941 resultados para Asia, Central
Resumo:
近十年,植物群体遗传学的研究飞速发展,然而与海拔相关的植物群体遗传结构和遗传变异研究却相对较少。到目前为止,还不清楚遗传变异与海拔之间是否有一个通用的格局。在山区,各种生态因子,如温度、降水、降雪、紫外线辐射强度以及土壤成分都随海拔梯度急剧变化,造成了即使在一个小的空间区域,植被类型变化显著,这种高山环境的异质性和复杂性为我们研究植物群体遗传结构和分化提供了方便。沙棘(Hippophea)属于胡颓子科(Elaeagnaceae)为多年生落叶灌木或乔木,雌雄异株,天然种群分布极为广泛。中国沙棘(H. rhamnoides subsp. sinensis)是沙棘属植物中分布较广的一个亚种,种内形态变异非常丰富,加之其具有独特的繁育系统和广泛的生态地理分布,是研究沙棘属植物遗传变异和系统分化的理想材料。本文从1,800 m 到3,400 m 分5 个海拔梯度进行取样,用RAPD 和cpSSR 分子标记研究了卧龙自然保护区中国沙棘天然群体的遗传结构和遗传变异。5 个取样群体依次标记为A、B、C、D 和E,它们分别代表分布在海拔1,800,2,200,2,600,3,000 和3,400 m 的5 个天然群体。RAPD实验用11 条寡核苷酸引物,扩增得到151 个重复性好的位点,其中143 个多态位点,多态率达94.7%。在5 个沙棘群体中,总遗传多样性值(HT)为0.289,B群体内的遗传多样性值为0.315,这完全符合沙棘这种多年生、远交的木本植物具有高遗传变异的特性。5 个群体内遗传多样性随海拔升高呈低-高-低变异趋势,在2,200 m海拔处的B群体遗传多样性达最大值0.315,3,400 m海拔处的E群体则表现最小仅0.098。5 个群体间的遗传分化值GST=0.406,也即是说有40.6%的遗传变异存在于群体间,1,800 m海拔处的A群体与其它群体的明显分离是造成群体间遗传分化大的原因。UPGMA聚类图和PCoA散点图进一步确证了5 个群体间的关系和所有个体间的关系。最后,经过Mantel检测,遗传距离与海拔表现了明显的相关性(r = 0.646, P = 0.011)。cpSSR 实验中,经过对24 对cpSSR 通用引物筛选,11 对引物能扩增出特异性条带,只有2 对引物(ccmp2 和ARCP4)呈现多态性。4 个等位基因共组合出4 种单倍型,单倍型Ⅰ出现在A 群体的所有个体和B 群体的8 个个体中,C、D、E 三个群体均不含有,而单倍型Ⅱ出现在C、D、E 三个群体的所有个体及B 群体的18 个个体中,A 群体不含有。另外两种单倍型Ⅲ和Ⅳ为稀有类型,仅B 群体中的4 个个体拥有。这种单倍型分布模式和TFPGA 群体聚类图揭示了,C、D、E 群体可能来源于同一祖先种,而A 群体却是由另一祖先种发展起来的,B 群体则兼具了这两种起源种的信息,这可能是因为在历史上的某一时期,在中国沙棘群体高山分化的过程中,B 群体处某个或者某些个体发生了基因突变,具备了适应高海拔环境的能力,产生了高海拔沙棘群体的祖先种。 In recent ten years, studies about population genetics of plants developed rapidly,whereas their genetic structure and genetic variation along altitudinal gradients have beenstudied relatively little. So far, it is uncleared whether there is a common pattern betweengenetic variation and altitudinal gradients. In the mountain environments, importantecological factors, e.g., temperature, rainfall, snowfall, ultraviolet radiation and soil substratesetc., change rapidly with altitudes, which cause the vegetation distribution varying typically,even on a small spatial scale. The mountain environments, which are heterogeneous andcomplex, facilitate and offer a good opportunity to characterize population genetic structureand population differentiation.The species of the genus Hippophae L. (Elaeagnaceae) are perennial deciduous shrubs ortrees, which are dioecious, wind-pollinated pioneer plants. The natural genus has a widedistribution extending from Northern Europe through Central Europe and Central Asia toChina. According to the latest taxonomy, the genus Hippophae is divided into six species and12 subspecies. The subspecies H. rhamnoides ssp. sinensis shows significant morphologicalvariations, large geographic range and dominantly outcrossing mating system. Thesecharacteristics of the subspecies are favourable to elucidate genetic variation and systemevolution. To estimate genetic variation and genetic structure of H. rhamnoides ssp. sinensisat different altitudes, we surveyed five natural populations in the Wolong Natural Reserve at altitudes ranging from 1,800 to 3,400 m above sea level (a.s.l.) using random amplifiedpolymorphic DNA markers (RAPDs) and cpSSR molecular methods. The five populations A,B, C, D, and E correspond to the altitudes 1,800, 2,200, 2,600, 3,000 and 3,400 m,respectively.Based on 11 decamer primers, a total of 151 reproducible DNA loci were yielded, ofwhich 143 were polymorphic and the percentage of polymorphic loci equaled 94.7%. Amongthe five populations investigated, the total gene diversity (HT) and gene