表型可塑性和局域适应在紫茎泽兰入侵不同海拔生境中的作用

紫茎泽兰是我国危害最严重的外来入侵物种之一,为探讨表型可塑性和局域适应在其入侵中的作用,在高、低海拔的两个样地内,测定了来自云南南部640～2450 m海拔范围的6个种源的紫茎泽兰种群的株高、冠宽、分枝数和高温半致死温度(HSLT).结果表明,在高海拔样地,各种群紫茎泽兰株高、冠宽、分枝数和HSLT(2130 m的哀牢山种群除外)均显著低于在低海拔样地,紫茎泽兰各种群的株高、冠宽和分枝数的可塑性指数(0.881～0.975)均较大,而HSLT的可塑性(0.052～0.200)较小.无论在高还是低海拔样地,紫茎泽兰的株高、冠宽和分枝数在种群间的差异均不显著,而HSLT在种群间的差异达极显著水平,表现出明显的遗传分化,但其在种群间的差异仍小于其在样地间的差异.在高海拔样地,紫茎泽兰各种群的分枝数与种源海拔呈显著正相关;在低海拔样地,紫茎泽兰的HSLT与种源海拔呈显著负相关,表现出明显的局域适应特征.表型可塑性和局域适应均与紫茎泽兰的入侵有关,但前者的作用可能更大.     

多地同质园实验揭示普通野生稻的表型可塑性与本地适应性

边缘种群的表型可塑性和本地适应性的解析是预测植物如何响应气候变化的基础。本研究以普通野生稻(Oryza rufipogon)的北缘种群为核心对象,以中心种群(中部、南部种群)为对照,进行3地同质园实验,以“空间换时间”的策略模拟未来气温变化情形下普通野生稻种群的表型表现。结果显示,普通野生稻各种群具有丰富的表型可塑性,种群间表型可塑性存在差异。北缘种群在株高、种子量和越冬存活率等指标上表现出明显的本地适应性,反映了对相对低温环境的适应。北缘种群无论北移还是南移都能够正常结实、完成生活史,说明在未来气温变化情形下也能生存;中部和南部种群在上海同质园都不能越冬存活,提示极端低温对于低纬度种群影响较大。这些结果表明,尽管普通野生稻存在明显的适应性分化,但有较强的表型可塑性能力,北缘种群和中心种群都能适应当前的气候变暖。     

海南凤仙花不同海拔种群的传粉生物学

凤仙花属(Impatiens)植物具有极为广泛的多样性和类型各异的特化传粉者, 被誉为“双子叶的兰花”, 受到众多传粉生物学家的关注。本文以海南特有种海南凤仙花(Impatiens hainanensis)为研究对象, 对3个不同海拔梯度的种群进行了开花生物学特性和开花物候、花器官结构、花粉活力和柱头活性、传粉者种类和访花行为及繁育系统的比较研究。结果表明: 单花花期4.10 ± 0.46 d, 雄性期和雌性期分别约为3.15 ± 0.24 d和0.95 ± 0.36 d; 种群开花峰期在8月初, 高海拔种群的花期高峰相对滞后。低、中海拔种群花粉活力呈现先升高后下降的趋势, 以开花第2 d花粉活力最高; 高海拔种群花粉活力随开花时间推移逐渐下降; 柱头活性随开花时间的推移而增强, 高海拔种群开花各天次均较低、中海拔种群低。主要传粉昆虫为黄黑无垫蜂(Amegilla leptocoma)和绿条无垫蜂(A. zonata), 低、中海拔种群以黄黑无垫蜂为主, 高海拔种群以绿条无垫蜂为主。未观察到自动自花授粉和无融合生殖现象, 人工授粉能明显增加坐果率(75-90%), 自然坐果率在高海拔种群相对较低(40-60%), 说明存在较强的传粉者限制。海南凤仙花的保护需要同时关注其有效传粉者的保护, 促进有效传粉昆虫在不同海拔种群之间的往来, 保证种群间的花粉流与种子流, 维持海南凤仙花的种群遗传多样性与有效种群大小。     

