珠穆朗玛峰自然保护区小叶金露梅灌丛群落数量分类和排序及其多样性垂直格局

根据野外样方调查数据,采用双向种指示分析(TWINSPAN)和典范对应分析(CCA),对珠穆朗玛峰国家级自然保护区小叶金露梅灌丛群落进行分类和排序,并分析物种多样性沿海拔梯度的分布格局。结果表明:(1)该区域24个样地中,记载的维管束植物共有23科45属80种,出现频度较高的种有小叶金露梅(Potentilla parvifolia)、高山嵩草(Kobresia pygmaea)、木根香青(Anaphalis xylorhiza)、垫状点地梅(Androsace tapete)、藏沙蒿(Artemisia wellbyi)、垫状雪灵芝(Arenaria pulvinata)和柴胡红景天(Rhodiola bupleuroides)等。(2)经TWINSPAN等级分类将该区域小叶金露梅灌丛24个样地划分为10个群丛类型。(3)样地和物种CCA二维排序结果表明,海拔和坡位是影响该区域小叶金露梅灌丛群落和物种分布格局的主要环境因子。(4)该区域小叶金露梅灌丛群落物种丰富度、Shannon-Wiener指数和Simpson指数均随海拔升高呈下降的趋势,而Pielou指数呈上升的趋势。(5)样地中优势种小叶金露梅的盖度和高度沿海拔梯度呈显著下降趋势。     

中国产委陵菜属叶表皮毛的研究

在光学显微镜和实体解剖镜下对28种和变种国产委陵菜属植物的叶表皮毛进行了研究,结果如下：1．在所观察的植物中,有4种类型叶表皮毛的存在,即：单细胞直的不分支的毛;单细胞卷曲的不分支的毛;腺毛;星状毛。2．叶表皮毛的种类在本属植物中具有稳定性,因此具有一定的分类学价值。3．腺毛在所观察的植物中普遍存在。4．讨论了金露梅、银露梅、小叶金露梅的系统归属。     

国产委陵菜属植物果实形态及其分类学意义

对国产委陵菜属Ｐｏｔｅｎｔｉｌｌａ３２种及变种植物的果实形态在实体解剖镜和扫描电镜下进行了研究。结果表明：在扫描电镜下,本属植物果实表面纹饰有５种不同的类型,即网纹型,疣状突起型,波纹型,帽状突起型和不规则型;果实形态特征,特别是电镜下的果皮表面纹饰对本属植物的分类具有较重要价值;根据果实形态,支持将金露梅、银露梅、小叶金露梅做为独立的种。     

祁连山金露梅灌丛草甸群落结构及主要种群的点格局分析

金露梅灌丛草甸是青藏高原高寒草地生态系统的重要组分之一,采用Ripley的K(t)函数分析方法,对放牧和封育条件下主要种群的分布格局和种间关系及金露梅灌丛群落动态变化进行了研究.结果表明:(1)经过6年的围栏封育,金露梅灌丛草甸主要种群的空间分布格局表现出从均匀或随机分布向随机或聚集分布转变的趋势,聚集强度随围封尺度增大而增加,随放牧的干扰而降低,聚集分布有利于种群发挥功能群整体效应.(2)围栏内不同功能群之间主要呈显著正关联;围栏外的禾草与杂类草、莎草与杂类草之间均呈显著负关联,出现种间资源竞争,且随着尺度的增大负关联逐渐增强,不同物种的生态位差异明显.(3)围栏内外种群点格局分布都具有明显的尺度依赖性,植物间相互作用和外界放牧干扰是影响物种不同尺度下空间格局变化的主要原因,点格局的变化驱动了群落结构的变化.可见,长期夏季放牧抑制了高寒金露梅灌丛草甸禾草和莎草植物的分株和生长,促进了杂类草的生长,金露梅灌丛草甸呈现出退化的趋势,围栏封育有利于草地生态系统的恢复演替.     

