Population variation and natural selection on leaf traits in cork oak throughout its distribution range

A central issue in evolutionary biology is the exploration of functional trait variation among populations and the extent to which this variation has adaptive value. It was recently proposed that specific leaf area (SLA), leaf nitrogen concentration per mass (N_mass) and water use efficiency in cork oak play an important role in adaptation to water availability in the environment. In order to investigate this hypothesis, we explored, first, whether there was population-level variation in cork oak (Quercus suber) for these functional traits throughout its distribution range; if this were the case, it would be consistent with the hypothesis that different rainfall patterns have led to ecotypic differentiation in this species. Second, we studied whether the population-level variation matched short-term selection on these traits under different water availability conditions using two fitness components: survival and growth. We found high population-level differentiation in SLA and N_mass, with populations from dry places exhibiting the lowest values for SLA and N_mass. Likewise, reduced SLA had fitness benefits in terms of growth for plants under dry conditions. However, contrary to our expectations, we did not find any pattern of association between functional traits and survival in nine-year-old saplings despite considerable drought during one year of the study period. These results together with findings from the literature suggest that early stages of development are the most critical period for this species. Most importantly, these findings suggest that cork oak saplings have a considerable potential to cope with dry conditions. This capacity to withstand aridity has important implications for conservation of cork oak woodlands under the ongoing climate change.     

Isolation by adaptation in Neochlamisus leaf beetles: host-related selection promotes neutral genomic divergence

This study tests how divergent natural selection promotes genomic differentiation during ecological speciation. Specifically, we use adaptive ecological divergence (here, population divergence in host plant use and preference) as a proxy for selection strength and evaluate the correlation between levels of adaptive and genetic differentiation across pairwise population comparisons. Positive correlations would reveal the pattern predicted by our hypothesis, that of 'isolation by adaptation' (IBA). Notably, IBA is predicted not only for selected loci but also for neutral loci. This may reflect the effects of divergent selection on neutral loci that are 'loosely linked' to divergently selected loci or on geneflow restriction that facilitates genetic drift at all loci, including neutral loci that are completely unlinked to those evolving under divergent selection. Here, we evaluate IBA in maple- and willow-associated populations of Neochlamisus bebbianae leaf beetles. To do so, we collected host preference data to construct adaptive divergence indices and used AFLPs (amplified fragment length polymorphisms) and mitochondrial sequences to quantify genetic differentiation. Partial Mantel tests showed significant IBA in 'pooled' analyses of putatively neutral and of putatively selected ('outlier') AFLP loci. This pattern was also recovered in 12% of 'locus-specific' analyses that separately evaluated genetic differentiation at individual neutral loci. These results provided evidence for widespread effects of selection on neutral genomic divergence. Our collective findings indicate that host-related selection may play important roles in the population genomic differentiation of both neutral and selected gene regions in herbivorous insects.     

西藏紫花针茅叶功能性状沿降水梯度的变化

植物叶功能性状与环境因子的关系是近10年来植物生态学的研究热点。该文以广泛分布于青藏高原干旱、半干旱草地的优势植物种紫花针茅(Stipa purpurea)为研究对象, 沿降水梯度(69-479 mm)系统测定了日土、改则、珠峰、当雄和纳木错5个调查地点紫花针茅比叶面积(SLA)、单位重量和单位面积叶氮含量(N_mass, N_area)、叶密度和厚度等叶功能性状以及土壤全氮含量等因子, 试图验证干旱胁迫地区同一物种内SLA-N_mass关系沿降水梯度的策略位移现象是否具有普遍性, 并对是否出现策略位移现象提出可能的解释。研究结果表明: 1) SLA和N_mass与生长季温度和降水以及土壤全氮含量均没有显著关系, SLA与N_mass的关系在干旱半干旱区(年降水/蒸发比< 0.11)与半湿润区(年降水/蒸发比> 0.11)之间并没有出现典型的位移现象; 2)叶密度是决定半湿润区SLA变化的主导因子, 而叶厚度则是干旱半干旱区SLA变化的控制因子, 两者与SLA均呈负相关, 随着温度增加或降水减少, 叶厚度增加而叶密度降低, 导致SLA随温度和降水变化不明显; 3)半湿润区的叶密度增加引起N_area增加, 而干旱半干旱区的叶厚度增加并没有造成N_area的显著变化, 导致N_area沿降水梯度没有显著变化; 4)紫花针茅地上生物量与N_area具有显著正相关关系, 表明N_area的增加有助于提高植被生产力。结果表明, 在干旱胁迫下, 植物通过增加叶厚度来维持不变的N_area可能有助于保持与较湿润地区相似的光合生产和水分利用效率。叶厚度和叶密度对比叶面积的相对影响在干旱半干旱区与半湿润区之间发生转变, 这为进一步检测高寒草地植被的水分限制阈值提供了新思路。     