基于野外调查和同质种植园实验的芦苇植物功能性状变异研究

研究表型可塑性和遗传变异在植物表型分化中的相对作用,有助于预测全球环境变化下的植物群落组成和生态系统功能的变化。芦苇（Phragmites australis）是全球性广布的草本植物,种内变异丰富,在我国西北和东部均存在多个分化稳定的生态型,但中国芦苇在更大尺度上的表型研究还非常匮乏。将位于黄河上游的宁夏平原和黄河下游的黄河三角洲作为研究区域,通过野外调查和同质种植园实验对芦苇自然种群的植物功能性状变异进行观测。结果表明,无论在野外还是同质种植园,黄河三角洲芦苇的基径、叶长和叶宽均显著大于宁夏平原芦苇,说明两个地区的芦苇种群之间存在着受遗传决定的表型分化,这可能与两个地区间的降水等气候差异有关。在野外,宁夏芦苇的株高和叶厚显著大于黄河三角洲芦苇,但在同质园中差异消失或相反,说明株高、叶厚受环境影响较大,表型可塑性也是芦苇适应环境变化的重要机制。在同质种植园中,宁夏平原芦苇的叶片氮磷含量较低,但株数却显著多于黄河三角洲芦苇,反映了不同地区芦苇之间存在不同的适应策略,宁夏平原芦苇更偏向于高扩散率的杂草策略,而黄河三角洲芦苇更偏向于竞争策略。此外,宁夏平原芦苇的株高、叶长两个性状以及基径-比叶面积相关性在野外和同质园两个环境中存在一致性,表明了性状变异和权衡策略的遗传稳定性。综上,位于黄河上下游的芦苇种群间存在着适应性分化,这是表型可塑性和遗传变异共同作用的结果,不同来源芦苇对全球变化下的多重环境因子的响应还需要进一步研究。     

中国东部沿海互花米草种群生活史特征的纬度变异与可塑性

互花米草(Spartina alterniflora)于20世纪70年代被引入中国,目前已在东部沿海盐沼湿地中广泛分布,成为海岸带盐沼中危害严重的入侵植物之一.为了研究互花米草在中国入侵区中的适应机制,揭示遗传分化和表型可塑性在该物种成功入侵中的作用,本研究沿纬度梯度在南起广东(22°N)、北至天津(39°N)的沿海样带上采集了10个种群的样本,通过同质园实验比较了不同纬度来源的种群在,生活史和生长特征方面是否存在遗传分化,并平行设置高低两个水位处理以比较互花米草对水位变化(不同高程生境条件)的可塑性反应.结果表明,在所研究的互花米草17个性状中有12个存在显著的种群间差异.其中,平均开花日期和相对生长率(植株高度)表现出显著的纬度梯度变异:随着纬度的升高,开花时间提前,相对生长速率(植株高度)趋于增加.同时17个性状中有9个在不同水位处理之间存在显著差异.这些结果表明,遗传分化可能是互花米草能够快速占据广阔分布区的重要原因之一,而表型可塑性可能对互花米草在小尺度上占据不同高程环境的过程有重要作用.     

祁连山优势树木碳水化合物资源分配的海拔和树种效应

海拔梯度造成的温度、水分和土壤肥力等环境异质性会影响树木的生长, 但是不同树木的生理差异也决定了树木资源分配权衡还存在很多不确定性。为探明祁连山地区青海云杉(Picea crassifolia)和祁连圆柏(Juniperus przewalskii)非结构碳水化合物(NSC)含量的海拔和树种效应差异, 该研究以祁连山两个优势针叶树种青海云杉和祁连圆柏为研究对象, 分别设置了高海拔(3 300 m)和低海拔(2 850 m)两个采样高度, 采用t检验和多因素方差分析比较了不同海拔两个树种NSC及其组分(可溶性糖、淀粉)含量的差异, 分析叶、树干、粗根和细根间的NSC及其组分含量的资源分配权衡特征, 明晰不同海拔青海云杉和祁连圆柏生长限制因素和生理生态适应机理。结果表明: (1)低海拔青海云杉整株和各器官NSC及其组分含量均显著高于高海拔青海云杉; 而低海拔祁连圆柏整株和各器官NSC及其组分含量均显著低于高海拔祁连圆柏; (2)不同海拔两个优势针叶树种可溶性糖主要投资在叶, 而淀粉主要投资在粗根和树干; (3)高海拔青海云杉的可溶性糖与淀粉的比值显著高于低海拔青海云杉, 说明高海拔青海云杉将更多的碳用于生长, 而低海拔青海云杉将更多的碳用于储存; (4)海拔、树种、器官以及它们的交互作用显著地影响NSC及其组分含量及可溶性糖:淀粉, 其中树种分别解释了NSC、淀粉总变异的38%和37%; 器官分别解释了可溶性糖、可溶性糖:淀粉总变异的68%和42%。该研究结果阐明了不同海拔青海云杉和祁连圆柏的生长限制因素及其资源分配权衡, 为理解不同海拔和不同树种的生态适应机理以及祁连山森林生态系统的保护提供了科学参考。     

