Adaptive Population Differentiation Research Articles

Developmental bias predicts 60 million years of wing shape evolution

The degree to which developmental biases affect trait evolution is subject to much debate. Here, we first quantify fluctuating asymmetry as a measure of developmental variability, i.e., the propensity of developmental systems to create some phenotypic variants more often than others, and show that it predicts phenotypic and standing genetic variation as well as deep macroevolutionary divergence in wing shape in sepsid flies. Comparing our data to the findings of a previous study demonstrates that developmental variability in the sepsid fly Sepsis punctum strongly aligns with mutational, standing genetic, and macroevolutionary variation in the Drosophilidae--a group that diverged from the sepsid lineage ca. 64 My ago. We also find that developmental bias in S. punctum wing shape aligns with the effects of allometry, but less so with putatively adaptive thermal plasticity and population differentiation along latitude. Our findings demonstrate that developmental bias in fly wings predicts evolvability and macroevolutionary trajectories on a much greater scale than previously appreciated but also suggest that causal explanations for such alignments may go beyond simple constraint hypotheses.

2b-RAD Genotyping of the Seagrass Cymodocea nodosa Along a Latitudinal Cline Identifies Candidate Genes for Environmental Adaptation.

Plant populations distributed along broad latitudinal gradients often show patterns of clinal variation in genotype and phenotype. Differences in photoperiod and temperature cues across latitudes influence major phenological events, such as timing of flowering or seed dormancy. Here, we used an array of 4,941 SNPs derived from 2b-RAD genotyping to characterize population differentiation and levels of genetic and genotypic diversity of three populations of the seagrass Cymodocea nodosa along a latitudinal gradient extending across the Atlantic-Mediterranean boundary (i.e., Gran Canaria—Canary Islands, Faro—Portugal, and Ebro Delta—Spain). Our main goal was to search for potential outlier loci that could underlie adaptive differentiation of populations across the latitudinal distribution of the species. We hypothesized that such polymorphisms could be related to variation in photoperiod-temperature regime occurring across latitudes. The three populations were clearly differentiated and exhibited diverse levels of clonality and genetic diversity. Cymodocea nodosa from the Mediterranean displayed the highest genotypic richness, while the Portuguese population had the highest clonality values. Gran Canaria exhibited the lowest genetic diversity (as observed heterozygosity). Nine SNPs were reliably identified as outliers across the three sites by two different methods (i.e., BayeScan and pcadapt), and three SNPs could be associated to specific protein-coding genes by screening available C. nodosa transcriptomes. Two SNPs-carrying contigs encoded for transcription factors, while the other one encoded for an enzyme specifically involved in the regulation of flowering time, namely Lysine-specific histone demethylase 1 homolog 2. When analyzing biological processes enriched within the whole dataset of outlier SNPs identified by at least one method, “regulation of transcription” and “signalling” were among the most represented. Our results highlight the fundamental importance signal integration and gene-regulatory networks, as well as epigenetic regulation via DNA (de)methylation, could have for enabling adaptation of seagrass populations along environmental gradients.

Crossbreeding reveals the scale and genetic architecture of local adaptation in a predominantly selfing Hordeum spontaneum

ABSTRACT Environmental variation can be large across a wide range of spatial scales resulting in complex patterns of local adaptation across species ranges. We analyzed the scale, genetic mechanism and direct climatic causes of local adaptation in a widely distributed grass Hordeum spontaneum. We performed artificial crosses of maternal plants representing the same Negev desert population with plants originating elsewhere. Pollen donors were plants from other Negev desert populations, non-desert Israeli populations sampled along an aridity gradient, and accessions covering the entire species range. Our study included planting of inter-population hybrids under favorable and simulated desert experimental conditions, followed by analysis of their performance, variation in adaptive traits and relationship with climatic parameters at sampling locations. The combined results of parental phenotypic variation and performance of hybrids were consistent with local selection, reflecting the importance of both regional and local climates. The adaptive genetic differentiation of barley desert populations had a complex architecture. None of the three effects (additive, dominance and epistasis) were fully responsible for this differentiation. Although genetic effects not related to extrinsic selection appear to contribute to genetic differentiation in barley, epistatic effects arising from local selection clearly predominated. The short-term effect of gene flow by pollen was generally negative, indicating that a majority of the new allele combinations created by recombination were maladaptive. However, the long-term effect of occasional pollen flow from other desert populations appears to be positive, as some new recombined genotypes were superior in fitness to the maternal plants even in the F2 generation.

