Experimental evolution in fluctuating environments: tolerance measurements at constant temperatures incorrectly predict the ability to tolerate fluctuating temperatures

The ability to predict the consequences of fluctuating environments on species distribution and extinction often relies on determining the tolerances of species or genotypes in different constant environments (i.e. determining tolerance curves). However, very little is known about the suitability of measurements made in constant environments to predict the level of adaptation to rapidly fluctuating environments. To explore this question, we used bacterial clones adapted to constant or fluctuating temperatures and found that measurements across a range of constant temperatures did not indicate any adaptation to fluctuating temperatures. However, adaptation to fluctuating temperatures was only apparent if growth was measured during thermal fluctuation. Thus, tolerance curves based on measurements in constant environments can be misleading in predicting the ability to tolerate fast environmental fluctuations. Such complications could lead to false estimates of the genetic merits of genotypes and extinction risks of species due to climate change‐induced thermal fluctuations.     

Mutations, evolutionary theory and cancer

When searching for mutations that may be responsible for tumourigenesis and interpreting their significance, molecular oncologists often make a number of implicit assumptions about how and why tumour genotypes develop. These assumptions are based on an underlying classical model of tumourigenesis. The classical model has a number of similarities to models of evolution: given the parallels between the growth of tumours and the evolution of whole organisms, this is to be expected. However, consideration of tumourigenesis as an evolutionary process also suggests some modifications that might be made to the classical model. The experimental methods and data analysis of molecular oncology must take full account of the potential contribution of evolutionary theory. As the study of mutations in cancer expands, molecular oncologists are starting to do this.     

Distribution of MR-induced sex-linked recessive lethal mutations in Drosophila melanogaster

In the ‘doubling-dose’ method currently used in genetic risk evaluation, two principle assumptions are made and these are: (1) there is proportionality between spontaneous and induced mutations and (2) the lesions that lead to spontaneous and induced mutations are essentially similar. The studies reported in this paper were directed at examining the validity of these two assumptions in Drosophila. An analysis was made of the distribution of sex-linked recessive lethals induced by MR, one of the well-studied mutator systems in Drosophila.

Appropriate genetic complementation tests with 15 defined X-chromosome duplications showed that MR-induced lethals occurred at many sites along the X-chromosome (in contrast to the known locus specificity of MR-induced visible-mutations); some, but not all these sites at which recessive lethals arose in the MR-system are the same as those known to be hot-spots for X-ray-induced lethals. With in situ hybridization we were able to demonstrate that a majority of MR-induced lethals is associated with a particular mobile DNA sequence, the P-element, i.e. they arose as a result of transposition.

The differences between the profiles of MR-induced and X-ray-induced recessive lethals, and the nature of MR-induced and X-ray-induced mutations, thus raise questions about the validity of the assumptions involved in the use of the ‘doubling-dose’ method.     

Classifying genotypic data from plant breeding trials: a preliminary investigation using repeated checks

Summary Several subjective choices must be made when classifying genotypes based on data from plant breeding trials. One choice involves the method used to weight the contribution each environment makes to the classification. A second involves the use of either genotype-means for each environment or genotypevalues for each block, i.e., considering each block to be a different environment. Another involves whether environments (or blocks) in which genotypes are nonsignificantly different should be included or excluded from such classifications. An alternative to the use of raw or standardized data, is proposed in which each environment is weighted by a discrimination index (DI) that is based on the concept of repeatability. In this study the effect of three weighting methods (raw, standardized and DI), the choice of using environments or blocks, and the choice of including or excluding environments or blocks in which genotypic effects were not significant, were considered in factorial combination to give 12 options. A data set comprised of five check cultivars each repeated six times in each of three blocks at six environments was used. The effect of these options on the ability of a hierarchical clustering technique to correctly classify the repeats into five groups, each consisting of all the six repeats of a particular check cultivar, was investigated. It was found that the DI weighting method generally led to better recovery of the known structure. Using block data rather than environmental data also improved structure recovery for each of the three weighting methods. The exclusive use of environments in which genotypic effects were significant decreased structure recovery while the contrary generally occurred for blocks. The best structure recovery was obtained from the DI weighting applied to blocks (whether genotypes were significant or not).     