亚高寒草甸不同坡向金露梅种群的空间分布格局及空间关联

植物种群在不同干扰条件下对环境的适应对策可以通过其空间分布格局及关联性反映出来.金露梅是甘南亚高寒草甸中的优势种,通过野外群落调查,运用空间点格局分析的Ripley K函数,分析甘南不同坡向亚高寒草甸中金露梅种群的空间分布格局及其关联性.结果表明: 金露梅种群在3个坡向的分布具有显著差异,重要值、生物量和盖度在北坡均高于南坡;北坡和西坡金露梅种群Ⅰ、Ⅱ级个体在小尺度上呈显著聚集分布,随着龄级和空间尺度的增加,金露梅种群聚集强度逐渐减弱,趋向于随机分布;南坡由于其生境条件对金露梅生长的抑制,种间竞争激烈,各级金露梅种群的分布格局均以随机分布为主;不同坡向的金露梅种群Ⅰ、Ⅱ级个体之间在一定尺度内表现为负关联,随着尺度增加,关联度降低,Ⅲ、Ⅳ级与Ⅰ、Ⅱ级个体之间的关联性呈显著负关联;较大个体之间无明显关系,表明年龄接近的个体在空间分布上是相互独立的,利于其充分利用环境资源.     

青海高寒区域金露梅灌丛草甸灌木和草本植物固碳量的比较

以青海海北高寒区域金露梅（PotentillafruticosaLinn.）灌丛草甸为研究对象,分析了6月至9月金露梅灌丛草甸灌木和草本植物不同部位的生物碳量,并据此对灌木及草本植物的年净初级生产碳量进行了比较。结果显示：金露梅灌丛草甸灌木植物地上部和地下部不同层次的生物量和碳含量均有明显差异,根据生物量所占比例确定其地上部和地下部的平均碳含量分别为0.50和0.48。依据不同月份灌丛冠面最大长度、最小宽度和最大高度,采用方程“Wij=e〔aln（A·B·H）＋b〕”计算灌木地上当年新生生物碳量、地上多年累积生物碳量和地下多年累积生物碳量,相关性均极显著（P〈0.01）,表明利用该方程评估金露梅灌丛草甸灌木不同部位的生物碳量是可行的。不同月份金露梅灌丛草甸灌木地上当年新生生物碳量、地上多年累积生物碳量和地下多年累积生物碳量分别为9.36-21.15、78.07-90.12和74.37-101.22g·m-2,差异不明显;其地上部和地下部净初级生产碳量分别为33.20和26.85g·m-2,总计为60.05g·m-2。金露梅灌丛草甸草本植物地上部和地下部净初级生产碳量分别为111.41和445.41g·m-2,总计为556.82g·m-2。如果根据草本和灌木占地面积78%和22%进行加权计算,则金露梅灌丛草甸当年的总净初级生产碳量为447.53g·m-2,其中灌木的净初级生产碳量仅占2.95%,且金露梅灌丛草甸地下部与地上部净初级生产碳量的比值为3.75。研究结果显示：青海高寒区域金露梅灌丛草甸以草本固碳为主,且地下部净初级生产碳量明显高于其地上部。     

HPLC测定不同海拔小叶金露梅17种氨基酸含量

目的:建立高效液相色谱法测定小叶金露梅中17种氨基酸含量的方法。方法:采用N,N-二硝基氟苯柱前衍生氨基酸后测定;色谱柱为Phenomenex Gemini 5μC18,流动相为醋酸钠(pH=6.40)和50%乙腈,采用梯度洗脱,检测波长为360nm,外标法计算含量。结果:17种氨基酸线性关系、精密度、稳定性、重复性良好,加样回收率为97.4%～102.8%(RSD为1.21%～2.5%)。各海拔的氨基酸总量范围是10%～14.79%,在不同海拔小叶金露梅中氨基酸平均质量分数顺序为亮氨酸>天冬氨酸>异亮氨酸>组氨酸>丝氨酸>精氨酸>丙氨酸>苯丙氨酸>缬氨酸>谷氨酸>脯氨酸>甘氨酸>赖氨酸。小叶金露梅含有7种必需氨基酸,它们的含量均大于48%。本方法简便、准确,为小叶金露梅药理提供理论依据。     