diversity within population B equaled 0.289 and 0.315, respectively, which are modest for a subspecies of H.rhamnoides, which is an outcrossing, long-lived, woody plant. The amount of geneticvariation within populations varied from 0.098 within population E (3,400 m a.s.l.) to 0.315within population B (2,200 m a.s.l.). The coefficient of gene differentiation (GST) amongpopulations equaled 0.406 and revealed that 40.6% of the genetic variance existed amongpopulations and 59.4% within populations. The population A (1,800 m a.s.l.) differed greatlyfrom the other four populations, which contributes to high genetic differentiation. A UPGMAcluster analysis and principal coordinate analyses based on Nei's genetic distances furthercorroborated the relationships among the five populations and all the sampling individuals,respectively. Mantel tests detected a significant correlation between genetic distances andaltitudinal gradients (r = 0.646, P = 0.011).Eleven of the original 24 cpSSR primer pairs tested produced good PCR products, onlytwo (ccmp2 and ARCP4) of which were polymorphic. Four total length variants (alleles) werecombined resulting in 4 haplotypes. The haplotype was present in all individuals of Ⅰpopulation A and 8 individuals of populations B, the other three populations (C, D and Epopulations) did not share. The haplotype was present in all individuals of populations C, D Ⅱand E and 18 individuals of populations B, population A did not share. The other twohaplotypes and were rare haplotypes, which were only shared in 4 individuals of Ⅲ Ⅳpopulation B. The distribution of haplotypes and TFPGA population clustering map showedthat the populations C, D and E might be origined from one ancestor seed and population Amight be from another, whereas population B owned information of the two ancestor seeds. Itwas because that gene mutation within some individual or seed in the location of population Bwas likely to happen in the history of H. rhamnoides, which was the original ancestor of thehigh-altitude populations.
Resumo:
角蟾科(Megophryidae)是以角蟾属(Megophrys Kuhl and Van Hasselt, 1822)为模式属而建立的,隶于无尾目(Anura),变凹型亚目(Anomocoela)。角蟾科包括2 亚科11 属142 种,分布于东洋界,从巴基斯坦、中国西部向东直到菲律宾和苏达群岛;中国有9 属75 种分布于华中和华南地区。角蟾科被认为是原始的两栖动物之一,其分类学、系统学、生态学、动物地理学的研究均深受中外科学家的瞩目。近年来,通过形态学、古生物学、细胞学、生态学、支序系统学的研究,角蟾科的分类与系统学研究取得了较大进展。与成体形态和分子系统学研究结果相比较,蝌蚪的研究存在更多的问题和挑战,尚需深入研究:(1)角蟾科蝌蚪的形态多样性分析;(2)角蟾科的系统发育关系与蝌蚪的演化,以及口漏斗的起源;(3)角蟾科蝌蚪表型分化与栖息环境和觅食行为的适应演化。针对上述问题,本文对角蟾科9 属30 种蝌蚪的形态特征,包括外部宏观形态和口器外部结构特征、口器内部显微结构、唇齿和角质颌的亚显微结构作了深入细致、多层次的比较研究;通过12s rRNA 和cytochrome b 基因构建最大简约树,采用贝叶斯系统发育进行分析,蝌蚪型的演化采用祖先性状的重建方法分析;得到如下结论:1)初步将角蟾科蝌蚪分为4 种类型;并且建立了2 种新的角蟾科蝌蚪类型。A 型:拟髭蟾型蝌蚪,该型蝌蚪包括拟髭蟾属、髭蟾属、齿蟾属和齿突蟾属的物种;B 型:新类型,掌突蟾型蝌蚪,该型蝌蚪在本文中包括掌突蟾属、小臂蟾属的物种;C 型:新类型,短腿蟾型蝌蚪,一种特化类型,该型蝌蚪在本文中仅包括短腿蟾属的物种;D 型:角蟾型蝌蚪,该型蝌蚪在本文中包括无耳蟾属、小口拟角蟾属和异角蟾属的物种。2)对角蟾科的分类进行了修订:(1)支持角蟾科两个亚科的分类系统;(2)角蟾亚科包括拟角蟾属、异角蟾属、无耳蟾属和短腿蟾属;该亚科形态差异小,系统学关系比较复杂,暂不作族级分类的再划分;(3)拟髭蟾亚科分为2 个族:拟髭蟾族,该族物种具有类型A 的蝌蚪,包括4 个属:拟髭蟾属、髭蟾属、齿蟾属、齿突蟾属;掌突蟾族,该族物种具有类型B 的蝌蚪,包括2 个属:掌突蟾属和小臂蟾属。3)结合分子系统进化关系探讨了4 种蝌蚪类型的演化。(1)角蟾科蝌蚪的最近共同祖先来自于一类具有拟髭蟾型蝌蚪性状的蝌蚪;(2)掌突蟾型蝌蚪和角蟾亚科的蝌蚪是由具有拟髭蟾型蝌蚪性状的祖先蝌蚪分别演化而来;(3)短腿蟾型蝌蚪是角蟾型蝌蚪的一种特化类型;(4)外群蝌蚪具有与拟髭蟾型蝌蚪相似的性状,进一步印证了类拟髭蟾型蝌蚪是角蟾科蝌蚪的最近共同祖先的假说;(5)具有口漏斗的蝌蚪类型是由不具口漏斗的蝌蚪类型演化而来,在角蟾科中口漏斗是一种衍生性状。4)分析了角蟾科四种蝌蚪类型与栖息环境的适应演化。(1)角蟾科蝌蚪的口部和体形的变化反映了该科蝌蚪由缓流向类似静水生境的回水凼的渐变式适应,角蟾科蝌蚪的形态显示了多方面的适应变化;(2)随着蝌蚪类型由A 向D的演化,当水速较大时,拟髭蟾型的蝌蚪营流水攀吸型生活方式;当水速递减时,掌突蟾型蝌蚪营流水附着型生活方式;当水速进一步递减时,具有较小口漏斗的短腿蟾型蝌蚪和具有大漏斗的角蟾型蝌蚪营流水浮泳型生活。