Morphology of tulips (Tulipa, Liliaceae) in its primary centre of diversity

Environmental variation can be large across a wide range of spatial scales resulting in complex patterns of local adaptation across species ranges. We analyzed the scale, genetic mechanism and direct climatic causes of local adaptation in a widely distributed grass Hordeum spontaneum. We performed artificial crosses of maternal plants representing the same Negev desert population with plants originating elsewhere. Pollen donors were plants from other Negev desert populations, non-desert Israeli populations sampled along an aridity gradient, and accessions covering the entire species range. Our study included planting of inter-population hybrids under favorable and simulated desert experimental conditions, followed by analysis of their performance, variation in adaptive traits and relationship with climatic parameters at sampling locations. The combined results of parental phenotypic variation and performance of hybrids were consistent with local selection, reflecting the importance of both regional and local climates. The adaptive genetic differentiation of barley desert populations had a complex architecture. None of the three effects (additive, dominance and epistasis) were fully responsible for this differentiation. Although genetic effects not related to extrinsic selection appear to contribute to genetic differentiation in barley, epistatic effects arising from local selection clearly predominated. The short-term effect of gene flow by pollen was generally negative, indicating that a majority of the new allele combinations created by recombination were maladaptive. However, the long-term effect of occasional pollen flow from other desert populations appears to be positive, as some new recombined genotypes were superior in fitness to the maternal plants even in the F2 generation.

Manipulation of trait expression and pollination regime reveals the adaptive significance of spur length

Understanding the mechanisms of adaptive population differentiation requires that both the functional and adaptive significance of divergent traits are characterized in contrasting environments. Here, we (a) determined the effects of floral spur length on pollen removal and receipt using plants with artificial spurs representing the species-wide variation in length, and (b) quantified pollinator-mediated selection on spur length and three traits contributing to floral display in two populations each of the short-spurred and the long-spurred ecotype of the orchid Platanthera bifolia. Both pollen receipt and removal reached a maximum at 28-29mm long spurs in a short-spurred population visited by short-tongued moths. In contrast, pollen receipt increased linearly across the tested range (4-52mm) and pollen removal was unrelated to spur length in a long-spurred population predominantly visited by a long-tongued moth. The experimentally documented effects on pollen transfer were not reflected in pollinator-mediated selection through female fitness or pollen removal indicating that the natural within-population variation in spur length was insufficient to result in detectable variation in pollen limitation. Our study illustrates how combining trait manipulation with analysis of causes and strength of phenotypic selection can illuminate the functional and adaptive significance of trait expression when trait variation is limited.

Effects of marker type and filtering criteria on Q ST-F ST comparisons.

Comparative studies of quantitative and neutral genetic differentiation (QST-FST tests) provide means to detect adaptive population differentiation. However, QST-FST tests can be overly liberal if the markers used deflate FST below its expectation, or overly conservative if methodological biases lead to inflated FST estimates. We investigated how marker type and filtering criteria for marker selection influence QST-FST comparisons through their effects on FST using simulations and empirical data on over 18 000 in silico genotyped microsatellites and 3.8 million single-locus polymorphism (SNP) loci from four populations of nine-spined sticklebacks (Pungitius pungitius). Empirical and simulated data revealed that FST decreased with increasing marker variability, and was generally higher with SNPs than with microsatellites. The estimated baseline FST levels were also sensitive to filtering criteria for SNPs: both minor alleles and linkage disequilibrium (LD) pruning influenced FST estimation, as did marker ascertainment. However, in the case of stickleback data used here where QST is high, the choice of marker type, their genomic location, ascertainment and filtering made little difference to outcomes of QST-FST tests. Nevertheless, we recommend that QST-FST tests using microsatellites should discard the most variable loci, and those using SNPs should pay attention to marker ascertainment and properly account for LD before filtering SNPs. This may be especially important when level of quantitative trait differentiation is low and levels of neutral differentiation high.