承德光秃山不同海拔油松居群遗传多样性

为了解光秃山不同海拔下天然油松居群之间的遗传差异及探求遗传多样性与土壤因子的关联,为油松种源筛选和管理提供参考,本研究运用ISSR技术,对位于河北承德辽河源光秃山4个不同海拔油松天然居群共118个植株个体的遗传多样性进行分析。13个引物共扩增出177条清晰的条带,种群多态位点百分比(PPL)为60.2775%,Nei’s基因多样性指数(h)为0.2171,Shannon信息指数(I)为0.3222;不同种群遗传变异水平随海拔差异呈规律性变化,表现为沿海拔升高而呈低-高-低的分布规律,其中1354～1274 m范围的遗传多样性水平最高;在物种水平上油松具有较高的遗传多样性(PPL=98.33%,h=0.38142,I=0.5550),种群间的遗传分化系数Gst=0.6562。利用AMOVA软件对遗传变异的等级剖分结果表明,种群间有显著的遗传分化,约2/5的遗传变异存在种群间,种群内占3/5。Pearson相关分析表明,油松居群内遗传多样性与海拔、土壤养分(有机质、速效磷、速效钾含量)之间存在显著或一定的相关关系。Mantel检验结果显示,油松居群遗传距离与海拔差距、土壤养分因子的分异存在一定相关性。以上结果表明不同海拔区域的生态因子、低基因流等对油松居群间的遗传分化影响较大。     

Environmental pressure rather than ongoing hybridization is responsible for an altitudinal cline in the morphologies of two oaks

在相关类群之间的接触区,表型变异可能源于遗传和/或环境梯度。本研究旨在阐明两种栎属植物[Quercus crispula (QCC)和 Q. mongolicoides (QCM)],在其接触区沿海拔梯度形成叶片形态表型变异的原因。对于源自接触区48个个体和QCC和QCM的每个参照 种群24个个体样本,我们测定其6种叶片形态特征,同时记录13个核微卫星位点的基因型。我们通过构建模型解释表型变异(叶片形态)与环境(海拔)和遗传(参照种群世系)梯度的关系。研究结果表明,形态和遗传标记均能较好地区分参照种群中的两个品种。我们能够确认形态和遗传标记的作用。接触区种群内的个体具有略偏向QCM分支的中间世系,其形态分布与参照种群中两个变种的形态分布重叠。海拔会显著影响叶片形态性状,而遗传对叶片形态性状无显著影响。接触区种群的世系和种间杂合性分布与F₂或后代杂交种中的分布相似。这些结果表明,在两种栎属植物QCC和QCM之间的接触区,并没有发生杂交,但环境压力通过表型可塑性和/或功能基因的变异,导致了其在形态性状上的海拔梯度效应。     

