Adaptation to an extraordinary environment by evolution of phenotypic plasticity and genetic assimilation

Adaptation to a sudden extreme change in environment, beyond the usual range of background environmental fluctuations, is analysed using a quantitative genetic model of phenotypic plasticity. Generations are discrete, with time lag τ between a critical period for environmental influence on individual development and natural selection on adult phenotypes. The optimum phenotype, and genotypic norms of reaction, are linear functions of the environment. Reaction norm elevation and slope (plasticity) vary among genotypes. Initially, in the average background environment, the character is canalized with minimum genetic and phenotypic variance, and no correlation between reaction norm elevation and slope. The optimal plasticity is proportional to the predictability of environmental fluctuations over time lag τ. During the first generation in the new environment the mean fitness suddenly drops and the mean phenotype jumps towards the new optimum phenotype by plasticity. Subsequent adaptation occurs in two phases. Rapid evolution of increased plasticity allows the mean phenotype to closely approach the new optimum. The new phenotype then undergoes slow genetic assimilation, with reduction in plasticity compensated by genetic evolution of reaction norm elevation in the original environment.     

A conceptual model for the origin of worker behaviour and adaptation of eusociality

In a model based on the wasp family Vespidae, the origin of worker behaviour, which constitutes the eusociality threshold, is not based on relatedness, therefore the origin of eusociality does not depend on inclusive fitness, and workers at the eusociality threshold are not altruistic. Instead, incipient workers and queens behave selfishly and are subject to direct natural selection. Beyond the eusociality threshold, relatedness enables 'soft inheritance' as the framework for initial adaptations of eusociality. At the threshold of irreversibility, queen and worker castes become fixed in advanced eusociality. Transitions from solitary to facultative, facultative to primitive, and primitive to advanced eusociality occur via exaptation, phenotypic accommodation and genetic assimilation. Multilevel selection characterizes the solitary to highly eusocial transition, but components of multilevel selection vary across levels of eusociality. Roles of behavioural flexibility and developmental plasticity in the evolutionary process equal or exceed those of genotype.     

The fitness costs of developmental canalization and plasticity

Organisms are capable of an astonishing repertoire of phenotypic responses to the environment, and these often define important adaptive solutions to heterogeneous and unpredictable conditions. The terms ‘phenotypic plasticity’ and ‘canalization’ indicate whether environmental variation has a large or small effect on the phenotype. The evolution of canalization and plasticity is influenced by optimizing selection‐targeting traits within environments, but inherent fitness costs of plasticity may also be important. We present a meta‐analysis of 27 studies (of 16 species of plant and 7 animals) that have measured selection on the degree of plasticity independent of the characters expressed within environments. Costs of plasticity and canalization were equally frequent and usually mild; large costs were observed only in studies with low sample size. We tested the importance of several covariates, but only the degree of environmental stress was marginally positively related to the cost of plasticity. These findings suggest that costs of plasticity are often weak, and may influence phenotypic evolution only under stressful conditions.     

Selection in a fluctuating environment leads to decreased genetic variation and facilitates the evolution of phenotypic plasticity

Changes in the environment are expected to induce changes in the quantitative genetic variation, which influences the ability of a population to adapt to environmental change. Furthermore, environmental changes are not constant in time, but fluctuate. Here, we investigate the effect of rapid, continuous and/or fluctuating temperature changes in the seed beetle Callosobruchus maculatus, using an evolution experiment followed by a split-brood experiment. In line with expectations, individuals responded in a plastic way and had an overall higher potential to respond to selection after a rapid change in the environment. After selection in an environment with increasing temperature, plasticity remained unchanged (or decreased) and environmental variation decreased, especially when fluctuations were added; these results were unexpected. As expected, the genetic variation decreased after fluctuating selection. Our results suggest that fluctuations in the environment have major impact on the response of a population to environmental change; in a highly variable environment with low predictability, a plastic response might not be beneficial and the response is genetically and environmentally canalized resulting in a low potential to respond to selection and low environmental sensitivity. Interestingly, we found greater variation for phenotypic plasticity after selection, suggesting that the potential for plasticity to evolve is facilitated after exposure to environmental fluctuations. Our study highlights that environmental fluctuations should be considered when investigating the response of a population to environmental change.     

