Old wine in new bottles: reaction norms in salmonid fishes

Genetic variability in reaction norms reflects differences in the ability of individuals, populations and ultimately species to respond to environmental change. By increasing our understanding of how genotype × environment interactions influence evolution, studies of genetic variation in phenotypic plasticity serve to refine our capacity to predict how populations will respond to natural and anthropogenic environmental variability, including climate change. Given the extraordinary variability in morphology, behaviour and life history in salmonids, one might anticipate the research milieu on reaction norms in these fishes to be empirically rich and intellectually engaging. Here, I undertake a review of genetic variability in continuous and discontinuous (threshold) norms of reaction in salmonid fishes, as determined primarily (but not exclusively) by common-garden experiments. Although in its infancy from a numerical publication perspective, there is taxonomically broad evidence of genetic differentiation in continuous, threshold and bivariate reaction norms among individuals, families and populations (including inter-population hybrids and backcrosses) for traits as divergent as embryonic development, age and size at maturity, and gene expression. There is compelling inferential evidence that plasticity is heritable and that population differences in reaction norms can reflect adaptive responses, by natural selection, to local environments. As a stimulus for future work, a series of 20 research questions are identified that focus on reaction-norm variability, selection, costs and constraints, demographic and conservation consequences, and genetic markers and correlates of phenotypic plasticity.     

AFLP标记与玉米杂种产量,产量杂种优势的预测

以１７个玉米（Ｚｅａ ｍａｙｓ Ｌ．）自交系及其按双列杂效配制的１３６个单交种为材料,研究ＡＦＬＰ分子标记与玉米杂效种产量、产量杂种优势的关系。结果表明,基于ＡＦＬＰ数据计算的遗传距离与１９９７年杂种产量、产量中优势的相关系数（ｒ）分别为０．４５０３、０．３７１４,与１９９８年杂种产量、产量杂种优势的相关系数分别为０．４３５２、０．３２５３,均达到显水平,但决定系数（ｒ＾２）都很小。当亲本材料改     

High-Altitude Adaptation: Mechanistic Insights from Integrated Genomics and Physiology

Population genomic analyses of high-altitude humans and other vertebrates have identified numerous candidate genes for hypoxia adaptation, and the physiological pathways implicated by such analyses suggest testable hypotheses about underlying mechanisms. Studies of highland natives that integrate genomic data with experimental measures of physiological performance capacities and subordinate traits are revealing associations between genotypes (e.g., hypoxia-inducible factor gene variants) and hypoxia-responsive phenotypes. The subsequent search for causal mechanisms is complicated by the fact that observed genotypic associations with hypoxia-induced phenotypes may reflect second-order consequences of selection-mediated changes in other (unmeasured) traits that are coupled with the focal trait via feedback regulation. Manipulative experiments to decipher circuits of feedback control and patterns of phenotypic integration can help identify causal relationships that underlie observed genotype–phenotype associations. Such experiments are critical for correct inferences about phenotypic targets of selection and mechanisms of adaptation.     

不同光环境下胡桃楸幼苗的形态可塑性及其响应研究

为了探求不同光环境对胡桃楸幼苗生长的影响,对全光照（100%光照）、中光照（60%光照）、低光照（30%光照）3种光环境下胡桃楸幼苗的生长、形态特征、生物量分配和水分特征开展研究。结果表明：胡桃楸幼苗的形态在不同光环境下存在明显的可塑性变化,生物量分配和水分特征均表现出不同程度的差异。低光照条件下,胡桃楸幼苗生长快,株高、地径分别为全光照的1.14、1.18倍;冠幅大,是全光照的1.40倍;叶片宽大,叶长和叶宽分别是全光照的1.25和1.36倍;分枝数明显少于全光照。3种光环境下胡桃楸幼苗地上部分生物量存在显著差异,地下部分差异不显著。叶片相对含水量随着光照的降低呈现逐渐增加的趋势;全光照条件下叶片饱和持水量与叶片干鲜比明显大于遮阴条件;3种光照条件下叶绿素含量差异显著。胡桃楸幼苗为适应不同光环境,在形态和生理方面都做出适应性调整,适当的遮阴处理有利于胡桃楸幼苗生长。     