唐古特白刺叶功能性状沿气候梯度的变异特征

叶功能性状与植物的生长对策及资源利用效率密切相关,研究叶功能性状沿气候梯度的变异特征能为理解植物对气候变化的响应机制提供一种简便可行的测定指标。以我国西北荒漠地区广泛分布的唐古特白刺（Nitraria tangutorum）为研究对象,对其比叶面积（SLA）、单位质量和单位面积叶氮含量（N_mass、N_area）、单位质量和单位面积叶建成成本（CC_mass、CC_area）进行测定,分析这些叶功能性状及性状相关关系沿气候梯度的变异特征。结果表明,唐古特白刺叶功能性状（CC_area除外）在气候梯度下存在显著差异,其中,温度是决定唐古特白刺SLA变化的主要因子,SLA随着温度的增加而增加;降水和温度对唐古特白刺N_mass、N_area和CC_mass均有显著影响,N_mass和N_area随着降水和温度的增加而降低,而CC_mass呈增加趋势。沿气候梯度,唐古特白刺SLA-N_mass、CC_mass-N_mass和CC_area-N_area的线性正相关关系发生平移,导致在相同SLA、CC_mass和CC_area下,降水和温度较低的地区具有更高的N_mass和N_area。这一结果表明唐古特白刺能通过调节叶功能性状之间的关系来适应气候的变化,并形成性状间的最佳功能组合。     

Linked selection,ancient polymorphism,and ecological adaptation shape the genomic landscape of divergence in Quercus dentata

Multiple evolutionary forces contribute to heterogeneous genomic landscapes; however, disentangling their relative contributions is challenging. We sampled nine populations across the distribution of Quercus dentata, a dominant forest tree in East Asia, and used whole-genome sequencing data to investigate mechanisms underlying divergence. We identified two genetic groups (north and south) that diverged ~1.84 million years ago, consistent with the uplift of the Qinling Mountains during the Pleistocene. The north group experienced a bottleneck during the middle–late Pleistocene and expanded from multiple refugia. The south group experienced a more severe bottleneck and showed high population differentiation, probably due to long-term isolation and habitat fragmentation. We detected genomic islands with elevated relative differentiation (F_ST) scattered across the genome. Among these, 65.9% showed reduced absolute divergence (d_XY) consistent with linked selection, while the remaining (34.1%) showed elevated d_XY suggestive of divergent sorting of ancient polymorphisms. The recombination rate in genomic islands was lower than background, suggesting the importance of genome structure in shaping the genomic landscape. We detected 108 single nucleotide polymorphisms significantly associated with environmental factors, 12 of which clustered in a region of ~500 kb. This region showed multiple signals of positive selection in the north group, including the enrichment of XP-extended haplotype homozygosity scores, an elevated population branch statistic, and an excess of high-frequency derived alleles. In addition, we found that linkage disequilibrium was low and derived haplotypes declined rapidly in this region, indicating selection on standing variation. Our results clarify the evolutionary processes driving genomic divergence in Q. dentata.     

羌塘高原降水梯度植物叶片、根系性状变异和生态适应对策

叶片和根系是植物获取资源的最重要的器官,其性状随环境梯度的变化反映了植物光合碳获取和水分与养分的吸收能力及其对环境变化适应的生态对策。羌塘高原降水梯度带高寒草地群落叶片和根系成对性状关系研究不仅能揭示环境梯度对植物性状的塑造作用,也可为理解寒、旱和贫瘠等极端环境下植物的适应策略提供依据。为此,选择3组具有代表性的叶片和根系成对性状:比叶面积(SLA)和比根长(SRL);单位质量叶氮含量(LN_mass)和单位质量根氮含量(RN_mass);单位面积叶氮含量(LN_area)和单位长度根氮含量(RN_length),分析不同优势植物地上、地下成对性状变异特征及其与环境因子的关系,探讨植物性状对高寒生态系统水分和养分限制因素的适应策略。研究表明,区域气候和土壤环境导致的叶片性状变异大于根系性状的变异,干旱端的植物既具有高的SRL,又具有高的叶片和根系的养分含量(LN_mass,LN_area和RN_mass)。SLA-SRL、LN_mass     

Climatic drivers of leaf traits and genetic divergence in the tree Annona crassiflora: a broad spatial survey in the Brazilian savannas