Evolution in heterogeneous environments: Effects of migration on habitat specialization

Summary Richard Levins introduced fitness sets as a tool for investigating evolution within heterogeneous environments. Evolutionary game theory permits a synthesis and generalization of this approach by considering the evolutionary response of organisms to any scale of habitat heterogeneity. As scales of heterogeneity increase from fine to coarse, the evolutionary stable strategy (ESS) switches from a single generalist species to several species that become increasingly specialized on distinct habitats. Depending upon the organisms' ecology, the switch from one to two species may occur at high migration rates (relatively fine-grained environment), or may only occur at very low migration rates (coarse-grained environment). At the ESS, the evolutionary context of a species is the entire landscape, while its ecological context may be a single habitat.Evolution towards the ESS can be represented with adaptive landscapes. In the absence of frequency-dependence, shifting from a single strategy ESS to a two strategy ESS poses the problem of evolving across valleys in the adaptive surface to occupy new peaks (hence, Sewell Wright's shifting balance theory). Frequency-dependent processes facilitate evolution across valleys. If a system with a two strategy ESS is constrained to possess a single strategy, the population may actually evolve a strategy that minimizes fitness. Because the population now rests at the bottom of a valley, evolution by natural selection can drive populations to occupy both peaks.     

核桃楸种源选择试验及其环境因子的影响

收集13个核桃楸种源,在黑龙江省林口和兴隆两个试验点进行播种育苗,调查1年生苗木生长量,进行核桃楸种源选择试验。结果表明,不同种源核桃楸在两地播种后,苗高、基径生长量差异显著,适合林口地区的优良种源为绥阳、和龙、帽儿山和辉南种源;适合兴隆地区的优良种源为抚松、和龙、兴隆和绥阳种源。林口试验点核桃楸种子直径与1年生苗木基径生长量的相关系数达到了显著水平,可以作为反映基径生长量的指标,用线性函数模型来模拟核桃楸种子与苗木的生长情况的相关系数和拟合度相对较高。通过对13个种源地和2个试验点的主要环境因子差异与其1年生苗生长性状差异相关性分析,核桃楸不同种源1年生苗高与纬度和年平均气温等环境因子有一定相关性,且呈负相关,核桃楸种源可适当向北移栽。     

Phenotypic plasticity: linking molecular mechanisms with evolutionary outcomes

We argue that phenotypic plasticity should be broadly construed to encompass a diversity of phenomena spanning several hierarchical levels of organization. Despite seemingly disparate outcomes among different groups of organisms (e.g., the opening/closing of stomata in leaves, adjustments of allocation to growth/reproduction, or the production of different castes in social insects), there are underlying shared processes that initiate these responses. At the most fundamental level, all plastic responses originate at the level of individual cells, which receive and process signals from their environment. The broad variations in physiology, morphology, behavior, etc., that can be produced by a single genotype, can be accounted for by processes regulating gene expression in response to environmental variation. Although evolution of adaptive plasticity may not be possible for some types of environmental signals, in many cases selection has molded responses to environmental variation that generate precise and repeatable patterns of gene expression. We highlight the example of responses of plants to variation in light quality and quantity, mediated via the phytochrome genes. Responses to changes in light at particular stages of plants' life cycles (e.g., seed germination, competition, reproduction) are controlled by different members of this gene family. The mechanistic details of the cell and molecular biology of phytochrome gene action (e.g., their effects on expression of other genes) is outlined. Plasticity of cells and organisms to internal and external environmental signals is pervasive, and represents not just an outcome of evolutionary processes, but also a potentially important molder of them. Phenotypes originally initiated via a plastic response, can be fixed through genetic assimilation as alternate regulatory pathways are shut off. Evolution of mechanisms of plasticity and canalization can both reduce genetic variation, as well as shield it. When the organism encounters novel environmental conditions, this shielded variation may be expressed, revealing hidden reaction norms that represent the raw material for subsequent evolution.     