Adaptive differentiation among populations of the Mediterranean dry grassland species Brachypodium retusum: The role of soil conditions, grazing, and humidity.

Genetic differentiation in plant species may result from adaptation to environmental conditions, but also from stochastic processes. The drivers selecting for local adaptation and the contribution of adaptation to genetic differentiation are often unknown. Restoration and succession studies have revealed different colonization patterns for Brachypodium retusum, a common Mediterranean grass. In order to understand these patterns, we tested population differentiation and adaptation to different environmental factors. Structured sampling of 12 populations from six sites and two soil types within site was used to analyze the spatial and environmental structure of population differentiation. Sampling sites differ in grazing intensity and climate. We tested germination and growth in a common garden. In subsets, we analyzed the differential response to stone cover, grazing and soil moisture. We found significant differences among populations. The site explained population differentiation better than soil, suggesting a dominant influence of climate and/or genetic drift. Stone cover had a positive influence on seedling establishment, and populations showed a differential response. However, this response was not related to environmental differences between collection sites. Regrowth after clipping was higher in populations from the more intensively grazed Red Mediterranean soils suggesting an adaptation to grazing. Final germination was generally high even under drought, but germination response to differences in soil moisture was similar across populations. Adaptive population differentiation in germination and early growth may have contributed to different colonization patterns. Thus, the provenance of B. retusum needs to be carefully considered in ecological restoration.

Adaptive genetic variability and differentiation of Croatian and Austrian Quercus robur L. populations at a drought prone field trial

Provenance trials, where populations of different geographical origin are tested in a common environment (common garden test), are a tool suited to allow the study of intraspecific adaptive genetic variation. Research of pedunculate oak (Quercus robur L.) adaptive genetic variability through analyses of populations in common garden tests has a long tradition. However, pedunculated oak populations originating south-eastern from the Alps have been scarcely studied in this way. This study addresses the adaptive genetic variability and differentiation of pedunculate oak populations originating from Austria and Croatia in a provenance/progeny field trial. Studied plants were six years old and were growing at the trial for three years. After two years of unusually low precipitations height and survival were analysed. The total mean height of all plants in the trial was 137.8 cm and ranged from 123.0 cm to 151.8 cm. The overall mean survival rate was rather high (0.85). Mean population survival ranged from 0.64 to 0.94. Individual narrow-sense heritabilities (hi2), family mean heritabilities (hf2), the coefficients of additive genetic variation (CVA) and quantitative genetic differentiation coefficients (QST) were calculated. A multivariate regression tree (MRT) analysis was used to determine the pattern of genetic differentiation of the populations. Individual heritabilities for height ranged between 0.00 and 0.39. Family mean heritabilities for height were rather low in most populations as well (<0.5). Family mean heritabilities for survival were higher than for height (ranging between 0.00 and 0.77). Calculated QST coefficients (0.25 for height and 0.14 for survival) indicated between-population genetic differentiation. The populations were separated into two clusters by MRT analysis regarding a climatic variable, namely Hargreaves’ reference evapotranspiration. Populations originating from comparatively more humid habitats were grouped in the first cluster. The first cluster had a lower mean height and survival compared to the second one. The differences between these clusters were highly statistically significant. The observed quantitative genetic differentiation might have been driven by natural selection caused by differences in the relative moisture of the habitats from which the progeny populations originate. The results suggest ecotypic pattern of the quantitative genetic differentiation among studied populations.

Evidence of correlated evolution and adaptive differentiation of stem and leaf functional traits in the herbaceous genus, Helianthus.