角蟾科蝌蚪对于水流递减的适应演化说明蝌蚪的生态学适应是具有进化意义的;(3)蝌蚪口器内部结构的分化揭示了蝌蚪和食性的适应关系,蝌蚪以口部的唇齿与角质颌刮取或吞吸水中的物质,然后,通过口乳突有选择地过滤进入口腔中食物。拟髭蟾亚科蝌蚪的唇齿多而窄,唇齿间距宽,颌鞘粗而稀,反映了其植食性为主的特点;它们的舌前乳突一般为指状,在口腔入口处所占面积小,其机械过滤的作用很多被唇齿和角质颌分担了;而角蟾亚科的蝌蚪,其角质颌弱,其舌前乳突一般为匙状,几乎填满了口腔入口处,因此舌前乳突起了主要的机械过滤作用。The family Megophryidae is the largest and most diverse families inArchaeobatrachia, and most of its species occur in India, Pakistan, and eastward intoChina, Southeast Asia, Borneo and the Philippines to the Sunda Islands. Currently thefamily includes 142 species have been grouped into two subfamilies, Megophryinaeand Leptobrachiinae. The mountains of central and southern China are rich in speciesof Megophryidae, 75 species belong to 9 genera and two subfamilies.The family was supposed to be ideal materials of studies in many fields of biology,such as taxonomy, evolution, systematics, ecology, and biogeography. Recently, therehave a great development in taxonomy and systematics of megophryids throughstudied by morphology, paleontology, cytology, ecology, and cladistics. However,larvae of megophryids were generally unknown, although the tadpoles might be veryimportant for above studies.In this paper, we examined the evolutionary scenario of the tadpoles’ morphologyin the context of a phylogenetic framework. Our objectives are (1) to evaluate thedivergence of larval body shape and oral discs in the family Megophryidae, (2) toexplore the evolutionary trends of the larvae in megophryidae, and test if thefunnel-shaped oral disc is apomorphic, and (3) to explore the relationship of the larvalstructure, diet and microhabitat.We examined larval morphology of 30 megophryid species, the larval body shape,oral discs, the buccopharyngeal cavity, and jaw sheaths and denticles of the Chinesemegophryid frogs were re-examined. We constructed a phylogeny of the species on thebasis of published mitochondrial cytochrome b and 16S rRNA gene segments usingpartitioned Bayesian analyses. Furthermore, hypothetical changes of larval morphologywere inferred using parsimony principle on the phylogeny. The results showed that:1) Four tadpole types in Megophryidae. The larval morphological charactersseries in Chinese megophryids fall into four general categories according to the bodyshape and oral discs: (A) Leptobrachiini type, species from genera Leptobrachium,Oreolalax, Scutiger and, Vibrissaphora share this type of tadpoles. (B) Leptolalax type,species of genus Leptolalax have this type of tadpoles. (C) Brachytarsophrys type,species of the genus Brachytarsophrys have this type of tadpoles. (D) Megophryinitype, species of the genera Atympanophrys, Ophryophryne, and Xenophrys share this type of tadpoles. Of which B and C are two novel types.2)Taxonomic implications. The present study leads us to reconsider the generalclassification of tribes attributed to members of Megophryidae. More specifically,concerning the phylogenetic relationships and the two novel tadpole types describedherein, we propose a provisional taxonomy for the family but suggest that further taxasampling of other megophryids be performed to confirm this taxonomic change. TheMegophryidae is composed of two subfamilies (Leptobrachiinae and Megophryinae).The Leptobrachiinae was recogonized the two tribes: (1) tribe Leptobrachiini sensuDubois, corresponding to the tadpole of type A, including four genera, i.e.,Leptobrachium, Oreolalax, Scutiger