基于Ripley K(r)函数的金露梅与蒲公英种群空间分布格局及其关联性

物种空间分布格局和种间关联性是研究种群特征、种间相互影响以及物种同环境关系的重要方法。金露梅(Potentilla fruticosa L.)与蒲公英(Taraxacum mongolicum Hand.-Mazz.)是甘南亚高寒草甸的主要物种,通过野外群落调查,应用点格局方法中的Ripley K函数,分析不同坡向的金露梅与蒲公英种群的空间分布格局及种间关联性。结果表明:金露梅与蒲公英在3个坡向上的分布具有明显差异,随着坡向由北坡向南坡的转变,金露梅种群呈现递减趋势,蒲公英种群则呈现出递增趋势;在北坡,金露梅在0~4.8 m内呈现聚集分布且聚集程度比较高,在4.8~5 m呈现随机分布;在西坡,金露梅在0~0.6 m尺度内表现出随机分布,在0.6~1.6 m尺度内为聚集分布,在1.6~5 m尺度内则又再次呈现随机分布;而在南坡,金露梅在整个研究尺度范围内均呈现随机分布;北坡的蒲公英在0~2.8 m尺度上呈现聚集分布,在2.8~5 m内呈现随机分布;西坡和南坡上蒲公英在整个研究尺度内都呈现随机分布;北坡的金露梅与蒲公英在整个研究尺度内呈现负关联,西坡两种群在整个尺度内呈现无关联;南坡的金露梅与蒲公英在0~2.4 m尺度内呈现无关联,在2.4~5 m尺度内呈现负关联。     

五种西北旱区植物对干旱胁迫的生理响应

以黑果枸杞、金露梅、蒙古莸、柠条和花棒5种西北旱区植物为材料,采用盆栽试验研究了干旱胁迫对植物叶片光合、水分和生理生化的影响.结果表明: 随干旱胁迫时间的延长,5种植物叶片的保水力均升高,叶片相对含水量先升后降,叶绿素含量和叶绿素荧光参数均不同程度地降低,花棒和黑果枸杞的光合系统受到的影响最小,金露梅的光合系统受损最严重.干旱胁迫后金露梅叶片的电导率和丙二醛含量先降低后急剧上升,超氧化物歧化酶(SOD)和过氧化物酶(POD)活性先升高后急剧降低.其余4种植物随土壤含水量的降低电导率先升后降,丙二醛含量先升后降或先降后升,SOD和POD活性总体呈上升趋势,其中蒙古莸SOD活性和花棒POD活性在所有植物中升幅最大,干旱28 d后分别升至为胁迫前的2.1、14.1倍.随胁迫时间的延长,5种植物各有机溶质的积累量存在较大差异,金露梅积累最多的可溶性糖和游离脯氨酸,28 d后分别增至为原来的2.1、23.4倍,花棒积累最多的可溶性蛋白,28 d后增至为原来的2.0倍.隶属函数法表明,植物耐旱能力为花棒>黑果枸杞>柠条>蒙古莸>金露梅.5种旱区植物可通过提高抗氧化酶活性和积累有机溶质对干旱胁迫进行积极的反馈,以减轻细胞受到的伤害.金露梅对水分亏缺的敏感性最高,试验期间其干旱程度已经超出了自身调节能力的阈值.     

放牧影响下不同盖度金露梅灌丛草本植物叶功能性状与土壤因子的关系

植物功能性状与环境因子的研究对认识不同环境梯度下植物群落的形成及对环境的适应机制具有重要意义。该研究以高寒金露梅(Potentilla fruticosa)灌丛草甸为研究对象,分析放牧影响下金露梅灌丛不同盖度梯度(金露梅灌丛盖度为0%、30%、60%和90%的灌草斑块)灌草斑块中草本植物叶功能性状与土壤因子的变化趋势及其相互关系。结果表明:(1)土壤容重(BD)、土壤pH和土壤养分在金露梅灌丛盖度梯度下差异显著,干柴滩金露梅灌丛具有明显的"沃岛效应";随着金露梅灌丛盖度的递增,土壤理化性质变化趋势各有分异。(2)随着金露梅灌丛盖度的增加,比叶面积(SLA)显著上升,叶片厚度(LT)和叶片磷含量(LPC)呈显著下降趋势(P0.05)。(3)比叶面积与叶片厚度呈负相关关系,与叶片氮磷含量(LNC、LPC)呈极显著负相关关系(P0.01),与叶片碳含量(LCC)呈极显著正相关关系(P0.01)。(4)RDA冗余分析表明,土壤因子对叶功能性状变异的平均解释变量达72.25%,影响植物功能性状的主导因子有土壤有机质(SOM)、土壤全氮(TN)、土壤有效氮(AN)、土壤有效钾(AK)和土壤容重(BD),但不同盖度金露梅灌丛群落中影响叶性状的主导因子存在差异。金露梅灌丛盖度较小(30%)时受SOM和TN、AN的影响较大,而金露梅灌丛盖度较大时(60%)时主要受AK和BD的影响。     

