described in genetic terms, the same as to human languages cannot be translated one into another whenever the meaning of words is constantly changing. Consequently, Weismannian germ-plasma concept adequately describes the relation of characters to genes only during stasis, but is inapplicable to evolution.     

Interference Competition Set Limits to the Fundamental Theorem of Natural Selection

The relationship between Fisher's fundamental theorem of natural selection and the ecological environment of density regulation is examined. Using a linear model, it is shown that the theorem holds when density regulation is caused by exploitative competition and that the theorem fails with interference competition. In the latter case the theorem holds only at the limit of zero population density and/or at the limit where the competitively superior individuals cannot monopolise the resource. The results are discussed in relation to population dynamics and life history evolution, where evidence suggests that the level of interference competition in natural populations is so high that the fundamental theorem does not apply.     

Survival strategies of <Emphasis Type="Italic">Escherichia coli</Emphasis> and <Emphasis Type="Italic">Vibrio</Emphasis> spp.: contribution of the viable but nonculturable phenotype to their stress-resistance and persistence in adverse environments

In their natural ecosystems, bacteria are continuously exposed to changing environmental factors including physicochemical parameters (e.g. temperature, pH, etc.), availability of nutrients as well as interaction(s) with other organisms. To increase their tolerance and survival under adverse conditions, bacteria trigger a number of adaptation mechanisms. One of the well-known adaptation responses of the non-spore-forming bacteria is the acquisition of the viable but non-culturable (VBNC) state. This phenotype is induced by different stress factors (e.g. low temperature) and is characterized by the temporal loss of culturability, which can potentially be restored. Moreover, this response can be combined with the bust and boom strategy, which implies the death of the main population of the stressed cells (or their entry into the VBNC state) upon stress, thus enabling the remaining cells (i.e. residual culturable population) to subsist at the expense of the dead or/and VBNC cells. In this review, we discuss the characteristics of the VBNC state, its biological significance and contribution to bacterial survival.     

Indirect Evolution of Hybrid Lethality Due to Linkage with Selected Locus in Mimulus guttatus

Most species are superbly and intricately adapted to the environments in which they live. Adaptive evolution by natural selection is the primary force shaping biological diversity. Differences between closely related species in ecologically selected characters such as habitat preference, reproductive timing, courtship behavior, or pollinator attraction may prevent interbreeding in nature, causing reproductive isolation. But does ecological adaptation cause reproductive incompatibilities such as hybrid sterility or lethality? Although several genes causing hybrid incompatibilities have been identified, there is intense debate over whether the genes that contribute to ecological adaptations also cause hybrid incompatibilities. Thirty years ago, a genetic study of local adaptation to copper mine soils in the wildflower Mimulus guttatus identified a locus that appeared to cause copper tolerance and hybrid lethality in crosses to other populations. But do copper tolerance and hybrid lethality have the same molecular genetic basis? Here we show, using high-resolution genome mapping, that copper tolerance and hybrid lethality are not caused by the same gene but are in fact separately controlled by two tightly linked loci. We further show that selection on the copper tolerance locus indirectly caused the hybrid incompatibility allele to go to high frequency in the copper mine population because of hitchhiking. Our results provide a new twist on Darwin''s original supposition that hybrid incompatibilities evolve as an incidental by-product of ordinary adaptation to the environment.     

植物病原物的群体遗传学

品种单一化、生产密集型和一年多茬的现代农业特点导致病原物呈现出进化速度加快、致病力增强及流行风险增大趋势。深入研究病原物群体遗传学对认识病害的流行、有效选育和使用抗性品种乃至控制病害具有重要意义。文章阐述了植物病原物群体遗传学的研究目标和内容、突变、基因迁移、基因重组、随机遗传漂变和自然选择5大遗传机制在植物病原物进化过程中的作用,以及目前植物病原物群体遗传学研究的现状。     

植物病原物的群体遗传学

品种单一化、生产密集型和一年多茬的现代农业特点导致病原物呈现出进化速度加快、致病力增强及流行风险增大趋势。深入研究病原物群体遗传学对认识病害的流行、有效选育和使用抗性品种乃至控制病害具有重要意义。文章阐述了植物病原物群体遗传学的研究目标和内容、突变、基因迁移、基因重组、随机遗传漂变和自然选择5大遗传机制在植物病原物进化过程中的作用, 以及目前植物病原物群体遗传学研究的现状。     

Polymorphism in multiallelic migration–selection models with dominance

The evolution of the gene frequencies at a single multiallelic locus under the joint action of migration and viability selection with dominance is investigated. The monoecious, diploid population is subdivided into finitely many panmictic colonies that exchange adult migrants independently of genotype. Underdominance and overdominance are excluded. If the degree of dominance is deme independent for every pair of alleles, then under the Levene model, the qualitative evolution of the gene frequencies (i.e., the existence and stability of the equilibria) is the same as without dominance. In particular: (i) the number of demes is a generic upper bound on the number of alleles present at equilibrium; (ii) there exists exactly one stable equilibrium, and it is globally attracting; and (iii) if there exists an internal equilibrium, it is globally asymptotically stable. Analytic examples demonstrate that if either the Levene model does not apply or the degree of dominance is deme dependent, then the above results can fail. A complete global analysis of weak migration and weak selection on a recessive allele in two demes is presented.     