海拔梯度上青海云杉径向生长与气候关系稳定性研究

以祁连山北坡海拔2 700m(分布下限)、3 000m(中部)和3 300m(分布上限)的青海云杉为研究对象,采用长短两种不同步长的样条函数去趋势对1952～1980年和1981～2009年两个时段青海云杉径向生长与气候的关系进行对比分析,以揭示青海云杉年轮中包含的不同频率生长趋势与气候之间的关系及其在海拔梯度上的差异,探讨海拔梯度上气候变化对树木径向生长的影响。结果显示:(1)随着海拔升高,限制青海云杉径向生长的气候因子从降水转变为温度,不同时期的限制因子也发生变化。(2)在海拔2 700m处,上一年12月份平均温度与青海云杉生长的关系发生了分异,小步长的样条函数得到的年表中保留了较多降水信号;海拔3 000m处青海云杉径向生长与上年9月份降水量、平均温度表现出显著正相关关系;海拔3 300m处青海云杉生长与当年夏季降水量呈显著正相关关系,而在后一时段相关关系不显著,与当年春季、夏季的平均温度相关关系不稳定,大步长的样条函数得到的年表中表现出温度与生长关系的分异,且不同海拔青海云杉径向生长的降水量制约趋势变得明显。研究表明,祁连山北坡青海云杉与气候的关系在海拔梯度上存在差异,其对气候变化的响应也存在差异,水分的限制作用随着温度的逐年升高逐渐突出;采用不同步长的样条函数去趋势可以在年表中保留不同的气候信息,从而对分异现象分析结果产生影响,选择合理步长的样条函数进行去趋势才能得到合理准确的分析结果。     

植物群体表观遗传学研究进展

植物表观遗传多样性是对生物多样性的遗传多样性层面的补充,也是植物表型多样性的重要来源,研究表观遗传多样性的群体表观遗传学应运而生。在植物自然种群的研究中,群体表观遗传学弥补了群体遗传学对不符合孟德尔遗传定律的表型遗传的认识,是对现代综合进化论的重要补充。分子生物学技术的发展,为研究表观遗传变异提供了甲基化敏感的扩增片段长度多样(MS-AFLP)、结合二代测序的亚硫酸氢钠测序法等有力的技术手段。在生态和进化领域,自然种群中表观遗传变异与遗传变异、表型可塑性、生境分化、物种形成的关系成为研究的热点。表观遗传机制在生态系统和生物进化中的作用也逐渐得到揭示,本综述回顾近年来植物群体表观遗传学的实验研究和理论观点,展望了植物群体表观遗传学在研究方法和研究主题上的前景。     

Adaptive responses for seed and leaf phenology in natural populations of sessile oak along an altitudinal gradient

We assessed the adaptive potential of seed and leaf phenology in 10 natural populations of sessile oak (Quercus petraea) sampled along two altitudinal transects using common garden experiments. Population differentiation for both phenological traits was observed with high-altitude populations germinating and flushing later than low altitude ones. However, high genetic variation and heritability values were also maintained within populations, despite slightly decreasing for dates of leaf unfolding with increasing altitude. We suggest that biotic and abiotic fluctuating selection pressures within populations and high gene flow are the main mechanisms maintaining high genetic variation for these fitness related traits. Moreover, changes in selection intensity and/or selection pressures along the altitudinal gradient can explain the reduction in genetic variation observed for leaf phenology. We anticipate that the maintenance of high genetic variation will be a valuable resource for future adaptation of sessile oak populations undergoing an upslope shift caused by climate change.     

天目山不同海拔檫木群体遗传多样性和遗传结构

采用13对SSR引物,运用Bioptic Qsep100全自动核酸分析系统,分析了天目山5个海拔檫木群体的遗传多样性和遗传结构及其在不同海拔下的变化模式.结果表明: 天目山檫木群体具有较高的遗传多样性水平,其中期望杂合度和观察杂合度分别为0.84和0.61.根据Shannon指数,天目山檫木中海拔(500～800 m)群体的遗传多样性水平大于低海拔(200 m)和高海拔(1100～1400 m)群体的遗传多样性水平.由基因分化系数和AMOVA分析可知,檫木种群的遗传变异主要存在于群体内.STRUCTURE分析和UPGMA聚类分析表明,中、低海拔被划为一个群体,而高海拔被划为另一个群体.低海拔和中海拔檫木群体遗传距离的差异表明,人为干扰对物种多样性具有负面效应,而自然保护区对物种多样性的保护起到了积极作用.     