麦瓶草种群中的花药黑粉菌感染:花的形态学变异、菌孢子散布式样及病原菌-传媒调节性选择

花药黑粉菌(Microbotryum violaceum)可系统侵染其寄主植物麦瓶草(Silene dioica),使其不育而代之以在植物花中布满菌孢子.这些菌孢子主要由花传媒昆虫带到健康植株.花传媒昆虫对花部特征变异的行为反应不仅可能影响花粉的输入/输出率,而且影响菌孢子的着落率和植株的发病与否.为研究S.dioica雌株花部性状特征与花粉传布、菌孢子着落及由此而导致的植株染病之间的相关性,用采自4个不同S.dioica种群的植物进行了移栽实验.该4个种群均位于瑞典北部的Skeppsvik群岛,代表了从健康(岛1,种群1),低度发病(岛2,种群2)到高度发病(岛3、4,种群3、4)的发病梯度.从上述4个不同种群中各采50个植株移栽至岛4的中部.来自健康种群的植株较来自高发病种群的植株具有较大的花,较长的花柱.研究发现,着落在花上的花粉粒数和菌孢子数呈强正相关.来自健康种群的植株每朵花上着落的花粉粒和菌孢子数分别是高发病种群(种群4)植株的4倍和9倍,导致来自4个不同种群的植株的发病率存在着显著差异,种群1、2、3、4的发病率分别为37%、20%、18%、0.在涉及10个种群的田间调查研究中,发现种群中健康植株的平均花柱长度(与花冠大小,胚珠数目正相关)与植株的发病率显著负相关.讨论了病原体-传媒调节的植物花部性状特征选择潜势及由其导致的麦瓶草S. dioica种群间的基因漂移.     

Evolution of phenotypic plasticity: patterns of plasticity and the emergence of ecotypes

Phenotypic plasticity itself evolves, as does any other quantitative trait. A very different question is whether phenotypic plasticity causes evolution or is a major evolutionary mechanism. Existing models of the evolution of phenotypic plasticity cover many of the proposals in the literature about the role of phenotypic plasticity in evolution. I will extend existing models to cover adaptation to a novel environment, the appearance of ecotypes and possible covariation between phenotypic plasticity and mean trait value of ecotypes. Genetic assimilation does not sufficiently explain details of observed patterns. Phenotypic plasticity as a major mechanism for evolution--such as, invading new niches, speciation or macroevolution--has, at present, neither empirical nor model support.     

Selection,structure and the heritability of behaviour

Characters which are closely linked to fitness often have low heritabilities (V_A/V_P). Low heritabilities could be because of low additive genetic variation (V_A), that had been depleted by directional selection. Alternatively, low heritabilities may be caused by large residual variation (V_R=V_P – V_A) compounded at a disproportionately higher rate than V_A across integrated characters. Both hypotheses assume that each component of quantitative variation has an independent effect on heritability. However, V_A and V_R may also covary, in which case differences in heritability cannot be fully explained by the independent effects of elimination‐selection or compounded residual variation. We compared the central tendency of published behavioural heritabilities (mean=0.31, median=0.23) with morphological and life history data collected by 26 ). Average behavioural heritability was not significantly different from average life history heritability, but both were smaller than average morphological heritability. We cross‐classified behavioural traits to test whether variation in heritability was related to selection (dominance, domestic/wild) or variance compounding (integration level). There was a significant three‐way interaction between indices of selection and variance compounding, related to the absence of either effect at the highest integration level. At lower integration levels, high dominance variance indicated effects of selection. It was also indicated by the low CV_A of domestic species. At the same time CV_R increased disproportionately faster than CV_A across integration levels, demonstrating variance compounding. However, neither CV_R nor CV_A had a predominant effect on heritability. The partial regression coefficients of CV_R and CV_A on heritability were similar and a path analysis indicated that their (positive) correlation was also necessary to explain variation in heritability. These results suggest that relationships between additive genetic and residual components of quantitative genetic variation can constrain their independent direct effects on behavioural heritability.     

