Evolution of 4-coumarate:coenzyme A ligase (4CL) gene and divergence of Larix (Pinaceae)

The evolutionary dynamics of the 4CL gene encoding 4-coumarate:coenzyme A ligase was investigated in the genus Larix (Pinaceae) by comparing copy number, GC content and codon usage, sequence divergence, and phylogenetic analysis. All 4CL clones of Larix formed a strongly supported monophyletic group, in which two robust clades (4clA and 4clB) derived from an ancient gene duplication event in the common ancestor of Larix were identified. Further gene duplication in the 4clA clade gave rise to two subclades 4clA(1) and 4clA(2). Frequent duplication/deletion appears to be a common evolutionary phenomenon in the 4CL gene family and paralogous genes differ greatly in their evolution rate. The existence of L. speciosa in subclades 4clA(1) and 4clA(2) suggests that this species may represent a primitive form of Larix or the closest relative of the common ancestor of the Eurasian Sect. Multiserialis. In addition, cpDNA and nrDNA ITS analyses support the hypothesis of an early separation of Larix into a North American and a Eurasian clade, which is congruent with the results of previous allozyme and very recent AFLP analyses. The unexpected close relationship between North American larches and the short-bracted species L. gmelinii in East Asia, based on the 4CL gene tree, may stem from lineage sorting.     

Phylogeography of Larix sukaczewii Dyl. and Larix sibirica L. inferred from nucleotide variation of nuclear genes

We investigated phylogeography of Larix sukaczewii and Larix sibirica using nucleotide variation at three following nuclear gene regions: 5.8 S rDNA including two internal transcribed spacers (ITS), cinnamyl alcohol dehydrogenase (CAD), and phytochrome-O (PHYO). We also included sequences of the 4-coumarate: coenzyme A ligase (4CL) gene region obtained in our recent study. CAD and PHYO showed very low nucleotide variation, but ITS and 4CL had levels of variation similar to those reported for other conifers. Pleistocene refugia have been hypothesized to exist in the Southern Urals and South Central Siberia, where four out of nine of the investigated populations occur. We found moderate to high levels of population differentiation (F _ST = 0.115 – 0.531) in some pairwise comparisons suggesting limited gene flow and independent evolution of some refugial populations. In L. sukaczewii, low levels of differentiation were found among populations from areas glaciated during the Pleistocene, indicating their recent origin. Our results also suggest these populations were created by migrants from multiple, genetically distinct refugia. Furthermore, some haplotypes observed in populations from previously glaciated areas were not found in putative refugial populations, suggesting these populations might have contributed little to the extant populations created after the Last Glacial Maximum. Some authors regard L. sukaczewii and L. sibirica as a single species, while others consider them as separate species. The observed conspicuous differences in haplotype composition and distribution between L. sukaczewii and L. sibirica, together with high values of F _ST between populations of the two species, appear to support the latter classification. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Ismael A. Khatab and Kariyawasam K.G.U. Hemamali contributed equally to this work.     

Nuclear and cytoplasmic variation within and between Eurasian Larix (Pinaceae) species

The genetic variation in nuclear and cytoplasmic markers was investigated in 28 populations belonging to Eurasian Larix species (L. decidua, L. sibirica, L. gmelinii, L. olgensis, and L. kaempferi). Nuclear genetic variation was assessed at 214 AFLP loci, and both PCR-RFLP and four microsatellite loci were used to estimate variation of the chloroplast DNA. Variation of the mitochondrial genome was measured using RFLPs. Although population differentiation at both nuclear and chloroplast markers was much weaker than at mitochondrial DNA, it nonetheless corroborated the grouping observed with mitochondrial DNA. The AFLPs led to the same population grouping as mtDNA. Notably, the presence of two ancient western and eastern groups within L. sibirica was confirmed and possible postglacial routes inferred. The genetic composition of the northernmost L. sibirica population in our sample established that it is located at the confluence of the eastern and western recolonization routes. The joint use of the three markers also indicated that populations around Lake Baikal are hybrids between L. gmelinii and L. sibirica, with L. gmelinii primarily acting as the pollinator. Finally, AFLP-based estimates of nucleotide variation were an order of magnitude larger than the strikingly low estimates of nucleotide variation recently reported in Pinus sylvestris.     