Genotype–environment interaction and the ontogeny of diet-induced phenotypic plasticity in size and shape of Melanoplus femurrubrum (Orthoptera: Acrididae)

The developmental origin of phenotypic plasticity in morphological shape can be attributed to environment-specific changes in growth of overall body size, localized growth of a morphological structure or a combination of both. I monitored morphological development in the first four nymphal instars of grasshoppers (Melanoplus femurrubrum) raised on two different plant diets to determine the ontogenetic origins of diet-induced phenotypic plasticity and to quantify genetic variation for phenotypic plasticity. I measured diet-induced phenotypic plasticity in body size (tibia length), head size (articular width and mandible depth) and head shape (residual articular width and residual mandible depth) for grasshoppers from 37 full-sib families raised on either a hard plant diet (Lolium perenne) or a soft plant diet (Trifolium repens). By the second to third nymphal instar, grasshoppers raised on a hard plant diet had significantly smaller mean tibia length and greater mean residual articular width (distance between mandibles adjusted for body size) compared with full-sibs raised on a soft plant diet. However, there was no significant phenotypic plasticity in mean unadjusted articular width and mandible depth, and in mean residual mandible depth. At the population level, development of diet-induced phenotypic plasticity in grasshopper head shape is mediated by plastic changes in allocation to tissue growth that maintain growth of head size on hard, low-nutrient diets while reducing growth of body size. Within the population, there was substantial variation in the plasticity of growth trajectories since different full-sib families developed phenotypic plasticity of residual articular width through different combinations of head and body size growth. Genetic variation for diet-induced phenotypic plasticity of residual articular width, residual mandible depth and tibia length, as estimated by genotype–environment interaction, exhibited significant fluctuation through ontogeny (repeated measures MANOVA , family × plant × instar, P < 0.01). For example, there was significant genetic variation for phenotypic plasticity of residual articular width in the third nymphal instar, but not earlier or later in ontogeny. The observed patterns of genetic variation are discussed with reference to short-term constraints and the evolution of phenotypic plasticity.     

Phenotypic plasticity in gene expression contributes to divergence of locally adapted populations of Fundulus heteroclitus

We examine the interaction between phenotypic plasticity and evolutionary adaptation using muscle gene expression levels among populations of the fish Fundulus heteroclitus acclimated to three temperatures. Our analysis reveals shared patterns of phenotypic plasticity due to thermal acclimation as well as non‐neutral patterns of variation among populations adapted to different thermal environments. For the majority of significant differences in gene expression levels, phenotypic plasticity and adaptation operate on different suites of genes. The subset of genes that demonstrate both adaptive differences and phenotypic plasticity, however, exhibit countergradient variation of expression. Thus, expression differences among populations counteract environmental effects, reducing the phenotypic differentiation between populations. Finally, gene‐by‐environment interactions among genes with non‐neutral patterns of expression suggest that the penetrance of adaptive variation depends on the environmental conditions experienced by the individual.     

Phenotypic plasticity and divergence in gene expression

The extent to which phenotypic plasticity, or the ability of a single genotype to produce different phenotypes in different environments, impedes or promotes genetic divergence has been a matter of debate within evolutionary biology for many decades (see, for example, Ghalambor et al. 2007 ; Pfennig et al. 2010 ). Similarly, the role of evolution in shaping phenotypic plasticity remains poorly understood (Pigliucci 2005 ). In this issue of Molecular Ecology, Dayan et al. ( 2015 ) provide empirical data relevant to these questions by assessing the extent of plasticity and divergence in the expression levels of 2272 genes in muscle tissue from killifish (genus Fundulus) exposed to different temperatures. F. heteroclitus (Fig.  1 A) and F. grandis are minnows that inhabit estuarine marshes (Fig.  1 B) along the coasts of the Atlantic Ocean and Gulf of Mexico in North America. These habitats undergo large variations in temperature both daily and seasonally, and these fish are known to demonstrate substantial phenotypic plasticity in response to temperature change (e.g. Fangue et al. 2006 ). Furthermore, the range of F. heteroclitus spans a large latitudinal gradient of temperatures, such that northern populations experience temperatures that are on average ~10°C colder than do southern populations (Schulte 2007 ). By comparing gene expression patterns between populations of these fish from different thermal habitats held in the laboratory at three different temperatures, Dayan et al. ( 2015 ) address two important questions regarding the interacting effects of plasticity and evolution: (i) How does phenotypic plasticity affect adaptive divergence? and (ii) How does adaptive divergence affect plasticity?     