麦瓶草种群中的花药黑粉菌感染：花的形态学变异、菌孢子散布式样及病原菌—传媒调节性选择

花药黑粉菌（Microbotryum violaceum)可系统侵染其寄主植物麦瓶草（Silene dioica),使其不育而代之以在植物花中布满菌孢子,这些菌孢子主要由花传媒昆虫带到健康植株,花传媒昆虫对花部特征变异的行为反应不仅可能影响花粉的输入/输出率,而且影响菌孢子的着落率和植株的发病与否,为研究S.dioica雌株花部性状特征与花粉传布,菌孢子着落及由此而导致的植株染病之间的相关性,用采自4个不同S.dioica种群的植物进行了移栽实验,讠4个种群均位于瑞典北部的Skeppsvik群岛,代表了从健康（岛1,种群1）,低度发病（岛2,种群2）到高度发病（岛3、4,种群3、4）的发病梯度,从上述4个不同种群中各采50个植株移栽至岛4的中部,来自健康种群的植株较来自高发病种群的植株具有较大的花,较长的花柱,研究发现,着落在花上的花粉粒数和菌孢子数呈强正相关,来自健康种群的植株每朵花上着落的花粉粒和菌孢子数分别是高发病种群（种群4）植株的4倍和9倍,导致来自4个不同种群的植株的发病率存在着显差异。种群1、2、3、4的发病率分别为37%,20%,18%,0。在涉及10个种群的田间调查研究中,发现种群中健康植株的平均花柱长度（与花冠大小,胚珠数目正相关）与植株的发病率显负相关,讨论了病原体-传媒调节的植物花部性状特征选择潜势及其导致的麦瓶草S.dioica种群音质 基因漂移。     

Evolution of maternal care in diploid and haplodiploid populations

Maternal care has been suggested to evolve more readily in haplodiploid populations. Because maternal care appears to have been a prerequisite for the evolution of eusociality, this effect potentially explains the apparent preponderance of haplodiploidy among eusocial taxa. Here, I use a kin selection approach to model the evolution of maternal care in diploid and haplodiploid populations. In contrast to previous suggestions, I find that haplodiploidy may inhibit as well as promote the evolution of maternal care. Moreover, I find that the haplodiploidy effect vanishes in outbred populations if gene effects average rather than add together. I confirm these analytical results using numerical simulation of an explicit population genetics model. This analysis casts doubt upon the idea that haplodiploidy has promoted the evolution of maternal care and, consequently, the evolution of eusociality.     

Finding the Factors of Reduced Genetic Diversity on X Chromosomes of Macaca fascicularis: Male-Driven Evolution,Demography, and Natural Selection

The ratio of genetic diversity on X chromosomes relative to autosomes in organisms with XX/XY sex chromosomes could provide fundamental insight into the process of genome evolution. Here we report this ratio for 24 cynomolgus monkeys (Macaca fascicularis) originating in Indonesia, Malaysia, and the Philippines. The average X/A diversity ratios in these samples was 0.34 and 0.20 in the Indonesian–Malaysian and Philippine populations, respectively, considerably lower than the null expectation of 0.75. A Philippine population supposed to derive from an ancestral population by founding events showed a significantly lower ratio than the parental population, suggesting a demographic effect for the reduction. Taking sex-specific mutation rate bias and demographic effect into account, expected X/A diversity ratios generated by computer simulations roughly agreed with the observed data in the intergenic regions. In contrast, silent sites in genic regions on X chromosomes showed strong reduction in genetic diversity and the observed X/A diversity ratio in the genic regions cannot be explained by mutation rate bias and demography, indicating that natural selection also reduces the level of polymorphism near genes. Whole-genome analysis of a female cynomolgus monkey also supported the notion of stronger reduction of genetic diversity near genes on the X chromosome.     

Modifying effects of phenotypic plasticity on interactions among natural selection, adaptation and gene flow

Divergent natural selection, adaptive divergence and gene flow may interact in a number of ways. Recent studies have focused on the balance between selection and gene flow in natural populations, and empirical work has shown that gene flow can constrain adaptive divergence, and that divergent selection can constrain gene flow. A caveat is that phenotypic diversification may be under the direct influence of environmental factors (i.e. it may be due to phenotypic plasticity), in addition to partial genetic influence. In this case, phenotypic divergence may occur between populations despite high gene flow that imposes a constraint on genetic divergence. Plasticity may dampen the effects of natural selection by allowing individuals to rapidly adapt phenotypically to new conditions, thus slowing adaptive genetic divergence. On the other hand, plasticity may promote future adaptive divergence by allowing populations to persist in novel environments. Plasticity may promote gene flow between selective regimes by allowing dispersers to adapt to alternate conditions, or high gene flow may result in the selection for increased plasticity. Here I expand frameworks for understanding relationships among selection, adaptation and gene flow to include the effects of phenotypic plasticity in natural populations, and highlight its importance in evolutionary diversification.     