and, Vibrissaphora; (2) tribe Leptolalaxini,corresponding to the tadpole of novel type B, including two genera, i.e., Leptolalaxand Leptobrachella. However, the relationships among the genera of Megophryinaewere largely unresolved, they recognized no monophyletic groups above the generalevel. A more thorough sampling will likely foster a better taxonomic solution.3) The larval evolutionary scenario in Megophryidae.Type A is characteristicof normal-mouthed with multiple tooth rows, representing the tadpole type of theMRCA of Chinese megophryids. Type B is characteristic of normal-mouthed withreduced tooth rows, prolonging labium, and integumetary glands. Type C ischaracteristic of no labial teeth and smaller umbeliform oral disc. Type D ischaracteristic of no labial teeth, enlarged umbeliform oral disc, representing the tadpoleof the MRCA of subfamily Megophryinae. A previous hypothesis, referring tofunnel-shaped oral discs as an apomorphy, is supported.4) The larval adaptation to habitats in Megophryidae. Tadpoles generallyadhere to substrates using their mouths, and the microhabitat that the tadpoles occupyreflects the degree of adhesion and oral complexity. The morphological changes inmegophryid tadpoles virtually allow a progressive adaptation to a changing habitatfrom faster water to slower water. Within the tadpoles of Type A to type D, the TOTbecomes smaller and smaller, and the oral disc orientates from anteroventral toumbelliform upturned, and eye position orientates from dorsal to lateral, and the trunkis more and more depressed and tail becomes relatively longer and slender. Within therunning water, the normal-mouthed with multiple tooth rows of Leptobrachiini tadpoles are correlated with lotic-suctorial, benthic feeders with anteroventral oraldisc and the largest body. With the water’s velocity decreasing, the lotic-adherentfeeders of Leptolalax tadpoles have tube-shaped labium with reduced tooth rows andintegumetary glands. And then, the smaller umbeliform in Brachytarsophrys tadpolesand the enlarged umbeliform oral disc in the Megophryini tadpoles are inhabitmicrohabitats of non-flowing backwaters of rivers, indicative of adaptive traits oflotic-neustonic surface feeders. The scheme of megophryid tadpoles andmicrohabitats provided the first clear evidence which congruent with the hypothesis ofAltig and Johnston (1989). The ecological divergence plays a general role in thedivergence and evolution of megophrid larvae. There is a definite correlation amongthe buccopharyngeal cavity, diet and feeding mechanisms, the tadpole graze orswallow the food particles, then through papillae which like a sieve and sort out foodparticles to the oesophagus. The tadpole of Leptobrachiinae possess multiple toothrows, wide intertooth distance as well as thick and sparse jaw sheath, these tadpolesinhabit bottom of the streams and graze on epiphyton or major detritus of organicmatter on the substrates, their prelingual papillae like single finger, the mechanicalpurpose of papillae served share in by tooth and jaw. The tadpoles of Megophryinaeoccur near the water surface of small streams and are the filter feeder, their dietincludes plankton and organic debris floating on the water surface, those tadpolepossess weak jaw, their prelingual papillae like spoon, the mechanical purpose ofpapillae served mostly for sieve.