Short-term responses of phenology, shoot growth and leaf traits of four alpine shrubs in a timberline ecotone to simulated global warming, Eastern Tibetan Plateau, China

The short-term effects of artificial warming on phenology, growth and leaf traits were investigated in four alpine shrubs using the open-top chamber (OTC) method in a timberline ecotone (3240 m a.s.l.) on the Eastern Tibetan Plateau. The OTC enhanced the mean air temperature by 2.9°C throughout the growing season. In contrast, only a slight difference (0.4°C) in the mean soil temperature was observed in the OTC compared with the control plots (CP). Spiraea mongolica , Potentilla fruticosa , Conicera hispida (deciduous shrubs) and Daphne retusa (evergreen shrub) showed earlier bud break, flowering and fruit coloring as well as longer flower longevity in the OTC than in the CP. All deciduous shrubs in the OTC had a longer leaf lifespan. Daphne retusa had higher leaf survival rates in the OTC. No significant differences in the total number of flowers and fruits were noticed for most species between the two treatments. Warming stimulated the shoot and leaf growth for most species. The specific leaf area tended to increase for many species in the OTC. However, the leaf nitrogen concentration tended to decrease in P. fruticosa and S. mongolica. The results obtained in the present study indicate that warming conditions can have strong impacts on alpine shrubs in a timberline ecotone.     

Methane emissions by alpine plant communities in the Qinghai-Tibet Plateau

For the first time to our knowledge, we report here methane emissions by plant communities in alpine ecosystems in the Qinghai-Tibet Plateau. This has been achieved through long-term field observations from June 2003 to July 2006 using a closed chamber technique. Strong methane emission at the rate of 26.2+/-1.2 and 7.8+/-1.1microg CH4 m-2h-1 was observed for a grass community in a Kobresia humilis meadow and a Potentilla fruticosa meadow, respectively. A shrub community in the Potentilla meadow consumed atmospheric methane at the rate of 5.8+/-1.3microg CH4 m-2h-1 on a regional basis; plants from alpine meadows contribute at least 0.13Tg CH4 yr-1 in the Tibetan Plateau. This finding has important implications with regard to the regional methane budget and species-level difference should be considered when assessing methane emissions by plants.     

青藏高原地区刺山柑解剖特征研究

采用石蜡切片法对分布于海拔4 000 m的西藏札达的刺山柑解剖结构进行研究,进一步探讨青藏高原植物独特生理结构及其与环境的适应性。结果表明刺山柑具有适应高山荒漠草原环境的典型结构特征: 次生根周皮发达,具有较厚的木栓层和发达的次生木质部;茎具表皮毛和角质层,皮层较厚,厚角组织发达,髓较小;叶具角质层,气孔密集,栅栏组织多层,为双栅型等面叶;花冠较大,白色花瓣,适于虫媒传粉;侧膜胎座,子房多室,胚珠多数,花粉外壁较厚,具较强的抗腐蚀及抗酸碱性能。刺山柑形成上述结构特征是青藏高原特殊综合生态环境长期作用的结果。同时也是刺山柑对高原环境的高度适应。     

Ecological responses of Primula nutans to centimeter-scale topographic and environmental variability in an alpine wetland

Primula nutans Georgi is a herbaceous species broadly distributed in wetlands on the Qinghai–Tibetan Plateau. These wetlands are often spatially highly heterogeneous because of their hummock-and-hollow microtopography. To address how P. nutans can be so broadly distributed on these wetlands, we examined ramet distribution, abundance, and growth performance in environmentally variable microsites at the centimeter scale. P. nutans showed significantly higher ramet density on the south-orientated microsites than on the north-orientated microsites. With increasing relative microsite elevation on the hummocks, ramet density increased significantly, but individual leaf area decreased significantly. Principle component analysis and multiple regression analysis indicated that microsites at higher elevations tend to have higher light availability and higher ramet density. The study suggests that P. nutans shows high plasticity in the distribution, abundance, and growth performance in response to the microtopography at the centimeter scale, which may contribute to the broad distribution of the species in the hummock-and-hollow wetlands in harsh alpine environments on the Tibetan Plateau.     