A strong genetic correlation underlying a behavioural syndrome disappears during development because of genotype–age interactions

In animal populations, as in humans, behavioural differences between individuals that are consistent over time and across contexts are considered to reflect personality, and suites of correlated behaviours expressed by individuals are known as behavioural syndromes. Lifelong stability of behavioural syndromes is often assumed, either implicitly or explicitly. Here, we use a quantitative genetic approach to study the developmental stability of a behavioural syndrome in a wild population of blue tits. We find that a behavioural syndrome formed by a strong genetic correlation of two personality traits in nestlings disappears in adults, and we demonstrate that genotype–age interaction is the likely mechanism underlying this change during development. A behavioural syndrome may hence change during organismal development, even when personality traits seem to be strongly physiologically or functionally linked in one age group. We outline how such developmental plasticity has important ramifications for understanding the mechanistic basis as well as the evolutionary consequences of behavioural syndromes.     

Plasticity predicts evolution in a marine alga

Under global change, populations have four possible responses: ‘migrate, acclimate, adapt or die’ (Gienapp et al. 2008 Climate change and evolution: disentangling environmental and genetic response. Mol. Ecol. 17, 167–178. (doi:10.1111/j.1365-294X.2007.03413.x)). The challenge is to predict how much migration, acclimatization or adaptation populations are capable of. We have previously shown that populations from more variable environments are more plastic (Schaum et al. 2013 Variation in plastic responses of a globally distributed picoplankton species to ocean acidification. Nature 3, 298–230. (doi:10.1038/nclimate1774)), and here we use experimental evolution with a marine microbe to learn that plastic responses predict the extent of adaptation in the face of elevated partial pressure of CO₂ (pCO₂). Specifically, plastic populations evolve more, and plastic responses in traits other than growth can predict changes in growth in a marine microbe. The relationship between plasticity and evolution is strongest when populations evolve in fluctuating environments, which favour the evolution and maintenance of plasticity. Strikingly, plasticity predicts the extent, but not direction of phenotypic evolution. The plastic response to elevated pCO₂ in green algae is to increase cell division rates, but the evolutionary response here is to decrease cell division rates over 400 generations until cells are dividing at the same rate their ancestors did in ambient CO₂. Slow-growing cells have higher mitochondrial potential and withstand further environmental change better than faster growing cells. Based on this, we hypothesize that slow growth is adaptive under CO₂ enrichment when associated with the production of higher quality daughter cells.     