五种落叶松遗传关系的等位酶分析

由于种间形态上的微弱区别,落叶松属的系统分类一直很混乱,落叶松属的系统发生也知之甚少。本文分析了西伯利亚落叶松(Larix sibirica Ledeb.),卡氏落叶松(L.cajanderi Mayr.),兴安落叶松(L.gmelinii Rupr.),苏氏落叶松(L.sukaczewii Dil.)和杂交种切氏落叶松L.czekanowskii(L.gmelinii×L.sibirica)天然种群的遗传结构。采用水平切片淀粉凝胶电泳技术,等位酶分析手段对5个酶系统(AAT,IDH,DIA,PGM,SKDH)的8个基因位点进行了遗传结构分析。结果表明各种间遗传距离(D)在0.067~0.260之间,明显大于各种群内居群间的遗传距离。等位酶的分析结果揭示了5种落叶松的遗传关系。结合以上每种落叶松的形态学、生物学和生态学特性,等位酶的证据了支持兴安落叶松、西伯利亚落叶松、卡氏落叶松、苏氏落叶松作为独立种的观点。     

中国东北落叶松属植物rbcL 基因的序列分析及系统演化

长白落叶松的分类位置及与兴安落叶松的种间关系, 长期以来未有定论。本研究从大兴安岭、小兴安岭及张广才岭等6 个地点采集实验材料, 对其叶绿体DNA 的rbcL 基因进行PCR-RFLP 分析并测序。结果显示它们的rbcL 基因序列均没有差异, 表明东北地区落叶松属植物有共同起源。在系统演化上, 长白落叶松来自兴安落叶松(Larix gmelinii Rupr.), 应作为兴安落叶松的变种即L.gmelinii Rupr.var.olgensis Ostenfeld et S.Larsen     

中国东北落叶松属3种植物潜在分布对气候变化的敏感性分析

该文在东北地区多年平均的年均温、年降水分布图,海拔高程图、坡度图、坡向图和植被图的基础上,使用地理信息系统和Logistic回归模型的结合,预测3种落叶松(Larix sp.)的“气候-地形”潜在分布区。预测精度用敏感性、指定度和总正确率进行评价,3个树种的敏感性为61%~88%,指定度为80%~99.8%,总正确率为80%~99.8%。年均温、年降水和海拔是控制3种落叶松分布的主要环境因子。采用5种气温变化方案(+1 ℃、+2 ℃、+3 ℃、+4 ℃和+5 ℃)和6种降水变化方案(-30%、-20%、-10%、+10%、+20%和+30%),预测气候变化对各个树种潜在分布的影响,探索不同的树种对气候因子的敏感性。结果表明,气温每上升1 ℃,兴安落叶松(Larix gmelinii)将减少12%;长白落叶松(Larix olgensis var. changpaiensis)将增加23%;华北落叶松(Larix principis-rupprecntii)将增加500%。降水每增加10%,兴安落叶松将减少12.5%;长白落叶松将增加64%;华北落叶松将减少15%;随气候的“暖干化"(+5 ℃,-30%),兴安落叶松将向西北方退缩100 km左右;长白落叶松向西北方扩展100 km左右;华北落叶松将向东北方扩展800 km左右。随气候的“暖湿化"(+5 ℃,+30%),兴安落叶松将向西北退缩400 km左右;长白落叶松将向西北方扩展550 km;华北落叶松将向东北方扩展320 km左右。     

黑龙江漠河兴安落叶松与樟子松树轮生长特性及其对气候的响应

树木年轮 (简称树轮 ) 气候学是监测与重建全球气候变化的重要方法之一。针叶树树轮的生长能反馈出气温的变化, 在高纬度地带尤为明显。该文分析了生长在我国最北部的兴安落叶松 (Larixgmelinii) 与樟子松 (Pinussylvestrisvar.mongolica) 的树轮密度和宽度的特性。落叶松最大密度、晚材平均密度、早晚材宽度和轮宽都远高于樟子松。樟子松的所有密度变量的样本方差都明显高于兴安落叶松, 宽度变量的样本方差却明显低于兴安落叶松。两树种密度变量的差值年表显著相关, 宽度变量之间没有显著相关关系。落叶松与樟子松的晚材密度的形成受 7、8月的最高温控制。另外, 樟子松的晚材还与生长季节的长短相关。落叶松的年轮宽度对生长季节开始前的温度敏感, 而樟子松的轮宽对气候变量没有很好的响应。结果表明, 落叶松与樟子松的树轮最大密度都与生长季后期的温度显著相关, 两树种的树轮信息对气候变化的重建有很大的潜力。     

Response of larch species to climate changes

Global warming has changed the distributions of forests of northeastern China. Larix are very important species in this area. Predicting the potential distributions of Larix species and their responses to climate change would attract more and more attention.This paper predicted the potential distributions of three Larix species based on 'climatic-topographic' relationships by logistic regression. The results showed that L. gmelinii is predicted to retreat northwestward by 220 km by 2050 and by 270 km more by 2100; L. olgensis var. changpaiensis is predicted to retreat northwestward by 200 km by 2050 and by 190 to 300 km more by 2100; L. principis-rupprechtii is predicted to retreat northeastward by 200 km by 2050 and by 250 to 400 km more by 2100. This indicated that L. gmelinii could have its optimum latitude moved into Russia, L. olgensis var. changpaiensis could move to the Small Xing'an Mountains and L. principis-rupprechtii would move to the middle part of the Great Xing'an Mountains.     