Clustering Genotypes Based on Genotype-Environment Interaction When Error Variances Are Heterogeneous

A distance function between any two genotypes based on their interaction with environments, when error variances are heterogeneous is derived. Also a dissimilarity index for a group of genotypes which measures their interaction with environments, as an average of the pairwise distance functions is proposed. An iterative relocation algorithm is used to arrive at two different types of clustering based on their interaction with environments making use of the newly defined dissimilarity index.     

Fluctuating Environments Maintain Genetic Diversity through Neutral Fitness Effects and Balancing Selection

Genetic variation is the raw material upon which selection acts. The majority of environmental conditions change over time and therefore may result in variable selective effects. How temporally fluctuating environments impact the distribution of fitness effects and in turn population diversity is an unresolved question in evolutionary biology. Here, we employed continuous culturing using chemostats to establish environments that switch periodically between different nutrient limitations and compared the dynamics of selection to static conditions. We used the pooled Saccharomyces cerevisiae haploid gene deletion collection as a synthetic model for populations comprising thousands of unique genotypes. Using barcode sequencing, we find that static environments are uniquely characterized by a small number of high-fitness genotypes that rapidly dominate the population leading to dramatic decreases in genetic diversity. By contrast, fluctuating environments are enriched in genotypes with neutral fitness effects and an absence of extreme fitness genotypes contributing to the maintenance of genetic diversity. We also identified a unique class of genotypes whose frequencies oscillate sinusoidally with a period matching the environmental fluctuation. Oscillatory behavior corresponds to large differences in short-term fitness that are not observed across long timescales pointing to the importance of balancing selection in maintaining genetic diversity in fluctuating environments. Our results are consistent with a high degree of environmental specificity in the distribution of fitness effects and the combined effects of reduced and balancing selection in maintaining genetic diversity in the presence of variable selection.     

Distribution of MR-induced sex-linked recessove lethal mutations in Drosophila melanogaster

In the ‘doubling-dose’ method currently used in genetic risk evaluation, two principle assumptions are made and these are: (1) there is proportionality between spontaneous and induced mutations and (2) the lesions that lead to spontaneous and induced mutations are essentially similar. The studies reported in this paper were directed at examining the validity of these two assumptions in Drosophila. An analysis was made of the distribution of sex-linked recessive lethals induced by MR, one of the well-studied mutator systems in Drosophila.Appropriate genetic complementation tests with 15 defined X-chromosome duplications showed that MR-induced lethals occurred at many sites along the X-chromosome (in contrast to the known locus specificity of MR-induced visible-mutations); some, but not all these sites at which recessive lethals arose in the MR-system are the same as those known to be hot-spots for X-ray-induced lethals. With in situ hybridization we were able to demonstrate that a majority of MR-induced lethals is associated with a particular mobile DNA sequence, the P-element, i.e. they arose as a result of transposition.The differences between the profiles of MR-induced and X-ray-induced recessive lethals, and the nature of MR-induced and X-ray-induced mutations, thus raise questions about the validity of the assumptions involved in the use of the ‘doubling-dose’ method.     

The ecology and genetics of fitness in Chlamydomonas. X. The relationship between genetic correlation and genetic distance

Abstract.— A necessary condition for the maintenance of genetic variation in heterogenous environments is that the relative fitnesses of a collection of genotypes vary as conditions of growth change. This can be detected by estimating the amount of gene-by-environment interaction (G X E) when a range of types are tested across a range of conditions. However it is the sign and magnitude of the genetic correlation, which is a component of G X E, that governs the ultimate fate of variation. Whether genetic variation will be preserved, then, depends on how the genetic correlation changes as a function of the ecological differences among environments and the genetic differences among genotypes. To evaluate this, we assayed the performance of 15 chlorophyte species of known genetic relation in 20 environments. We found that the quantity of G X E increased as both the environmental variance across environments and the genetic distance increased. Moreover the genetic correlation declined as the environmental variance between pairs of environments and the genetic distance between pairs of genotypes increased. These results suggest that divergent selection will be more likely to maintain genetic variation when environments are strongly contrasted and genotypes widely divergent.     