披针叶茴香对变化光环境的表型可塑性

植物对变化光环境的表型可塑性大小影响其在林下生境中分布、生长和更新。为探讨披针叶茴香在不同光环境下的整体表型可塑性及其适应机制,采用遮荫试验模拟5种光照条件(100%、52%、33%、15%和6%相对光照强度),研究了不同光环境下披针叶茴香叶片形态、生理、解剖结构、根系形态以及生物量分配等的变化。结果表明:叶生物量在5种光照处理之间差异不显著,但叶面积和比叶面积均随光照强度减弱显著增加。遮荫处理增加了叶绿素a、叶绿素b和类胡萝卜素的含量,但叶绿素a/b比值随光照强度减弱而降低。遮荫降低了非结构性碳水化合物(淀粉和可溶性糖)和可溶性蛋白的含量,增加了叶片氮和磷含量,对叶片氮/磷比影响较小。在52%和33%相对光照处理下,叶片中硝酸盐含量最低,而在100%和6%相对光照处理下硝酸盐积累较多。根生物量、细根和粗根的长度、表面积以及比根长和比根表面积在5种光照处理之间均没有显著差异,根系氮含量在低光环境(15%和6%相对光照处理)中显著降低。随光照强度减弱,披针叶茴香采取保守生存策略,并没有增加叶生物量的分配,而是分配较多的生物量给枝条和树干,储存能量。综合来看,披针叶茴香具有较宽的光生态幅,在6%—100%光照强度下均能正常生长,遮荫有利于披针叶茴香地上和总生物量的积累,52%的相对光照条件下生长最佳。变化光环境下根系性状和整体结构的可塑性相对较低,叶片生理性状的可塑性在披针叶茴香适应光环境变化过程中发挥了主要作用。     

对生玉米产量性状的杂种优势和配合力分析

本文利用对生基因转育获得的对生与互生自交系及其杂交组合,研究了对生性状对产量性状杂种优势和配合力的遗传效应的影响。结果表明:对生玉米同互生玉米一样具有普遍的杂种优势,在产量性状上F1对生株优势互×互组合大于对×对组合和对×互(互×对)组合;对生性状的转育对产量性状的一般配合力有降低效应,而含有对生基因的互生玉米具有较高的一般配合力效应,因此,在对生玉米育种中可以利用含有不同对生基因的互生自交系作亲本来组配杂交组合(互×互),从而获得较高的制种产量和高产的对生杂交种。     

Integrated Environmental and Genomic Analysis Reveals the Drivers of Local Adaptation in African Indigenous Chickens

Breeding for climate resilience is currently an important goal for sustainable livestock production. Local adaptations exhibited by indigenous livestock allow investigating the genetic control of this resilience. Ecological niche modeling (ENM) provides a powerful avenue to identify the main environmental drivers of selection. Here, we applied an integrative approach combining ENM with genome-wide selection signature analyses (XPEHH and Fst) and genotype−environment association (redundancy analysis), with the aim of identifying the genomic signatures of adaptation in African village chickens. By dissecting 34 agro-climatic variables from the ecosystems of 25 Ethiopian village chicken populations, ENM identified six key drivers of environmental challenges: One temperature variable—strongly correlated with elevation, three precipitation variables as proxies for water availability, and two soil/land cover variables as proxies of food availability for foraging chickens. Genome analyses based on whole-genome sequencing (n = 245), identified a few strongly supported genomic regions under selection for environmental challenges related to altitude, temperature, water scarcity, and food availability. These regions harbor several gene clusters including regulatory genes, suggesting a predominantly oligogenic control of environmental adaptation. Few candidate genes detected in relation to heat-stress, indicates likely epigenetic regulation of thermo-tolerance for a domestic species originating from a tropical Asian wild ancestor. These results provide possible explanations for the rapid past adaptation of chickens to diverse African agro-ecologies, while also representing new landmarks for sustainable breeding improvement for climate resilience. We show that the pre-identification of key environmental drivers, followed by genomic investigation, provides a powerful new approach for elucidating adaptation in domestic animals.     

Pricklier with the proper predator? Predator‐induced small‐scale changes of spinescence in Daphnia