大兴安岭兴安落叶松森林不同林型AMF分布特性

为了初步了解内蒙古大兴安岭兴安落叶松森林生态系统丛枝菌根真菌(AMF)多样性状况, 调查了5种落叶松林型和火烧迹地土壤中AMF状况。从90份土样中共分离到AMF 4属53种, 其中无梗囊霉属Acaulospora 25种(47.17%), 球囊霉属Glomus 23种(43.40%), 此二属为优势属, 内养囊霉属Entrophospora 4种(7.55%), 巨孢囊霉属Gigaspora 1种(1.89%)。杜香落叶松林的优势种为浅窝无梗囊霉A. lacunosa; 草地落叶松林没有优势种, 最常见种为浅窝无梗囊霉A. lacunosa; 柴桦落叶松林的优势种为一种无梗囊霉Acaulospora sp. 3和缩球囊霉G. constrictum; 落叶松皆伐林的优势种为刺无梗囊霉A. spinosa; 落叶松渐伐林的优势种为一种球囊霉Glomus sp. 3; 火烧迹地的优势种为刺无梗囊霉A. spinosa。5种林型中以柴桦落叶松林的孢子密度(41.00个/50 g 风干土)、物种丰富度(12.66种/土样)、多样性指数(H = 2.12, D = 0.85)都为最高。孢子密度与有机质含量呈明显正相关(r = 0.956*), 物种丰富度与速效磷含量呈明显的负相关(r = -0.899*)。     

中国东北落叶松属植物亲缘关系的研究

本文用RAPD方法对中国东北落叶松属植物的亲缘关系进行了研究。从100个引物中筛选出41个引物,检测出120个多态位点。应用UPGMA法计算了种群内和种群间的遗传距离,并构建了系统树。实验结果表明,分布在长白山的落叶松与分布在大、小兴安岭的落叶松之间的差异,在遗传距离上还不能达到种间分化水平。分布在长白山的落叶松应视为兴安落叶松种下的变种,分布在东京城的落叶松可能是兴安落叶松与长白山落叶松的杂交种。     

Analysis of Genetic Diversity and Population Differentiation of Larix potaninii var. chinensis Using Microsatellite DNA

Larix potaninii var. chinensis is endemic to China and is found only on several peaks of the Qinling Mountains in Shaanxi Province. In China, it is classified in the second class of national protected rare plants. To estimate genetic diversity and to analyze population genetic structure of L. potaninii var. chinensis, 120 individual samples from six natural populations were assessed using seven Larix SSR primer pairs. The results indicate that the level of genetic diversity of L. potaninii var. chinensis is very high, with a mean number of alleles per locus of 4.71. On the other hand, correspondingly low genetic differentiation was found between populations, with an F (ST) value of 0.116, suggesting that more than four-fifths of the genetic variation resides within populations. Besides the influence of habitat heterogeneity and historical distribution, the high level of genetic diversity of L. potaninii var. chinensis is also attributed to its biological characteristics. The definite genetic differentiation among populations, however, is attributed to the effects of genetic drift and natural selection, which are most likely due to the spatial isolation and inclement climate of the species' habitat. This study also revealed evidence that L. potaninii var. chinensis could be endangered, and some conservation strategies are suggested.     

Recolonization and radiation in Larix (Pinaceae): evidence from nuclear ribosomal DNA paralogues

Gene paralogy frequently causes the conflict between gene tree and species tree, but sometimes the coexistence of a few paralogous copies could provide more markers for tracing the phylogeographical process of some organisms. In the present study, nrDNA ITS paralogues were cloned from all but one species of Larix, an Eocene genus having two sections, Larix and Multiserialis, with a huge circumboreal distribution and an Eastern Asia-Western North America disjunction, respectively. A total of 96 distinct clones, excluding five putative pseudogenes or recombinants, were obtained and used in the gene genealogy analysis. The clones from all Eurasian species of section Larix are mixed together, suggesting that recolonization and recent morphological differentiation could have played important roles in the evolution of this section. In contrast, the species diversification of the Eurasian section Multiserialis may result from radiation in the east Himalayas and its vicinity, considering extensive nrDNA founder effects in this group. Our study also suggests that the distribution pattern analysis of members of multiple gene family would be very useful in tracking the evolutionary history of some taxa with recent origin or rapid radiation that cannot be resolved by other molecular markers.     