Genotype — environment interaction in tissue cultures of birch

Summary In vitro culture experiments were carried out with three birch genotypes characterized by certain genealogical relationships which serve as indicators of genetic similarity or dissimilarity. Each genotype was grown in each of six different environments (medium types), and callus growth and colour were observed. The aim was to improve our understanding of the operation of genetic and environmental effects at the early stages of regeneration in vitro. For this purpose we tried to answer the question as to whether genetic differences exert effects that are consistently distinguishable under different environments or whether environmental differences exert effects that are consistently distinguishable between different genotypes. Since conventional analytical methods, such as the analysis of variance, are inappropriate for providing satisfactory answers to this question, we applied a new concept of interpretation. With the help of this concept we obtained the following results which appear to be unique among their kind. 1) For both characters, callus growth and callus colour, genetic differences are masked only slightly by the environments while environmental differences are almost completely masked by the genotypes. Thus, in the present experiment, interaction is one-sided in the sense that environmental effects interact strongly with genotypic effects but genotypic effects interact only slightly with the environmental ones. 2) Nuclear effects seem to be responsible for the differences observed in callus growth, while the differences in callus colour can be explained by the joint action of nuclear and extranuclear effects.     

Tsetse fly population genetics: an indirect approach to dispersal

Tsetse populations are distributed discontinuously, particularly the morsitans group. Dispersal among diverse populations cannot easily be measured directly because the geographical distances between them can be too great to have a reasonable expectation of recapturing experimentally released flies. Moreover, reproductive success of widely dispersed flies might be poor. The question of dispersal rates in tsetse is immediately important because area-wide eradication plans involving the sterile insect technique are under consideration. Dispersal and gene flow are important from evolutionary and historical viewpoints. An indirect method of estimating dispersal is to measure gene flow. Genetic data indicate surprisingly low rates of gene flow in the morsitans and palpalis groups studied to date. The underlying assumptions in making such estimates need to be examined carefully, however, before accepting firm conclusions, and further research is needed. Of particular interest is the question of tsetse adaptation to local environments.     

Genotype x environment interaction of crossover type: detecting its presence and estimating the crossover point

Genotype-environment interaction (GEI) introduces inconsistency in the relative rating of genotypes across environments and plays a key role in formulating strategies for crop improvement. GEI can be either qualitative (i.e., crossover type) or only quantitative (i.e., non-crossover type). Since the presence of crossover-type interaction has a strong implication for breeding for specific adaptation, it is important to assess the frequency of crossover interactions. This paper presents a test for detecting the presence of crossover-type interaction using the response-environment relationship and enumerates the frequency of crossovers and estimation of the crossover point (CP) on the environment axis, which serves as a cut-off point for the two environments groups where different/specific selections can be made. Sixty-four barley lines with various selection histories were grown in northern Syria and Lebanon giving a total of 21 environments (location-year combinations). Linear regression of the genotypic response on the environmental index represented a satisfactory model, and heterogeneity among regressions was significant. At a 5% level of significance, 38% and 19% of the pairs showed crossover interactions when the error variances were considered heterogeneous and homogeneous, respectively, implying that an appreciable number of crossovers took place in the case of barley lines responding to their environments. The CP of 1.64 t/ha, obtained as the CP of regression lines between the genotype numbers 19 and 31, provided maximum genotype x environment-group interaction. Across all environments, genotype nos. 59 and 12 stood first and second for high yield, respectively. The changes in the ranks of genotypes under the groups of environments can be used for selecting specifically adapted genotypes. Received: 25 January 1999 / Accepted: 16 March 1999     

What do evolutionary researchers believe about human psychology and behavior?