Patterns of plant stem traits are expected to align with a "fast-slow" plant economic spectrum across taxa. Although broad patterns support such tradeoffs in field studies, tests of hypothesized correlated trait evolution and adaptive differentiation are more robust when taxa relatedness and environment are taken into consideration. Here we test for correlated evolution of stem and leaf traits and their adaptive differentiation across environments in the herbaceous genus, Helianthus. Stem and leaf traits of 14 species of Helianthus (28 populations) were assessed in a common garden greenhouse study. Phylogenetically independent contrasts were used to test for evidence of correlated evolution of stem hydraulic and biomechanical properties, correlated evolution of stem and leaf traits, and adaptive differentiation associated with source habitat environments. Among stem traits, there was evidence for correlated evolution of some hydraulic and biomechanical properties, supporting an expected tradeoff between stem theoretical hydraulic efficiency and resistance to bending stress. Population differentiation for suites of stem and leaf traits was found to be consistent with a "fast-slow" resource-use axis for traits related to water transport and use. Associations of population traits with source habitat characteristics supported repeated evolution of a resource-acquisitive "drought-escape" strategy in arid environments. This study provides evidence of correlated evolution of stem and leaf traits consistent with the fast-slow spectrum of trait combinations related to water transport and use along the stem-to-leaf pathway. Correlations of traits with source habitat characteristics further indicate that the correlated evolution is associated, at least in part, with adaptive differentiation of Helianthus populations among native habitats differing in climate.

Phenotypic differentiation of Streptococcus pyogenes populations is induced by recombination-driven gene-specific sweeps.

Genomic recombination plays an important role in driving adaptive evolution and population differentiation in bacteria. However, controversy exists as to the effects of recombination on population diversity and differentiation, i.e., recombination is frequent enough to sweep through the population at selected gene loci (gene-specific sweeps), or the recombination rate is low without interfering genome-wide selective sweeps. Observations supporting either view are sparse. Pathogenic bacteria causing infectious diseases are promising candidates to provide observations of recombination. However, phenotype-associated differentiations are usually vague among them due to diverse disease manifestations. Here we report a population genomic study of the group A Streptococcus pyogenes (GAS), a human pathogen with highly recombining genomes. By employing a genome-wide association study on single nucleotide polymorphisms (SNPs), we demonstrate a phenotypic differentiation of GAS, represented by separate clustering of two sublineages associated with niche-specific infections, i.e., skin infection and pharyngitis-induced acute rheumatic fever. By quantifying SNPs associated with the differentiation in a statistical and phylogenetic context, we propose that the phenotype-associated differentiation arose through recombination-driven gene-specific sweeps, rather than genome-wide sweeps. Our work provides a novel paradigm of phenotype-associated differentiation induced by gene-specific sweeps in a human pathogen and has implications for understanding of driving forces of bacterial evolution.

Adaptive flexibility of enzymatic antioxidants SOD, APX and CAT to high light stress: The clonal perennial monocot Iris pumila as a study case

High solar radiation has been recognized as one of the main causes of the overproduction of reactive oxygen species (ROS) and oxidative stress in plants. To remove the excess of ROS, plants use different antioxidants and tune their activity and/or isoform number as required for given light conditions. In this study, the adaptiveness of light-induced variation in the activities and isoform patterns of key enzymatic antioxidants SOD, APX and CAT was tested in leaves of Iris pumila clonal plants from two natural populations inhabiting a sun exposed dune site and a forest understory, using a reciprocal-transplant experiment. At the exposed habitat, the mean enzymatic activity of total SODs was significantly greater than that in the shaded one, while the amount of the mitochondrial MnSOD was notably higher compared to the plastidic Cu/ZnSOD. However, the number of Cu/ZnSOD isoforms was greater in the forest understory relative to the exposed site (three vs. two, respectively). An inverse relationship recorded between the quantities of MnSOD and Cu/ZnSOD in alternative light habitats might indicate that the two enzymes compensate each other in maintaining intracellular ROS and redox balance. The adaptive population differentiation in APX activity was exclusively recorded in the open habitat, which indicated that the synergistic effect of high light and temperature stress could be the principal selective factor, rather than high light alone. The enzymatic activity of CAT was similar between the two populations, implicating APX as the primary H2O2 scavenger in the I. pumila leaves exposed to high light intensity.