Population history and genomic signatures for high-altitude adaptation in Tibetan pigs

Background

The Tibetan pig is one of domestic animals indigenous to the Qinghai-Tibet Plateau. Several geographically isolated pig populations are distributed throughout the Plateau. It remained an open question if these populations have experienced different demographic histories and have evolved independent adaptive loci for the harsh environment of the Plateau. To address these questions, we herein investigated ~ 40,000 genetic variants across the pig genome in a broad panel of 678 individuals from 5 Tibetan geographic populations and 34 lowland breeds.

Results

Using a series of population genetic analyses, we show that Tibetan pig populations have marked genetic differentiations. Tibetan pigs appear to be 3 independent populations corresponding to the Tibetan, Gansu and Sichuan & Yunnan locations. Each population is more genetically similar to its geographic neighbors than to any of the other Tibetan populations. By applying a locus-specific branch length test, we identified both population-specific and -shared candidate genes under selection in Tibetan pigs. These genes, such as PLA2G12A, RGCC, C9ORF3, GRIN2B, GRID1 and EPAS1, are involved in high-altitude physiology including angiogenesis, pulmonary hypertension, oxygen intake, defense response and erythropoiesis. A majority of these genes have not been implicated in previous studies of highlanders and high-altitude animals.

Conclusion

Tibetan pig populations have experienced substantial genetic differentiation. Historically, Tibetan pigs likely had admixture with neighboring lowland breeds. During the long history of colonization in the Plateau, Tibetan pigs have developed a complex biological adaptation mechanism that could be different from that of Tibetans and other animals. Different Tibetan pig populations appear to have both distinct and convergent adaptive loci for the harsh environment of the Plateau.

Electronic supplementary material

The online version of this article (doi:10.1186/1471-2164-15-834) contains supplementary material, which is available to authorized users.     

2种珍稀裂腹鱼类线粒体DNA部分序列片段的比较分析

对扁吻鱼(Aspiorhynchus laticeps)及塔里木裂腹鱼(Schizothorax biddulphi)线粒体CO I(624 bp)、Cyt b(712 bp)和D-loop(457 bp)基因片段进行了PCR扩增和序列测定.通过分析比较发现,3个片段A+T含量与其他鱼类相比均较高,G含量偏低这一现象在...     

雌性藏羚迁徙对青藏高原降水时空分布的适应性分析

藏羚主要分布在青藏高原,雌性有季节性的生殖迁徙行为。产仔前,分布在各地的雌性藏羚沿着比较固定的路线向青藏高原西北部可可西里的太阳湖、卓乃湖、阿尔金山保护区的慕孜塔格峰等产羔地迁徙,产羔后一段时间又返回原栖息地。对这种特殊的迁徙现象,已有一些假说用来解释,但都没有得到普遍认可。青藏高原由于其海拔高,常年温度低,在藏羚分布区降水主要以固态形式为主,并且青藏高原降水量由东南部向西北部递减,结合青藏高原地质历史时期植被演变的特点和藏羚迁徙的方向,本文提出了一个新假设,以此来解释藏羚迁徙的原因。我们的假设是:藏羚迁往产羔地产羔是为了躲避固态降水相对丰富的地区,以保证新生羚羊有高的存活率;藏羚迁往的产羔地曾是它们的故乡。笔者主要分析了提出这个假设的根据。     

Maternal investment increases with altitude in a frog on the Tibetan Plateau

Reproducing females can allocate energy between the production of eggs or offspring of different size or number, both of which can strongly influence fitness. The physical capacity to store developing offspring imposes constraints on maximum clutch volume, but individual females and populations can trade off whether more or fewer eggs or offspring are produced, and their relative sizes. Harsh environments are likely to select for larger egg or offspring size, and many vertebrate populations compensate for this reproductive investment through an increase in female body size. We report a different trade‐off in a frog endemic to the Tibetan Plateau, Rana kukunoris. Females living at higher altitudes (n = 11 populations, 2000–3500 m) produce larger eggs, but without a concomitant increase in female body size or clutch size. The reduced diel and seasonal activity at high altitudes may impose constraints on the maximum body size of adult frogs, by limiting the opportunity for energy accumulation. Simultaneously, producing larger eggs likely helps to increase the rate of embryonic development, causing tadpoles to hatch earlier. The gelatinous matrix surrounding eggs, more of which is produced by large females, may help buffer developing embryos from temperature fluctuations or offer protection from ultraviolet radiation. High‐altitude frogs on the Tibetan Plateau employ a reproductive strategy that favours large egg size independent of body size, which is unusual in amphibians. The harsh and unpredictable environmental conditions at high altitudes can thus impose strong and opposing selection pressures on adult and embryonic life stages, both of which can simultaneously influence fitness.     