We investigated the prevalence of beliefs in several key and contested aspects of human psychology and behavior in a broad sample of evolutionary-informed scholars (N = 581). Nearly all participants believed that developmental environments substantially shape human adult psychology and behavior, that there are differences in human psychology and behavior based on sex differences from sexual selection, and that there are individual differences in human psychology and behavior resulting from different genotypes. About three-quarters of participants believed that there are population differences from dissimilar ancestral ecologies/environments and within-person differences across the menstrual cycle. Three-fifths believed that the human mind consists of domain-specific, context-sensitive modules. About half of participants believed that behavioral and cognitive aspects of human life history vary along a unified fast-slow continuum. Two-fifths of participants believed that group-level selection has substantially contributed to human evolution. Results indicate that there are both shared core beliefs as well as phenomena that are accepted by varying proportions of scholars. Such patterns represent the views of contemporary scholars and the current state of the field. The degree of acceptance for some phenomena may change over time as evolutionary science advances through the accumulation of empirical evidence.     

Genomic control for association studies

A dense set of single nucleotide polymorphisms (SNP) covering the genome and an efficient method to assess SNP genotypes are expected to be available in the near future. An outstanding question is how to use these technologies efficiently to identify genes affecting liability to complex disorders. To achieve this goal, we propose a statistical method that has several optimal properties: It can be used with case control data and yet, like family-based designs, controls for population heterogeneity; it is insensitive to the usual violations of model assumptions, such as cases failing to be strictly independent; and, by using Bayesian outlier methods, it circumvents the need for Bonferroni correction for multiple tests, leading to better performance in many settings while still constraining risk for false positives. The performance of our genomic control method is quite good for plausible effects of liability genes, which bodes well for future genetic analyses of complex disorders.     

Adult Hippocampal Neurogenesis in Mammals (and Humans): The Death of a Central Dogma in Neuroscience and its Replacement by a New Dogma

The review is a critical appraisal of the history and present status of the phenomenon of adult hippocampal neurogenesis (AHN) in the mammalian and human brain. Previous as well as most current studies of AHN have focused on highly inbred domestic mice and rats that are examined in rigorously controlled laboratory environments using psychology‐based behavioral tests. However, this approach cannot reveal the adaptive significance of AHN, a key unresolved question in the field. After the publication of several thousand articles in the field over the last 20 years, little progress has been made in our understanding of the biological utility of AHN in the real world. To accomplish this will require comparative studies employing a greater diversity of species and species‐specific behaviors that are investigated in a more naturalistic, evolutionary context. Although efforts along these lines are on the rise, they remain “voices in the wilderness.” This review is an attempt to hasten and increase those efforts.     

Contributions to the theory of organic form: The intestinal tract

By making some assumptions concerning the symmetry of certain “classes” of vertebrates and other assumptions concerning the mode of absorption of food by the small intestine, an equation is developed which gives a relation between the length of the small intestine and the total mass of an animal. The equation contains parameters which depend upon the category of vertebrates (carnivorous, omnivorous, etc.) with which the animal is to be associated. The paper also contains a brief discussion of the parameters of the equation and some of the implications which stem from assumptions about their relative magnitudes. In particular, it is noted that the constant of proportionality in the equation which relates intestine length to a power of the body mass is found to be inversely proportional to the square of the “energy content” of the food which makes up the animal's diet. Some suggestions are offered which may lead to an experimental evaluation of the parameters.     

Natural genetic variation in social environment choice: context-dependent gene-environment correlation in Drosophila melanogaster

Gene-environment correlation (rGE) occurs when an individual's genotype determines its choice of environment, generating a correlation between environment and genotype frequency. In particular, social rGE, caused by genetic variation in social environment choice, can critically determine both individual development and the course of social selection. Despite its foundational role in social evolution and developmental psychology theory, natural genetic variation in social environment choice has scarcely been examined empirically. Drosophila melanogaster provides an ideal system for investigating social rGE. Flies live socially in nature and have many opportunities to make social decisions; and natural, heterozygous genotypes may be replicated, enabling comparisons between genotypes across environments. Using this approach, I show that all aspects of social environment choice vary among natural genotypes, demonstrating pervasive social rGE. Surprisingly, genetic variation in group-size preference was density dependent, indicating that the behavioral and evolutionary consequences of rGE may depend on the context in which social decisions are made. These results provide the first detailed investigation of social rGE, and illustrate that that genetic variation may influence organismal performance by specifying the environment in which traits are expressed.     