The Cerrado is the largest South American savanna and encompasses substantial species diversity and environmental variation. Nevertheless, little is known regarding the influence of the environment on population divergence of Cerrado species. Here, we searched for climatic drivers of genetic (nuclear microsatellites) and leaf trait divergence in Annona crassiflora, a widespread tree in the Cerrado. The sampling encompassed all phytogeographic provinces of the continuous area of the Cerrado and included 397 individuals belonging to 21 populations. Populations showed substantial genetic and leaf trait divergence across the species' range. Our data revealed three spatially defined genetic groups (eastern, western and southern) and two morphologically distinct groups (eastern and western only). The east‐west split in both the morphological and genetic data closely mirrors previously described phylogeographic patterns of Cerrado species. Generalized linear mixed effects models and multiple regression analyses revealed several climatic factors associated with both genetic and leaf trait divergence among populations of A. crassiflora. Isolation by environment (IBE) was mainly due to temperature seasonality and precipitation of the warmest quarter. Populations that experienced lower precipitation summers and hotter winters had heavier leaves and lower specific leaf area. The southwestern area of the Cerrado had the highest genetic diversity of A. crassiflora, suggesting that this region may have been climatically stable. Overall, we demonstrate that a combination of current climate and past climatic changes have shaped the population divergence and spatial structure of A. crassiflora. However, the genetic structure of A. crassiflora reflects the biogeographic history of the species more strongly than leaf traits, which are more related to current climate.     

Patterns in leaf traits of leguminous and non-leguminous dominant trees along a rainfall gradient in Ghana

Aims Both dominance distribution of species and the composition of the dominant species determine the distribution of traits within community. Leaf carbon (C) and nitrogen (N) isotopic composition are important leaf traits, and such traits of dominant species are associated with ecosystem C, water and N cycling. Very little is known how dominant species with distinct traits (e.g. N-fixing leguminous and non-leguminous trees) mediate resource utilization of the ecosystems in stressful environment.Methods Leaves of 81 dominant leguminous and non-leguminous trees were collected in forest (moist semi-deciduous and dry semi-deciduous ecosystems) and savanna (costal savanna, Guinean savanna and west Sudanian savanna ecosystems) areas and the transitional zone (between the forest and the savanna) along the transect from the south to the north of Ghana. We measured leaf traits, i.e. leaf δ 13 C, leaf δ 15 N, leaf water content, leaf mass per area (LMA) and C and N concentration. Correlation analyses were used to examine trait–trait relationships, and relationships of leaf traits with temperature and precipitation. We used analysis of covariance to test the differences in slopes of the linear regressions between legumes and non-legumes.Important findings Leaf δ 13 C, δ 15 N, leaf water content and LMA did not differ between leguminous and non-leguminous trees. Leaf N concentration and C:N ratio differed between the two groups. Moreover, leaf traits varied significantly among the six ecosystems. δ 13 C values were negatively correlated with annual precipitation and positively correlated with mean annual temperature. In contrast, leaf δ 15 N of non-leguminous trees were positively correlated with annual precipitation and negatively correlated with mean annual temperature. For leguminous trees, such correlations were not significant. We also found significant coordination between leaf traits. However, the slopes of the linear relationships were significantly different between leguminous and non-leguminous trees. Our results indicate that shifts in dominant trees with distinct water-use efficiency were corresponded to the rainfall gradient. Moreover, leguminous trees, those characterized with relative high water-use efficiency in the low rainfall ecosystems, were also corresponded to the relative high N use efficiency. The high proportion of leguminous trees in the savannas is crucial to mitigate nutrient stress.     

Adaptive population differentiation in phenology across a latitudinal gradient in European aspen (Populus tremula, L.): a comparison of neutral markers, candidate genes and phenotypic traits

A correct timing of growth cessation and dormancy induction represents a critical ecological and evolutionary trade-off between survival and growth in most forest trees (Rehfeldt et al. 1999; Horvath et al. 2003; Howe et al. 2003). We have studied the deciduous tree European Aspen (Populus tremula) across a latitudinal gradient and compared genetic differentiation in phenology traits with molecular markers. Trees from 12 different areas covering 10 latitudinal degrees were cloned and planted in two common gardens. Several phenology traits showed strong genetic differentiation and clinal variation across the latitudinal gradient, with Q(ST) values generally exceeding 0.5. This is in stark contrast to genetic differentiation at several classes of genetic markers (18 neutral SSRs, 7 SSRs located close to phenology candidate genes and 50 SNPs from five phenology candidate genes) that all showed F(ST) values around 0.015. We thus find strong evidence for adaptive divergence in phenology traits across the latitudinal gradient. However, the strong population structure seen at the quantitative traits is not reflected in underlying candidate genes. This result fit theoretical expectations that suggest that genetic differentiation at candidate loci is better described by F(ST) at neutral loci rather than by Q(ST) at the quantitative traits themselves.     