Beyond fruit-flies: population genomic advances in non-Drosophila arthropods.

Understanding the evolutionary processes driving the adaptive differentiation of populations is of broad interest in biology. Genome-wide nucleotide polymorphisms provide the basis for population genetic studies powered by advances in high-throughput sequencing technologies. These advances have led to an extension of genome projects to a variety of non-genetic model organisms, broadening our view on the evolution of gene families and taxonomic-restricted novelties. Here, we review the progress of genome projects in non-Drosophila arthropods, focusing on advances in the analysis of large-scale polymorphism data and functional genomics and examples of population genomic studies.

Population size, habitat fragmentation, and the nature of adaptive variation in a stream fish.

Whether and how habitat fragmentation and population size jointly affect adaptive genetic variation and adaptive population differentiation are largely unexplored. Owing to pronounced genetic drift, small, fragmented populations are thought to exhibit reduced adaptive genetic variation relative to large populations. Yet fragmentation is known to increase variability within and among habitats as population size decreases. Such variability might instead favour the maintenance of adaptive polymorphisms and/or generate more variability in adaptive differentiation at smaller population size. We investigated these alternative hypotheses by analysing coding-gene, single-nucleotide polymorphisms associated with different biological functions in fragmented brook trout populations of variable sizes. Putative adaptive differentiation was greater between small and large populations or among small populations than among large populations. These trends were stronger for genetic population size measures than demographic ones and were present despite pronounced drift in small populations. Our results suggest that fragmentation affects natural selection and that the changes elicited in the adaptive genetic composition and differentiation of fragmented populations vary with population size. By generating more variable evolutionary responses, the alteration of selective pressures during habitat fragmentation may affect future population persistence independently of, and perhaps long before, the effects of demographic and genetic stochasticity are manifest.

Conservation genetics of tilapias: Seeking to define appropriate units for management

Tilapias comprise 112 species and subspecies of cichlid fishes of the genera Oreochromis, Sarotherodon, and Tilapia. Because of the importance of tilapias to aquatic ecosystem structure and function, fisheries, and aquaculture, it is critically important to conserve wild genetic resources. Of the 112 recognized tilapia taxa, 41 are regarded as imperiled. In order to manage adaptively important genetic variation in tilapias, we need to recognize and define biologically appropriate units of conservation. An evolutionary significant unit (ESU) can be defined as a population or group of populations that merits priority for conservation and separate management because of high genetic and ecological distinctiveness from other such units. Management units (MUs) are defined as populations that are demographically independent of one another; that is, their population dynamics depend mostly on local birth and death rates, and not on genetically effective migration. Identification of MUs – similar to “stocks” widely referred to in fisheries management – is useful for short-term management, such as managing habitat, delineating fishing areas, setting harvest rates, and monitoring population status. Against this background, the goal of our review and synthesis was to summarize knowledge and recommend critical work yet to be done regarding conservation of tilapias, approaching this task in a species-by-species manner. With the exceptions of Oreochromis niloticus, O. mossambicus, and Sarotherodon melanotheron, the body of existing work regarding genetic and adaptive differentiation of populations of tilapias is insufficient to inform identification of evolutionary significant units, a knowledge gap that should be addressed by targeted research. We note that competition and interspecific hybridization caused by introductions of tilapias for aquaculture purposes jeopardizes the genetic resources of certain native tilapia species. We hope that our review and synthesis spark critical discussion of tilapia conservation within the tilapia aquaculture, management, and conservation communities.