Phenotypic plasticity in defensive traits is a common response of prey organisms to variable and unpredictable predation regimes and risks. Cladocerans of the genus Daphnia are keystone species in the food web of lentic freshwater bodies and are well known for their ability to express a large variety of inducible morphological defenses in response to invertebrate and vertebrate predator kairomones. The developed defenses render the daphnids less susceptible to predation. So far, primarily large‐scale morphological defenses, like helmets, crests, and tail‐spines, have been documented. However, less is known on whether the tiny spinules, rather inconspicuous traits which cover many Daphnia’s dorsal and ventral carapace margins, respond to predator kairomones, as well. For this reason, we investigated two Daphnia species (D. magna and D. longicephala) concerning their predator kairomone‐induced changes in dorsal and ventral spinules. Since these small, inconspicuous traits may only act as a defense against predatory invertebrates, with fine‐structured catching apparatuses, and not against vertebrate predators, we exposed them to both, an invertebrate (Triops cancriformis or Notontecta maculata) and a vertebrate predator (Leucaspius delineatus). Our results show that the length of these spinules as well as spinules‐covered areas vary, likely depending on the predator the prey is exposed to. We further present first indications of a Daphnia species‐specific elongation of the spinules and an increase of the spinules‐bearing areas. Although we cannot exclude that spinescence is altered because it is developmentally connected to changes in body shape in general, our results suggest that the inducible alterations to the spinule length and spinules‐covered areas disclose another level of predator‐induced changes in two common Daphnia species. The predator‐induced changes on this level together with the large‐scale and ultrastructural defensive traits may act as the overall morphological defense, adjusted to specific predator regimes in nature.     

表型可塑性与外来植物的入侵能力

外来植物的入侵能力与其性状之间的关系是入侵生态学中的基本问题之一。成功的入侵种常常能占据多样化的生境,并以广幅的环境耐受性为特征。遗传分化(包括生态型分化)和表型可塑性是广布性物种适应变化、异质性生境的两种不同但并不矛盾和排斥的策略。越来越多的实验证据表明,表型可塑性具有确定的遗传基础,本身是一种可以独立进化的性状。许多入侵种遗传多样性比较低,但同时又占据了广阔的地理分布区和多样化的生境,表型可塑性可能在这些物种的入侵成功和随后的扩散中起到了关键作用。本文首先介绍表型可塑性的含义,简述表型可塑性和生物适应的关系,然后从理论分析和实验证据两个方面论述了表型可塑性与外来植物入侵能力的相关性,最后针对进一步的研究工作进行了讨论。当然,并非所有入侵种的成功都能归因于表型可塑性,作者认为对于那些遗传多样性比较低同时又占据多样化生境的入侵种,表型可塑性和入侵能力的正相关可能是一条普遍法则,而非特例。     

对生玉米籽粒品质性状的杂种优势和配合力分析

本文利用对生基因转育获得的对生与互生自交系及其杂交组合,研究了对生性状对玉米主要品质性状的杂种优势和配合力的遗传效应的影响。结果表明:对生自交系籽粒品质性状的配合力效应一般高于互生玉米,在蛋白质含量上,对生F1杂种优势与普通互生F1无明显差异,而在油份和淀粉含量上对生F1较互生表现出明显的优势。不同组配方式对F1对生群体籽粒品质性状的杂种优势存在差异,在对生组合利用中通过互×互组配较对×对组配能使F1获得更高的蛋白质含量;选育含油量较高的对生系,利用对×对组合有助于选育高油杂交种;选育淀粉含量较高的含有不同对生基因的互生自交系作亲本,利用互×互组配有助于选育高淀粉对生杂交组合。     

Genetic and environmental components of growth in nestling blue tits (Parus caeruleus)

We investigated the effect of brood‐size mediated food availability on the genetic and environmental components of nestling growth in the blue tit (Parus caeruleus), using a cross‐fostering technique. We found genetic variation for body size at most nestling ages, and for duration of mass increase, but not of tarsus growth. Hence, nestling growth in our study population seems to have the potential to evolve further. Furthermore, significant genotype–environment interactions indicated heritable variation in reaction norms of growth rates and growth periods, i.e. that our study population had a heritable plasticity in the growth response to environmental conditions. The decreasing phenotypic variance with nestling age indicated compensatory growth in all body traits. Furthermore, the period of weight increase was longer for nestlings growing up in enlarged broods, while there was no difference to reduced broods in the period of tarsus growth. At fledging, birds in enlarged broods had shorter tarsi and lower weights than birds in reduced broods, but there was no difference in wing length or body condition between the two experimental groups. The observed flexibility in nestling growth suggests that growing nestlings are able to respond adaptively to food constraint by protecting the growth of ecologically important traits.