Taxa distribution and RAPD markers indicate different origin and regional differentiation of hybrids in the invasive Fallopia complex in central-western Europe

Interspecific hybridization can be a driving force for evolutionary processes during plant invasions, by increasing genetic variation and creating novel gene combinations, thereby promoting genetic differentiation among populations of invasive species in the introduced range. We examined regional genetic structure in the invasive Fallopia complex, consisting of F. japonica var. japonica , F.   sachalinensis and their hybrid F.  ×  bohemica , in seven regions in Germany and Switzerland using RAPD analysis and flow cytometry. All individuals identified as F. japonica var . japonica had the same RAPD phenotype, while F. sachalinensis (11 RAPD phenotypes for 11 sampled individuals) and F.  ×  bohemica (24 RAPD phenotypes for 32 sampled individuals) showed high genotypic diversity. Bayesian cluster analysis revealed three distinct genetic clusters. The majority of F . ×  bohemica individuals were assigned to a unique genetic cluster that differed from those of the parental species, while the other F . ×  bohemica individuals had different degrees of admixture to the three genetic clusters. At the regional scale, the occurrence of male-fertile F. sachalinensis coincided with the distribution of F . ×  bohemica plants showing a high percentage of assignment to both parental species, suggesting that they originated from hybridization between the parental species. In contrast, in regions where male-fertile F. sachalinensis were absent, F . ×  bohemica belonged to the non-admixed genetic group, indicating multiple introductions of hybrids or sexual reproduction among hybrids. We also found regional differentiation in the gene pool of F.  ×  bohemica , with individuals within the same region more similar to each other than to individuals from different regions.     

Polymorphism of lectin genes in Lathyrus plants

The carbohydrate-binding sequences of the lectin genes from spring vetchling Lathyrus vernus (L.) Bernh., marsh vetchling L. palustris (L.), and Gmelin's vetchling L. gmelinii (Fitsch) (Fabaceae) were determined. Computer-aided analysis revealed substantial differences between nucleotide and predicted amino acid sequences of the lectin gene regions examined in each of the three vetchling species tested. In the phylogenetic trees based on sequence similarity of carbohydrate-biding regions of legume lectins, the sequences examined formed a compact cluster with the lectin genes of the plants belonging to the tribe Fabeae. In each plant, L. vernus, L. palustris, and L. gmelinii, three different lectin-encoding genes were detected. Most of the substitutions were identified within the gene sequence responsible for coding the carbohydrate-binding protein regions. This finding may explain different affinity of these lectins to different carbohydrates, and as a consequence, can affect the plant host specificity upon development of symbiosis with rhizobium bacteria.     

Hierarchical components of genetic variation at a species boundary: population structure in two sympatric varieties of Lupinus microcarpus (Leguminosae)

Lupinus microcarpus is a self-compatible annual plant that forms a species complex of morphologically variable but indeterminate varieties. In order to examine the hypothesis that varieties of L. microcarpus comprise genetically differentiated and reproductively isolated species, populations of L. microcarpus var. horizontalis and var. densiflorus were sampled from an area of sympatry in central California and genotyped using six microsatellite loci. Bayesian clustering divided the total sample into two groups corresponding to the named varieties with extremely low levels of inferred coancestry. Similarly, maximum likelihood and distance methods for genetic assignment placed individuals in two nonoverlapping groups. Evidence for isolation by distance (IBD) within each variety was found at shorter distance classes, but varieties remained differentiated in sympatry. Furthermore, coalescent estimates of divergence time indicate separation within the past 950-5050 generations, with minimal gene flow after divergence. A four-level hierarchical analysis of molecular variance (amova) found significant levels of genetic differentiation among varieties (theta(P) = 0.292), populations within varieties (theta(S) = 0.449), subpopulations within populations (theta(SS) = 0.623), and individuals within subpopulations (f = 0.421); but the greatest degree of differentiation was at the subpopulation level. Although it is sometimes assumed that the magnitude of genetic differences (e.g. F(ST)) should be greater between species than among populations or subpopulations of the same species, shared ancestral polymorphism may lead to relatively low levels of differentiation at the species level, even as the stochastic effects of genetic drift generate higher levels of differentiation at lower hierarchical levels. These results suggest that L. microcarpus var. horizontalis and var. densiflorus are recently diverged yet reproductively isolated species, with high levels of inbreeding resulting from the combined effects of limited gene flow, demographic bottlenecks, and partial selfing in finite, geographically structured populations.     