Genetic control of flowering in spotted gum, Corymbia citriodora subsp. variegata and C. maculata

Genetically controlled asynchrony in anthesis is an effective barrier to gene flow between planted and native forests. We investigated the degree of genetically controlled variation in the timing of key floral developmental stages in a major plantation species in subtropical Australia, Corymbia citriodora subsp. variegata K.D. Hill and L.A.S Johnson, and its relative C. maculata K.D. Hill and L.A.S. Johnson. Flowering observations were made in a common garden planting at Bonalbo in northern New South Wales in spring on 1855 trees from eight regions over three consecutive years, and monthly on a subset of 208 trees for 12 months. Peak anthesis time was stable over years and observations from translocated trees tended to be congruent with the observations in native stands, suggesting strong genetic control of anthesis time. A cluster of early flowering provenances was identified from the north-east of the Great Dividing Range. The recognition of a distinct flowering race from this region accorded well with earlier evidence of adaptive differentiation of populations from this region and geographically-structured genetic groupings in C. citriodora subsp. variegata. The early flowering northern race was more fecund, probably associated with its disease tolerance and greater vigour. Bud abundance fluctuated extensively at the regional level across 3 years suggesting bud abundance was more environmentally labile than timing of anthesis. Overall the level of flowering in the planted stand (age 12 years) was low (8–12% of assessed trees with open flowers), and was far lower than in nearby native stands. Low levels of flowering and asynchrony in peak anthesis between flowering races of C. citriodora subsp. variegata may partially mitigate a high likelihood of gene flow where the northern race is planted in the south of the species range neighbouring native stands.

BRINGING HABITAT INFORMATION INTO STATISTICAL TESTS OF LOCAL ADAPTATION IN QUANTITATIVE TRAITS: A CASE STUDY OF NINE-SPINED STICKLEBACKS

Detection of footprints of historical natural selection on quantitative traits in cross-sectional data sets is challenging, especially when the number of populations to be compared is small and the populations are subject to strong random genetic drift. We extend a recent Bayesian multivariate approach to differentiate between selective and neutral causes of population differentiation by the inclusion of habitat information. The extended framework allows one to test for signals of selection in two ways: by comparing the patterns of population differentiation in quantitative traits and in neutral loci, and by comparing the similarity of habitats and phenotypes. We illustrate the framework using data on variation of eight morphological and behavioral traits among four populations of nine-spined sticklebacks (Pungitius pungitius). In spite of the strong signal of genetic drift in the study system (average FST = 0.35 in neutral markers), strong footprints of adaptive population differentiation were uncovered both in morphological and behavioral traits. The results give quantitative support for earlier qualitative assessments, which have attributed the observed differentiation to adaptive divergence in response to differing ecological conditions in pond and marine habitats.

The relative importance of sexual reproduction and clonal propagation in rhizomatous herb Iris japonica Thunb. from two habitats of Jinyun Mountain, Southwest China

Reproductive behaviour of clonal plants might change in contrasting habitats. In field and simulated experiments, we studied the relative importance of sexual reproduction and clonal propagation in rhizomatous herb, Iris japonica Thunb. in two forest habitats (BF, bamboo forest and OAFE, open area of forest edge), and effects of population origin (BF vs. OAFE) and environmental effects (shading) on sexual vs. clonal reproduction. In field experiment, the relative importance of reproduction in I. japonica populations was different in two habitats, which showed predominantly sexual reproduction in OAFE and clonal propagation in BF. In simulated experiment, the effect of population origin and light treatment (shading) was significant for reproduction of I. japonica. Clonal propagation was only influenced by population origin, and sexual reproduction was determined both by population origin and light treatment. A trade-off between two reproductive modes exhibited in both experiments. The trade-offs was more obvious in OAFE than in BF because sexual reproduction, resource and inter-specific competition obviously lacked in BF. The results indicated that the selective forces shaping reproduction of I. japonica in contrasting habitats might demonstrate pronounce adaptive population differentiation among forest habitats. Thus, I. japonica populations formed local differentiation by adaptation of reproduction to local heterogeneous forest habitats.