Effects of dissolved organic matter from sewage sludge on the atrazine sorption by soils

The effects of dissolved organic matter (DOM), water soluble organic matter derived from sewage sludge, on the sorption of atrazine (2-chloro-4-ethylamino-6-isopropylamino-1,3,5-trazine) by soils were studied using a batch equilibrium technique. Six paddy soils, chosen so as to have different organic carbon contents, were experimented in this investigation. Atrazine sorption isotherms on soils were described by the linear equation, and the distribution coefficients without DOM (Kd) or with DOM (Kd*) were obtained. Generally, the values of Kd*/Kd initially increased and decreased thereafter with increasing DOM concentrations of 0-60 mg DOC ·L-1 in soil-solution system form. Critical concentrations of DOM (DOMnp) were obtained where the value of Kd* was equal to Kd. The presence of DOM with concentrations lower than DOMnp promoted atrazine sorption on soils (Kd* > Kd), whereas the presence of DOM with concentrations higher than DOMnp tended to inhibit atrazine sorption (Kd* < Kd). Interestingly, DOMnp for     

Effects of dissolved organic matter from sewage sludge on the atrazine sorption by soils

The effects of dissolved organic matter (DOM), water soluble organic matter derived from sewage sludge, on the sorption of atrazine (2-chloro-4-ethylamino-6-isopropylamino-1,3,5-trazine) by soils were studied using a batch equilibrium technique. Six paddy soils, chosen so as to have different organic carbon contents, were experimented in this investigation. Atrazine sorption isotherms on soils were described by the linear equation, and the distribution coefficients without DOM (Kd) or with DOM (Kd*) were obtained. Generally, the values of Kd*/Kd initially insoil-solution system form. Critical concentrations of DOM (DOMnp) were obtained where the value of Kd* was equal to Kd. The presence of DOM with concentrations lower than DOMnp promoted atrazine sorption on soils (Kd* ＞ Kd), whereas the presence of DOM with concentrations higher than DOMnp tended to inhibit atrazine sorption (Kd* ＜ Kd). Interestingly, DOMnp for tested soils was negatively correlated to the soil organic carbon content, and the maximum of Kd*/Kd (i.e.Kmax) correlated positively with the maximum of DOM sorption on soil (Xmax). Further investigations showed that the presence of hydrophobic fraction of DOM evidently promoted the atrazine sorption on soils, whereas the presence of hydrophilic DOM fraction obviously tended to inhibit the atrazine sorption. Interactions of soil surfaces with DOM and its fractions were suggested to be the major processes determining atrazine sorption on soils. The results of this work provide a reference to the agricultural use of organic amendment such as sewage sludge for improving the availability of atrazine in soils.     

Phenotypic plasticity and the possible role of genetic assimilation: Hypoxia-induced trade-offs in the morphological traits of an African cichlid

In this study we investigate the possible role of phenotypic plasticity and genetic assimilation in the process of adaptation and evolutionary change in the cichlid Pseudocrenilabrus multicolor victoriae . In the field we compared a population of a stable hypoxic habitat with one of a stable well-oxygenated habitat. In the laboratory, we compared individuals from the same mother raised under hypoxic or well-oxygenated conditions to examine phenotypic plasticity. Morphological parameters of three categories were measured: (a) the gill apparatus, (b) the surrounding structural elements, and (c) the outer shape of the fish. Swamp-dwelling fish had a 29% greater total gill surface area than fish from the well-oxygenated habitat due to their larger gill filament length and greater lamellar area. In the plasticity experiment, total gill surface area was 18% greater in the hypoxia group due to a larger number of longer filaments. Surrounding elements and outer shape also differed between the field populations and between fish grown under hypoxic and well-oxygenated conditions, but there was disparity between the field results and the plasticity experiment. The disparity between field and experimental fish may be due to: (a) differences in selection pressures between populations, (b) different constraints for genetic and plasticity changes, or (c) selection against plastic responses to hypoxia. Our results suggest that both (a) and (c) are involved.     

Science and the Concept of Evolution: From the Big Bang to the Origin and Evolution of Life

The common thread of evolution runs through all science disciplines, and the concept of evolution enables students to better understand the nature of the universe and our origins. “Science and the Concept of Evolution” is one of two interdisciplinary science Core courses taken by Dowling College undergraduates as part of their General Education requirements. The course examines basic principles and methods of science by following the concept of evolution from the big bang to the origin and evolution of life. Case studies of leading scientists illustrate how their ideas developed and contributed to the evolution of our understanding of the world. Evidences for physical, chemical, and biological evolution are explored, and students learn to view the evolution of matter and of ideas as a natural process of change over space and time.