Innate responses of the predatory mite <Emphasis Type="Italic">Phytoseiulus persimilis</Emphasis> to a herbivore-induced plant volatile

The responses of the predatory mite P. persimilis to herbivore-induced plant volatiles are at least partly genetically determined. Thus, there is potential for the evolution of this behaviour by natural selection. We tested whether distinct predator genotypes with contrasting responses to a specific herbivore-induced plant volatile, i.e. methyl salicylate (MeSa), could be found in a base population collected in the field (Sicily). To this end, we imposed purifying selection on individuals within iso-female lines of P. persimilis such that the lines were propagated only via the individual that showed either a preference or avoidance of MeSa. The responses of the lines were characterized as the mean proportion of individuals choosing MeSa when given a choice between MeSa and clean air. Significant variation in predator responses was detected among iso-female lines, thus confirming the presence of a genetic component for this behaviour. Nevertheless, we did not find a significant difference in the response to MeSa between the lines that were selected to avoid MeSa and the lines selected to prefer MeSa. Instead, in the course of selection the lines selected to avoid MeSa shifted their mean response towards a preference for MeSa. An inverse, albeit weaker, shift was detected for the lines selected to prefer MeSa. We discuss the factors that may have caused the apparent lack of a response to selection within iso-female line in this study and propose experimental approaches that address them.     

Natural Selection VS. Random Drift: Evidence from Temporal Variation in Allele Frequencies in Nature

We have obtained monthly samples of two species, Drosophila pseudoobscura and Drosophila persimilis, in a natural population from Napa County, California. In each species, about 300 genes have been assayed by electrophoresis for each of seven enzyme loci in each monthly sample from March 1972 to June 1975. Using statistical methods developed for the purpose, we have examined whether the allele frequencies at different loci vary in a correlated fashion. The methods used do not detect natural selection when it is deterministic (e.g., overdominance or directional selection), but only when alleles at different loci vary simultaneously in response to the same environmental variations. Moreover, only relatively large fitness differences (of the order of 15%) are detectable. We have found strong evidence of correlated allele frequency variation in 13-20% of the cases examined. We interpret this as evidence that natural selection plays a major role in the evolution of protein polymorphisms in nature.     

Genetic differences between populations of sardine, Sardina pilchardus, and anchovy, Engraulis encrasicolus, in the Aegean and Ionian seas

Genetic distances based on electrophoretic variation and multivariate analysis of several morpho-metric and meristic characters suggest that populations of sardine and anchovy from the Aegean and Ionian seas do not form one panmictic population. The distinction between these two classes of populations emerges only as a statistical property, and for some characters the within-sea variation is larger than the between-sea variation. Thus, the reproductive isolation between populations inhabiting the two seas appears to be only partial. Our findings do not support the' pure' or' discrete' stock concept. Instead, they provide evidence for the dynamic population structure model according to which physical (e.g., hydrographic) or biological (e.g., predation or behaviour) factors impose a population structure maintained in a semi-equilibrium state under the opposing influences of migration and selection. Such a dynamic state may not be stable in the long run, and it may not allow for the accumulation of genetic divergence necessary for the emergence of higher taxa. It must, nevertheless, be taken into account in the management of exploited natural populations.     

Elevated temperature increases genome-wide selection on de novo mutations

Adaptation in new environments depends on the amount of genetic variation available for evolution, and the efficacy by which natural selection discriminates among this variation. However, whether some ecological factors reveal more genetic variation, or impose stronger selection pressures than others, is typically not known. Here, we apply the enzyme kinetic theory to show that rising global temperatures are predicted to intensify natural selection throughout the genome by increasing the effects of DNA sequence variation on protein stability. We test this prediction by (i) estimating temperature-dependent fitness effects of induced mutations in seed beetles adapted to ancestral or elevated temperature, and (ii) calculate 100 paired selection estimates on mutations in benign versus stressful environments from unicellular and multicellular organisms. Environmental stress per se did not increase mean selection on de novo mutation, suggesting that the cost of adaptation does not generally increase in new ecological settings to which the organism is maladapted. However, elevated temperature increased the mean strength of selection on genome-wide polymorphism, signified by increases in both mutation load and mutational variance in fitness. These results have important implications for genetic diversity gradients and the rate and repeatability of evolution under climate change.     