The evolution of quantitative traits in complex environments

Species inhabit complex environments and respond to selection imposed by numerous abiotic and biotic conditions that vary in both space and time. Environmental heterogeneity strongly influences trait evolution and patterns of adaptive population differentiation. For example, heterogeneity can favor local adaptation, or can promote the evolution of plastic genotypes that alter their phenotypes based on the conditions they encounter. Different abiotic and biotic agents of selection can act synergistically to either accelerate or constrain trait evolution. The environmental context has profound effects on quantitative genetic parameters. For instance, heritabilities measured in controlled conditions often exceed those measured in the field; thus, laboratory experiments could overestimate the potential for a population to respond to selection. Nevertheless, most studies of the genetic basis of ecologically relevant traits are conducted in simplified laboratory environments, which do not reflect the complexity of nature. Here, we advocate for manipulative field experiments in the native ranges of plant species that differ in mating system, life-history strategy and growth form. Field studies are vital to evaluate the roles of disparate agents of selection, to elucidate the targets of selection and to develop a nuanced perspective on the evolution of quantitative traits. Quantitative genetics field studies will also shed light on the potential for natural populations to adapt to novel climates in highly fragmented landscapes. Drawing from our experience with the ecological model system Boechera (Brassicaceae), we discuss advancements possible through dedicated field studies, highlight future research directions and examine the challenges associated with field studies.

Potential for evolutionary responses to climate change – evidence from tree populations

Evolutionary responses are required for tree populations to be able to track climate change. Results of 250 years of common garden experiments show that most forest trees have evolved local adaptation, as evidenced by the adaptive differentiation of populations in quantitative traits, reflecting environmental conditions of population origins. On the basis of the patterns of quantitative variation for 19 adaptation-related traits studied in 59 tree species (mostly temperate and boreal species from the Northern hemisphere), we found that genetic differentiation between populations and clinal variation along environmental gradients were very common (respectively, 90% and 78% of cases). Thus, responding to climate change will likely require that the quantitative traits of populations again match their environments. We examine what kind of information is needed for evaluating the potential to respond, and what information is already available. We review the genetic models related to selection responses, and what is known currently about the genetic basis of the traits. We address special problems to be found at the range margins, and highlight the need for more modeling to understand specific issues at southern and northern margins. We need new common garden experiments for less known species. For extensively studied species, new experiments are needed outside the current ranges. Improving genomic information will allow better prediction of responses. Competitive and other interactions within species and interactions between species deserve more consideration. Despite the long generation times, the strong background in quantitative genetics and growing genomic resources make forest trees useful species for climate change research. The greatest adaptive response is expected when populations are large, have high genetic variability, selection is strong, and there is ecological opportunity for establishment of better adapted genotypes.

Latitudinal gradients as natural laboratories to infer species' responses to temperature

Summary Macroclimatic variation along latitudinal gradients provides an excellent natural laboratory to investigate the role of temperature and the potential impacts of climate warming on terrestrial organisms. Here, we review the use of latitudinal gradients for ecological climate change research, in comparison with altitudinal gradients and experimental warming, and illustrate their use and caveats with a meta‐analysis of latitudinal intraspecific variation in important life‐history traits of vascular plants. We first provide an overview of latitudinal patterns in temperature and other abiotic and biotic environmental variables in terrestrial ecosystems. We then assess the latitudinal intraspecific variation present in five key life‐history traits [plant height, specific leaf area (SLA), foliar nitrogen:phosphorus (N:P) stoichiometry, seed mass and root:shoot (R:S) ratio] in natural populations or common garden experiments across a total of 98 plant species. Intraspecific leaf N:P ratio and seed mass significantly decreased with latitude in natural populations. Conversely, the plant height decreased and SLA increased significantly with latitude of population origin in common garden experiments. However, less than a third of the investigated latitudinal transect studies also formally disentangled the effects of temperature from other environmental drivers which potentially hampers the translation from latitudinal effects into a temperature signal. Synthesis. Latitudinal gradients provide a methodological set‐up to overcome the drawbacks of other observational and experimental warming methods. Our synthesis indicates that many life‐history traits of plants vary with latitude but the translation of latitudinal clines into responses to temperature is a crucial step. Therefore, especially adaptive differentiation of populations and confounding environmental factors other than temperature need to be considered. More generally, integrated approaches of observational studies along temperature gradients, experimental methods and common garden experiments increasingly emerge as the way forward to further our understanding of species and community responses to climate warming.