取食落叶松的槌缘叶蜂属一新种(膜翅目,叶蜂科)

记述采自中国西北地区的突瓣叶蜂亚科1新种:西北槌缘叶蜂Pristiphora xibei sp.nov.,编制了古北区危害落叶松属植物的槌缘叶蜂属种类分种检索表。在宁夏地区,新种危害华北落叶松Larix gmelinii var.principisrupprechtii(Mayr.)Pilger,并与红环槌缘叶蜂(落叶松红腹叶蜂)Pristiphora erichsonii(Hartig,1837)混合发生,但种群数量显著小于后者。     

部分落叶松属植物核型研究

对落叶松属的兴安落叶松、长白落叶松、日本落叶松、华北落叶松、欧洲落叶松、美洲落叶松等6个种和日本落叶松×长白落叶松杂种进行了核型分析,所有材料的染色体数目均为2n=2x=24,2A核型。所有种的核型有着共同的构型,即二型核型。根据核型特征,可将以上各种分为三组:兴安落叶松、长白落叶松、日本落叶松为第一组,核型公式为2n=12m+10sm+2st;欧洲落叶松、美洲落叶松为第二组,核型公式为2n=12m+12sm;华北落叶松虽然具有与第二组相同的核型公式,但核型的对称性与其有一定差距,因此单独构成第三组。日本落叶松×长白落叶松杂种的核型数据和两个亲本的十分接近,表明这两个种杂交后染色体可能未发生明显的结构变异。作者认为长白落叶松和兴安落叶松的核型特征比较接近,从细胞学的角度作者支持长白落叶松作为兴安落叶松变种的观点。     

Refugia, differentiation and postglacial migration in arctic-alpine Eurasia, exemplified by the mountain avens (Dryas octopetala L.)

Many arctic-alpine organisms have vast present-day ranges across Eurasia, but their history of refugial isolation, differentiation and postglacial expansion is poorly understood. The mountain avens, Dryas octopetala sensu lato, is a long-lived, wind-dispersed, diploid shrub forming one of the most important components of Eurasian tundras and heaths in terms of biomass. We address differentiation and migration history of the species with emphasis on the western and northern Eurasian parts of its distribution area, also including some East Greenlandic and North American populations (partly referred to as the closely related D. integrifolia M. Vahl). We analysed 459 plants from 52 populations for 155 amplified fragment length polymorphisms (AFLP) markers. The Eurasian plants were separated into two main groups, probably reflecting isolation and expansion from two major glacial refugia, situated south and east of the North European ice sheets, respectively. Virtually all of northwestern Europe as well as East Greenland have been colonized by the Southern lineage, whereas northwest Russia, the Tatra Mountains and the arctic archipelago of Svalbard have been colonized by the Eastern lineage. The data indicate a contact zone between the two lineages in northern Scandinavia and possibly in the Tatra Mountains. The two single populations analysed from the Caucasus and Altai Mountains were most closely related to the Eastern lineage but were strongly divergent from the remaining eastern populations, suggesting survival in separate refugia at least during the last glaciation. The North American populations grouped with those from East Greenland, irrespective of their taxonomic affiliation, but this may be caused by independent hybridization with D. integrifolia and therefore not reflect the true relationship between populations from these areas.     

大兴安岭林区森林群落的演替

本文根据1954年设置的29块标准地的调查,分析了大兴安岭林区主要森林群落的演替趋势,指出兴安落叶松林是该区的演替顶极,樟子松林是土壤演替顶极,白桦林、黑桦林及蒙古栎林等次生林正向着兴安落叶松林演替。     

Phenetic analysis in paleoanthropology: phenogeography of peoples of the world

Phenetic diversity of peoples of the world in a system of nonmetric, discrete variable traits has been studied. Sixty-two populations from North, Central, and Southeast Asia, Eastern and Western Europe, America, East Africa, Australia and Melanesia have been examined. The estimates of phenetic diversity within regions (F(st)) and the distances of the regions from the global means (d) proved to be comparable to the corresponding estimates inferred from genetic data. This means that differentiation of populations in discrete variable traits is related to the history of formation of their gene pools. A classification tree of the world peoples constructed using bootstrap implemented in the PHYLIP program package (Felsenstein, 1993) showed that the Australo--Melanesian populations were close to the East African ones but separated from those of the Eurasian region. The results of phylogenetic analysis of the reconstructed phene pools of the regional ancestral populations support the assumptions on the early colonization of Australia and Melanesia and on the later time of divergence of the ancestors of modern Caucasoids and North Asian Mongoloids.