不同模拟增雨下白刺比叶面积和叶干物质含量的比较

以荒漠生态系统典型植物白刺(Nitraria tangutorum)为研究对象,根据内蒙古磴口多年平均降水量和植物生长规律,设计两个增雨时段(生长季前期与生长季后期),每个增雨时段设置两个增雨梯度(72.5mm/a(50%)、145mm/a(100%)),对天然白刺灌丛进行增雨实验,研究了不同模拟增雨处理下2012年与2013年生长季白刺叶片的比叶面积(SLA)与叶干物质含量(LDMC)的变化。结果表明,增雨处理可以增加白刺叶片的SLA及LDMC,同时期增雨100%处理对SLA及LDMC的影响大于50%处理,但同时期增雨的两个处理之间无显著差异;白刺叶片SLA在生长季前期对水分响应明显,LDMC则在生长季后期对水分反应敏感;相同增雨处理,2012年白刺叶片SLA及LDMC的净增加值高于2013年;SLA与LDMC在2012年呈显著负相关,在2013年虽呈负相关,但相关性不显著。在未来降雨增加的背景下,荒漠植物白刺叶片SLA与LDMC对增雨具有较强的协调适应能力,在不同生长季节可以通过改变不同的叶片性状来适应环境变化。     

中国东部南北样带暖温带区栎属树种叶片形态性状对气候条件的响应

植物叶片的形态性状能够有效地反映生存环境的变化, 并且影响植物的基本行为和功能。该研究通过获取植物标本提供的叶片形态信息, 结合相关分析和标准化主轴分析, 探讨了南北样带暖温带区栎属(Quercus)树种叶片形态性状对气候条件的响应及适应策略。结果表明: 在南北样带暖温带区, 随着气候条件的变化, 栎属树种的叶片形态性状发生显著的变化。随着年平均气温的降低和年日照时数的增加, 栎属树种叶面积增加, 以利于吸收更多的光照辐射, 并增加叶片的边界层阻力, 减少叶片热量的散失; 而叶片分裂程度的增加不仅可以降低热量的散失, 也可以提高叶片液流的波动以增强叶片的生理活动; 叶脉密度随温度的升高、光照强度和降水量的增加而增加, 以响应叶片蒸腾作用的增强, 提高水分的运输能力和叶片的支撑能力。此外, 为适应南北样带暖温带区气候条件的变化, 栎属树种的叶片形成了一系列的形态性状组合, 随着叶面积的增加, 叶柄长度和叶片分裂程度逐渐增加, 而叶脉密度降低; 随着叶片倾向于向长条状发展, 叶柄长度和叶脉密度也随之增加。     

Size‐dependent and environment‐mediated shifts in leaf traits of a deciduous tree species in a subtropical forest

AimsUnderstanding the joint effects of plant development and environment on shifts of intraspecific leaf traits will advance the understandings of the causes of intraspecific trait variation. We address this question by focusing on a widespread species Clausena dunniana in a subtropical broad‐leaved forest.MethodsWe sampled 262 individuals of C. dunniana at two major topographic habitat types, the slope and hilltop, within the karst forests in Maolan Nature Reserve in southwestern China. We measured individual plant level leaf traits (i.e., specific leaf area (SLA), leaf area, leaf dry‐matter content (LDMC), and leaf thickness) that are associated with plant resource‐use strategies. We adopted a linear mixed‐effects model in which the plant size (i.e., the first principal component of plant basal diameter and plant height) and environmental factors (i.e., topographic habitat, canopy height, and rock‐bareness) were used as independent variables, to estimate their influences on the shifts of leaf traits.Key ResultsWe found that (1) plant size and the environmental factors independently drove the intraspecific leaf trait shifts of C. dunniana, of which plant size explained less variances than environmental factors. (2) With increasing plant size, C. dunniana individuals had increasingly smaller SLA but larger sized leaves. (3) The most influential environmental factor was topographic habitat; it drove the shifts of all the four traits examined. Clausena dunniana individuals on hilltops had leaf traits representing more conservative resource‐use strategies (e.g., smaller SLA, higher LDMC) than individuals on slopes. On top of that, local‐scale environmental factors further modified leaf trait shifts.ConclusionsPlant size and environment independently shaped the variations in intraspecific leaf traits of C. dunniana in the subtropical karst forest of Maolan. Compared with plant size, the environment played a more critical role in shaping intraspecific leaf trait variations, and potentially also the underlying individual‐level plant resource‐use strategies.     