Mitochondrial phylogeography and genetic diversity of Tibetan gazelle (Procapra picticaudata): implications for conservation

The Tibetan gazelle (Procapra picticaudata) is a threatened species and distributed on the Qinghai-Tibet Plateau of China (Qinghai Province, Tibet Autonomous Region and the adjacent Gansu Province, Sichuan Province, and Xinjiang Uigur Autonomous Region). Small peripheral populations of Tibetan gazelle were once found in northern Sikkim and Ladakh, but now these are close to extinction. To describe the evolutionary history and to assess the genetic diversity within this monotypic species and population structure among different geographic locations in China, we sequenced mitochondrial DNA from the control region (CR) and cytochrome (cyt) b gene for 46 individuals from 12 geographic localities in Qinghai, Tibet, Xinjiang, Gansu, and Sichuan. A total of 25 CR haplotypes and 16 cyt b haplotypes were identified from these gazelle samples. CR haplotype diversity (0.98+/-0.01) and nucleotide diversity (0.08+/-0.009) were both high. Phylogenetic trees indicate that the Tibetan gazelle in China can be divided into three main clades: Tibet, Sichuan (SCH) and Qinghai-Arjin Shan-Kekexili (QH-ARJ-KKXL). Analysis of molecular variance (AMOVA) and network analysis consistently support this geographic structure in both datasets. Significant differentiation between populations argues for the presence of management units (MUs). Such differentiation may reflect a geographic separation resulting from the uplift of the Qinghai-Tibet Plateau during the Late Pliocene and Pleistocene. Mismatch distribution analysis implies that Tibetan gazelle has undergone complex population changes. We suggest that the present population structure has resulted from habitat fragmentation during the recent glacial period on the Qinghai-Tibet Plateau and population expansion from glacial refugia after the glacial period. It is likely that the present populations of Tibetan gazelle exhibit a pattern reminiscent of several bottlenecks and expansions in the recent past.     

Natural selection and adaptive evolution of leptin in the ochotona family driven by the cold environmental stress

Background

Environmental stress can accelerate the evolutionary rate of specific stress-response proteins and create new functions specialized for different environments, enhancing an organism''s fitness to stressful environments. Pikas (order Lagomorpha), endemic, non-hibernating mammals in the modern Holarctic Region, live in cold regions at either high altitudes or high latitudes and have a maximum distribution of species diversification confined to the Qinghai-Tibet Plateau. Variations in energy metabolism are remarkable for them living in cold environments. Leptin, an adipocyte-derived hormone, plays important roles in energy homeostasis.

Methodology/Principal Findings

To examine the extent of leptin variations within the Ochotona family, we cloned the entire coding sequence of pika leptin from 6 species in two regions (Qinghai-Tibet Plateau and Inner Mongolia steppe in China) and the leptin sequences of plateau pikas (O. curzonia) from different altitudes on Qinghai-Tibet Plateau. We carried out both DNA and amino acid sequence analyses in molecular evolution and compared modeled spatial structures. Our results show that positive selection (PS) acts on pika leptin, while nine PS sites located within the functionally significant segment 85-119 of leptin and one unique motif appeared only in pika lineages-the ATP synthase α and β subunit signature site. To reveal the environmental factors affecting sequence evolution of pika leptin, relative rate test was performed in pikas from different altitudes. Stepwise multiple regression shows that temperature is significantly and negatively correlated with the rates of non-synonymous substitution (Ka) and amino acid substitution (Aa), whereas altitude does not significantly affect synonymous substitution (Ks), Ka and Aa.

Conclusions/Significance

Our findings support the viewpoint that adaptive evolution may occur in pika leptin, which may play important roles in pikas'' ecological adaptation to extreme environmental stress. We speculate that cold, and probably not hypoxia, may be the primary environmental factor for driving adaptive evolution of pika leptin.