Proportions of different habitat types are critical to the fate of a resistance allele

We describe a simple deterministic theoretical framework for analysing the gene frequency evolution of two alternative alleles at a single genetic locus in a habitat comprising two environments in which the genotypes have different relative fitnesses. We illustrate this for adaptation of pest insects, where one allele (resistance to toxins expressed in transgenic crops) is favoured in one environment (transgenic plants) and the other allele (susceptibility to toxins) is favoured in the other environment (‘refuges’ of non-transgenic plants). The evolution of allele frequencies depends on selection pressure because of relative sizes of the environments and relative fitnesses of the genotypes in each environment. We demonstrate that there are critical threshold proportions for habitat division that determine equilibrium allele frequencies. The stability of the system depends on relationships between the relative genotype fitnesses. In some cases, the division of the habitat in exactly the threshold proportions removes selection pressure and maintains polymorphism at all allele frequencies.     

Potential gain from selection for yield stability in two grain sorghum populations

Summary Maximum yield under highly unpredictable environments should be associated with selection of genotypes with superior performance across good and poor environments. Several stability parameters have been proposed to identify superior genotypes over a wide range of environments. None of these has been used as selection criteria, however, because of their low heritability. The objective of the study presented here was to compare the relative efficiency of predicted gain from indirect selection among three stability parameters: the coefficient of regression (b), deviation from regression (S _d ² ), and principal components scores (PC) from the AMMI model; two indices including mean yield and a stability parameter; and three indices involving yield at the best, the worst, and an intermediate environment. Two hundred S₁ families from each of two sorghum populations (TP24D and KP9B) were evaluated at four dry-land evironments over 2 years. The low heritability estimates and the low genetic correlation between the various stability parameters and mean yield resulted in their low relative efficiency as indirect selection criteria for high yield across environments. However, when the parameters were combined with mean yield over all to create indices, the relative efficiency increased for all the environments. In terms of resource allocation, these indices were not as efficient as mean productivity, rank summation, and selection index that involved fewer environments in their estimation.Contribution no. 9820 of Agricultural Research Division, Univ. of Neb. and no. 92-203-J of Kansas Exp. Stn.     

Shade induced changes in biomechanical petiole properties in the stoloniferous herb <Emphasis Type="Italic">Trifolium repens</Emphasis>

Increased cell number and cell length both contribute to shade induced elongation of petioles which enables stoloniferous plants to place their leaf lamina higher up in the canopy. Although petiole elongation is assumed to be beneficial, it may also imply costs in terms of decreased biomechanical stability. We test the hypothesis that shade induced elongation changes the biomechanical properties of petioles and that the underlying mechanisms, cell division and cell elongation, differentially affect biomechanical properties. This was done by subjecting 14 genotypes differing in the relative contribution of cell size and cell number to shade induced elongation responses to high light conditions and to simulated canopy shade. Developmental traits (cell size and cell number), morphological traits characterizing the petioles, as well as biomechanical characteristics were measured. Our results show that, comparable to stems of non-clonal plants, the rigidity of a petiole’s tissue (the Young’s modulus) increases, leading to increased flexural stiffness of petioles subjected to shading. Increased flexural stiffness proved to be associated with increased performance under shaded conditions. Our results also indicate that cell number affected the material properties and the flexural stiffness of petioles. However, the degree and pattern of the effects differed between light environments. Shade induced increase in cell number translated into shade induced increase of Young’s modulus and flexural stiffness. Genotypes producing relatively larger cells under shaded conditions experienced a decrease in tissue rigidity. In concert our results indicate that the pattern of selection on flexural stiffness, and thereby also on shade induced changes of cell number and cell size differs among light environments. An erratum to this article can be found at