Speciation and the evolution of dispersal along environmental gradients

We analyze the joint evolution of an ecological character and of dispersal distance in asexual and sexual populations inhabiting an environmental gradient. Several interesting phenomena resulting from the evolutionary interplay of these characters are revealed. First, asexual and sexual populations exhibit two analogous evolutionary regimes, in which either speciation in the ecological character occurs in conjunction with evolution of short-range dispersal, or dispersal distance remains high and speciation does not occur. Second, transitions between these two regimes qualitatively differ between asexual and sexual populations, with the former showing speciation with long-range dispersal and the latter showing no speciation with short-range dispersal. Third, a phenotypic gradient following the environmental gradient occurs only in the last case, i.e., for non-speciating sexual populations evolving towards short-range dispersal. Fourth, the transition between the evolutionary regimes of long-range dispersal with no speciation and short-range dispersal with speciation is typically abrupt, mediated by a positive feedback between incipient speciation and the evolution of short-range dispersal. Fifth, even though the model of sexual evolution analyzed here does not permit assortative mating preferences, speciation occurs for a surprisingly wide range of conditions. This illustrates that dispersal evolution is a powerful alternative to preference evolution in enabling spatially distributed sexual populations to respond to frequency-dependent disruptive selection.     

The action of stabilizing selection on the perfection of new characters in evolution

In birds, fusion of the carpal elements to a carpometacarpus during morphogenesis takes place during the evolutionary process at different rates, i.e. heterochronically, in different groups (developmental radiations), like the elements of the nasal apparatus in Sauropsida. This is because new characters spread and are pushed back at different rates from the terminal to the initial levels of morphogenesis. The shift is most likely automatic, in that it does not require further mutation, but it is set in motion by selection. Selection does not take effect only when an adequate character (which at first is very imperfect) is accepted; it also causes the character to grow, pushes it back towards the threshold of morphogenesis and thereby increases its perfection and its fitness. The path along which the shift takes place is determined by the morphogenetic route of recapitulation of the ancestral structure and since this movement is caused by protracted stabilizing selection, we can describe it as orthoselective movement. This means that recapitulation is flexible. At the same time, cellular and epigenetic interactions with surrounding structures are reciprocally influenced. The shift continues until adequate adaptation and perfection have been achieved. Stabilizing selection then ceases to act. The shift shows the evolutionary trend or further developmental possibilities. Its chief role is probably in the development of specializations.     

On the evolution of dominance modifiers II: a non-equilibrium approach to the evolution of genetic systems

The evolution of dominance is both the simplest and best investigated example of the evolution of genetic systems. Nevertheless, there exists striking empirical material, e.g. industrial melanism, for which no satisfactory explanation could so far be provided. In this paper we take an approach to this classical problem based on a global analysis together with computer simulations. It reveals that during the evolution of dominance one has to distinguish a "nonequilibrium phase" and a "Fisherian phase". The non-equilibrium phase appears to be characterized by the fact that in general the selection intensity at the primary locus does not affect the degree of modifier selection but only the time necessary for passing through this phase. A further essential conclusion is that modifier evolution only obtains a reasonable amount of efficiency if the population reaches the Fisherian phase already with a high modifier frequency. Using these results, predictions on the population genetic prerequisites for the evolution of dominance are derived. From these we conclude that even in populations in which dominance evolution has occurred it cannot be expected that back-crosses into relics of the ancestral population lead to a breakdown of dominance within a few generations. These predictions are in accordance with empirical data on Biston betularia and Odontopera bidentata.     

Five questions on ecological speciation addressed with individual‐based simulations

We use an individual‐based simulation model to investigate factors influencing progress toward ecological speciation. We find that environmental differences can quickly lead to the evolution of substantial reproductive barriers between a population colonizing a new environment and the ancestral population in the old environment. Natural selection against immigrants and hybrids was a major contributor to this isolation, but the evolution of sexual preference was also important. Increasing dispersal had both positive and negative effects on population size in the new environment and had positive effects on natural selection against immigrants and hybrids. Genetic divergence at unlinked, neutral genetic markers was low, except when environmental differences were large and sexual preference was present. Our results highlight the importance of divergent selection and adaptive divergence for ecological speciation. At the same time, they reveal several interesting nonlinearities in interactions between environmental differences, sexual preference, dispersal and population size.     

Quantitative Aspekte der regressiven Evolution – dargestellt am Beispiel der Augenrückbildung bei Höhlenfischen

Quantitative aspects of regressive evolution – demonstrated by the reduction of the eyes in cave fishes According to our present understanding, the eyes of cave fishes are no longer subject to a stabilizing selection. Because of a strong mutation pressure in the destructive direction, this leads to a continual enrichment of elimination mutants and thereby to a progressive reduction of the eyes after separation of a cave population. Obviously a natural selection process does not play a significant role in the reduction of the eyes in cave fishes. Based on these findings, the degeneration process can be characterized in detail, especially concerning its course and duration.     

Zum konzept der »inneren« selektion: Stellungnahme zu einer evolutionstheoretischen kontroverse

Recently the concept of natural selection in Darwin’s sense has been criticized by some authors. It has been argued that this concept does not explain certain phenomena of evolutionary change, especially in the reach of macroevolution. Some biologists, therefore, demanded for evolution a new model of selection which focuses internal factors in phytogeny. — This paper is a brief discussion of some aspects of “internal” selection and its meaning in contemporary evolutionary biology. The argument of the paper is that evolution can only be explained by a theory taking cognizance of interactions between external and internal selective agencies. Such a theory would be a systems theory of evolution.