Morphological, genetic, and chromosomal variation at a small spatial scale within a mosaic hybrid zone of the grasshopper Dichroplus pratensis Bruner (Acrididae)

Hybrid zones are regions where genetically different populations meet and mate, resulting in offspring of mixed characteristics. In organisms with limited dispersal, such as melanopline grasshoppers, hybrid zones can occur at small spatial scales (i.e., <500 m). We assessed levels of morphological, chromosomal, and molecular variability in adult males of the grasshopper Dichroplus pratensis Bruner (N = 137 males, 188 females) collected at 12 sites within a mosaic hybrid zone in a heterogeneous environment in Sierra de la Ventana, Argentina. In this hybrid zone, 2 Robertsonian chromosomal races, polymorphic for different centric fusions, meet (the "Northern race" at low altitudes and the "Southern race" at higher altitudes), forming hybrids that show monobrachial homologies during meiosis. High morphometric variation in 6 traits was revealed among grasshoppers of both sexes, with male body size positively and significantly correlated with increasing altitude. Frequency of Robertsonian fusions characteristic of the Southern race increased significantly with altitude. Moreover, fusion frequencies covaried between samples. Considerable genetic variation was revealed by random amplification of polymorphic DNA markers, with heterozygosity ranging from 0.3477 to 0.3745. Insects from low-altitude and high-altitude populations showed significant genetic differentiation, as indicated by F(ST) values. The proposed model for D. pratensis, involving the generation and maintenance by chromosomal fusions, of gene complexes adaptive in different environments, could explain the observed clinal patterns within the contact zone.     

Fine‐grained adaptive divergence in an amphibian: genetic basis of phenotypic divergence and the role of nonrandom gene flow in restricting effective migration among wetlands

Adaptive ecological differentiation among sympatric populations is promoted by environmental heterogeneity, strong local selection and restricted gene flow. High gene flow, on the other hand, is expected to homogenize genetic variation among populations and therefore prevent local adaptation. Understanding how local adaptation can persist at the spatial scale at which gene flow occurs has remained an elusive goal, especially for wild vertebrate populations. Here, we explore the roles of natural selection and nonrandom gene flow (isolation by breeding time and habitat choice) in restricting effective migration among local populations and promoting generalized genetic barriers to neutral gene flow. We examined these processes in a network of 17 breeding ponds of the moor frog Rana arvalis, by combining environmental field data, a common garden experiment and data on variation in neutral microsatellite loci and in a thyroid hormone receptor (TRβ) gene putatively under selection. We illustrate the connection between genotype, phenotype and habitat variation and demonstrate that the strong differences in larval life history traits observed in the common garden experiment can result from adaptation to local pond characteristics. Remarkably, we found that haplotype variation in the TRβ gene contributes to variation in larval development time and growth rate, indicating that polymorphism in the TRβ gene is linked with the phenotypic variation among the environments. Genetic distance in neutral markers was correlated with differences in breeding time and environmental differences among the ponds, but not with geographical distance. These results demonstrate that while our study area did not exceed the scale of gene flow, ecological barriers constrained gene flow among contrasting habitats. Our results highlight the roles of strong selection and nonrandom gene flow created by phenological variation and, possibly, habitat preferences, which together maintain genetic and phenotypic divergence at a fine‐grained spatial scale.     

The influence of altitude and topography on genetic structure in the long-toed salamander (Ambystoma macrodactulym)