Variation in plant functional traits across and within four species of Western Australian Banksia (Proteaceae) along a natural climate gradient

Plant traits are fundamental components of the ecological strategies of plants, relating to how plants acquire and use resources. Their study provides insight into the dynamics of species geographical ranges in changing environments. Here, we assessed the variation in trait values at contrasting points along an environmental gradient to provide insight into the flexibility of species response to environmental heterogeneity. Firstly, we identified how commonly measured functional traits of four congeneric species (Banksia baxteri, B. coccinea, B. media and B. quercifolia) varied along a longitudinal gradient in the South Western Australian Floristic Region. This regional gradient provides significant variation in moisture, temperature and soil nutrients: soil nitrogen content decreases with declining rainfall and increasing temperature. We hypothesized that (i) the regional pattern in trait–environment associations across the species would match those observed on a global scale and (ii) that the direction and slopes of the within‐species relationships would be similar to those across species for each of the measured traits. Along the regional gradient we observed strong shifts in trait values, and cross‐species relationships followed the expected trend: specific leaf area was significantly lower, and leaf N_area and seed dry mass significantly higher, at the drier end of the rainfall gradient. However, traits within species were generally not well correlated with habitat factors: we found weak patterns among populations, either due to the small ecological gradient or perhaps because fine‐scale structuring among populations (at a micro‐evolutionary scale) was low due to high gene flow within species. Understanding how species traits shift as a result of climatic influences, both at the regional (across species) and local (within species) scale, provides insight into plant adaptation to the environment. Such studies have important applications for conservation biology and population management in the face of global change.     

Rapid reversal of a potentially constraining genetic covariance between leaf and flower traits in Silene latifolia

Genetic covariance between two traits generates correlated responses to selection, and may either enhance or constrain adaptation. Silene latifolia exhibits potentially constraining genetic covariance between specific leaf area (SLA) and flower number in males. Flower number is likely to increase via fecundity selection but the correlated increase in SLA increases mortality, and SLA is under selection to decrease in dry habitats. We selected on trait combinations in two selection lines for four generations to test whether genetic covariance could be reduced without significantly altering trait means. In one selection line, the genetic covariance changed sign and eigenstructure changed significantly, while in the other selection line eigenstructure remained similar to the control line. Changes in genetic variance–covariance structure are therefore possible without the introduction of new alleles, and the responses we observed suggest that founder effects and changes in frequency of alleles of major effect may be acting to produce the changes.     

海南岛热带山地雨林幼苗幼树光合与叶氮、叶磷及比叶面积的关系

以海南岛吊罗山热带山地雨林101个物种的幼苗幼树为试验材料,测定其光合、叶片氮、磷含量及比叶面积;检验其相关关系,并按乔木,乔灌木(小乔木至大灌木)和灌木3个生活型组进行分组检验。研究结果表明,单位叶面积(Aarea)和单位叶重量的光合速率(Amass)均表现出灌木>乔木>乔灌木,方差分析表明,灌木和乔灌木之间Aarea差异显著;灌木和乔木以及灌木和乔灌木之间Amass差异显著(p<0.05)。Aarea与叶氮含量之间的相关性在不同生态型组和所有物种之间均达到极显著水平(p<0.0001);与叶磷之间的相关关系在灌木(p=0.0038),乔灌木(p=0.0002)以及所有物种(p<0.0001)之间达到极显著水平,但是在乔木中未达到显著水平(p>0.05);与SLA之间在灌木(p=0.0006)、乔木(p<0.0001)和所有物种(p<0.0001)之间达到极显著水平,但是在乔灌木中未达到显著水平(p>0.05)。Amass与叶片氮含量、SLA的相关关系在不同生活型组和所有物种中都达到极显著水平(p<0.0001);与叶磷含量之间的相关性在灌木(p=0.0004),乔灌木(p=0.0018)及所有物种(p<0.0001)中极显著,在乔木生活型组中也达显著水平(p=0.0377)。逐步回归表明,与Aarea相比,Amass估计结果更接近于实际测值。由此可见,海南岛热带山地雨林林下幼苗幼树的光合和叶氮、磷含量及SLA之间相关关系与基于成树的研究非常相似,并且A比A更能稳定体现这种相关性。