A primary goal of molecular ecology is to understand the influence of abiotic factors on the spatial distribution of genetic variation. Features including altitudinal clines, topography and landscape characteristics affect the proportion of suitable habitat, influence dispersal patterns, and ultimately structure genetic differentiation among populations. We studied the effects of altitude and topography on genetic variation of long-toed salamanders (Ambystoma macrodactylum), a geographically widespread amphibian species throughout northwestern North America. We focused on 10 low altitude sites (< 1200 m) and 11 high-altitude sites in northwestern Montana and determined multilocus genotypes for 549 individuals using seven microsatellite loci. We tested four hypotheses: (1) gene flow is limited between high- and low-altitude sites; and, (2) gene flow is limited among high-altitude sites due to harsh habitat and extreme topographical relief between sites; (3) low-altitude sites exhibit higher among-site gene flow due to frequent flooding events and low altitudinal relief; and (4) there is a negative correlation between altitude and genetic variation. Overall F(ST) values were moderate (0.08611; P < 0.001). Pairwise F(ST) estimates between high and low populations and a population graphing method supported the hypothesis that low-altitude and high-altitude sites, taken together, are genetically differentiated from each other. Also as predicted, gene flow is more prominent among low-altitude sites than high-altitude sites; low-altitude sites had a significantly lower F(ST) (0.03995; P < 0.001) than high altitude sites (F(ST) = 0.10271; P < 0.001). Use of Bayesian analysis of population structure (BAPS) resulted in delineation of 10 genetic groups, two among low-altitude populations and eight among high-altitude populations. In addition, within high altitude populations, basin-level genetic structuring was apparent. A nonequilibrium algorithm for detecting current migration rates supported these population distinctions. Finally, we also found a significant negative correlation between genetic diversity and altitude. These results are consistent with the hypothesis that topography and altitudinal gradients shape the spatial distribution of genetic variation in a species with a broad geographical range and diverse life history. Our study sheds light on which key factors limit dispersal and ultimately species' distributions.     

Patterns of Genetic Variation across Altitude in Three Plant Species of Semi-Dry Grasslands

Background

Environmental gradients caused by altitudinal gradients may affect genetic variation within and among plant populations and inbreeding within populations. Populations in the upper range periphery of a species may be important source populations for range shifts to higher altitude in response to climate change. In this study we investigate patterns of population genetic variation at upper peripheral and lower more central altitudes in three common plant species of semi-dry grasslands in montane landscapes.

Methodology/Principal Findings

In Briza media, Trifolium montanum and Ranunculus bulbosus genetic diversity, inbreeding and genetic relatedness of individuals within populations and genetic differentiation among populations was characterized using AFLP markers. Populations were sampled in the Swiss Alps at 1800 (upper periphery of the study organisms) and at 1200 m a.s.l. Genetic diversity was not affected by altitude and only in B. media inbreeding was greater at higher altitudes. Genetic differentiation was slightly greater among populations at higher altitudes in B. media and individuals within populations were more related to each other compared to individuals in lower altitude populations. A similar but less strong pattern of differentiation and relatedness was observed in T. montanum, while in R. bulbosus there was no effect of altitude. Estimations of population size and isolation of populations were similar, both at higher and lower altitudes.

Conclusions/Significance

Our results suggest that altitude does not affect genetic diversity in the grassland species under study. Genetic differentiation of populations increased only slightly at higher elevation, probably due to extensive (historic) gene flow among altitudes. Potentially pre-adapted genes might therefore spread easily across altitudes. Our study indicates that populations at the upper periphery are not genetically depauperate or isolated and thus may be important source populations for migration under climate change.     

Cuticular melanisation and immune response in a butterfly: local adaptation and lack of correlation

1. Despite growing knowledge on the relationship between ecological variables and individual immune function, data on the spatial variability of immune defence in invertebrate natural populations are scarce. 2. Here, we use replicated populations of the butterfly Lycaena tityrus from different altitudes to investigate genetic variation in the melanin‐based encapsulation response. As high‐ and low‐altitude populations differ in cuticular pupal melanisation, we further tested for any associations between pupal melanisation and parasite resistance. 3. Although pupal melanisation was higher at higher compared with lower altitudes (and at a higher compared with a lower rearing temperature), any obvious relations to the encapsulation response were absent. Further phenotypic correlations within groups were significant in one out of four cases only, suggesting that in L. tityrus encapsulation operates largely independent of cuticular melanisation. 4. A significant interaction between altitude and temperature indicated that high‐altitude animals show a stronger melanisation response than low‐altitude ones at the lower temperature and vice versa, indicating local adaptation to different climates.     

Phenology of Festuca pallescens in relation to topography in north-western Patagonia

Abstract. The phenological changes in populations of Festuca pallescens (St. Yves) Parodi at different topographic positions and exposure along an altitudinal gradient (600 - 1100 m) were investigated during two growing seasons in northwestern Patagonia. Stepwise multiple regression analysis was used to describe the relationship between phenology and environment during the entire growing season. Analysis of variance was also performed at each sample date to detect significant environmental factors influencing phenology at different sites. The sum of maximum air temperatures was identified as the environmental variable best correlated with the seasonal variation of phenological events of Festuca pallescens over the period of two growing seasons, explaining 93.2 % of the total variance. Significant differences between sites were observed at each sample date. Main effects of altitude and topographic position and two-way interactions between altitude and topographic position, and topographic position and exposure were also detected as significant. Phenology was delayed at increased altitude. Differences in phenology between topographic sites at the same altitude were not detected during the entire growing season and were only observed in the reproductive phase. At this time, the phenology was significantly delayed at high topographic positions on the slopes as compared with low and mid positions. At high altitudes in the valley (950 m a. s. 1.), where steep slopes and humid conditions prevail, phenology was delayed on western exposures and low positions. The results adequately summarize and quantify the effect of spatial and temporal environmental variation on the phenological development of Festuca pallescens in northwestern Patagonia.     

Natural selection favors rapid reproductive phenology in Potentilla pulcherrima (Rosaceae) at opposite ends of a subalpine snowmelt gradient

In high altitude plants, flowering quickly ensures reproductive success within a short snow-free period, but limits maturation time and fecundity. Natural selection on prefloration intervals may therefore vary in contrasting snowmelt environments and could influence the outcome of phenological responses to climatic change. This study investigated adaptive differentiation and plasticity of prefloration intervals in the subalpine perennial Potentilla pulcherrima. Three years of in situ field observations were combined with phenotypic selection analyses and a common garden experiment. Plants from high, intermediate, and low altitudes expressed similar prefloration intervals and plasticity when grown at common altitude, indicating no evidence for adaptive differentiation. Selection on the prefloration interval was negative at both low and high altitudes before and after accounting for strong positive selection on size. Environmental differences between high and low altitudes indicated that long, dry seasons and short, wet seasons both favored rapid reproduction. Therefore, quicker reproduction was adaptive in response to late snowmelt, but slower reproduction in response to earlier snowmelt appeared to be maladaptive. Selection differed marginally between late snowmelt years and dry ones. Plastic responses to future precipitation patterns may therefore have positive or negative effects on fitness within a single species, depending upon altitude and year.     

Nitric oxide in adaptation to altitude

This review summarizes published information on the levels of nitric oxide gas (NO) in the lungs and NO-derived liquid-phase molecules in the acclimatization of visitors newly arrived at altitudes of 2500 m or more and adaptation of populations whose ancestors arrived thousands of years ago. Studies of acutely exposed visitors to high altitude focus on the first 24-48 h with just a few extending to days or weeks. Among healthy visitors, NO levels in the lung, plasma, and/or red blood cells fell within 2h, but then returned toward baseline or slightly higher by 48 h and increased above baseline by 5 days. Among visitors ill with high-altitude pulmonary edema at the time of the study or in the past, NO levels were lower than those of their healthy counterparts. As for highland populations, Tibetans had NO levels in the lung, plasma, and red blood cells that were at least double and in some cases orders of magnitude greater than other populations regardless of altitude. Red blood cell-associated nitrogen oxides were more than 200 times higher. Other highland populations had generally higher levels although not to the degree shown by Tibetans. Overall, responses of those acclimatized and those presumed to be adapted are in the same direction, although the Tibetans have much larger responses. Missing are long-term data on lowlanders at altitude showing how similar they become to the Tibetan phenotype. Also missing are data on Tibetans at low altitude to see the extent to which their phenotype is a response to the immediate environment or expressed constitutively. The mechanisms causing the visitors' and the Tibetans' high levels of NO and NO-derived molecules at altitude remain unknown. Limited data suggest processes including hypoxic upregulation of NO synthase gene expression, hemoglobin-NO reactions, and genetic variation. Gains in understanding will require integrating appropriate methods and measurement techniques with indicators of adaptive function under hypoxic stress.