992 resultados para Central Tibetan Plateau


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The biostratigraphic distribution and abundance of Eocene to Pleistocene silicoflagellates is documented from Ocean Drilling Program Leg 120 Holes 747A, 748A, 748B, 749B, and 751A on the Central Kerguelen Plateau. Well-preserved silicoflagellates are reported here from the middle Eocene Dictyocha grandis Zone to the Pleistocene Distephanus speculum speculum Zone. Assemblage diversity and abundance is variable, with many intervals either barren of silicoflagellates or containing only limited numbers.

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Improvement of end-use quality in bread wheat depends on a thorough understanding of current wheat quality and the influences of genotype (G), environment (E), and genotype by environment interaction (G x E) on quality traits. Thirty-nine spring-sown spring wheat (SSSW) cultivars and advanced lines from China were grown in four agro-ecological zones comprising seven locations during the 1998 and 1999 cropping seasons. Data on 12 major bread-making quality traits were used to investigate the effect of G, E, and G x E on these traits. Wide range variability for protein quantity and quality, starch quality parameters and milling quality in Chinese SSSW was observed. Genotype and environment were found to significantly influence all quality parameters as major effects. Kernel hardness, flour yield, Zeleny sedimentation value and mixograph properties were mainly influenced by the genetic variance components, while thousand kernel weight, test weight, and falling number were mostly influenced by the environmental variance components. Genotype, environment, and their interaction had important effects on test weight, mixing development time and RVA parameters. Cultivars originating from Zone VI (northeast) generally expressed high kernel hardness, good starch quality, but poor milling and medium to weak mixograph performance; those from Zone VII (north) medium to good gluten and starch quality, but low milling quality; those from Zone VIII (central northwest) medium milling and starch quality, and medium to strong mixograph performance; those from Zone IX (western/southwestern Qinghai-Tibetan Plateau) medium milling quality, but poor gluten strength and starch parameters; and those from Zone X (northwest) high milling quality, strong mixograph properties, but low protein content. Samples from Harbin are characterized by good gluten and starch quality, but medium to poor milling quality; those from Hongxinglong by strong mixograph properties, medium to high milling quality, but medium to poor starch quality and medium to low protein content; those from Hohhot by good gluten but poor milling quality; those from Linhe by weak gluten quality, medium to poor milling quality; those from Lanzhou by poor bread-making and starch quality; those from Yongning by acceptable bread-making and starch quality and good milling quality; and those from Urumqi by good milling quality, medium gluten quality and good starch pasting parameters. Our findings suggest that Chinese SSSW quality could be greatly enhanced through genetic improvement for targeted well-characterized production environments.

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Mid-Cretaceous sediments recovered during Ocean Drilling Program Leg 183 (Cores 183-1138A-69R to 73R) on the central Kerguelen Plateau have been analyzed palynologically and paleobotanically to determine the age of the strata and to reconstruct vegetational development and paleoecology. The lower strata (Cores 183-1138A-71R to 73R), a dark, organic-rich silty claystone with many wood fragments and fern remains (sedimentary Unit VI), certainly of terrestrial origin, directly overlies the volcanic basement, which is dated as latest Albian (~95 to 103 Ma) to earliest Cenomanian. The age of the terrestrial strata can be determined by sporomorphs as late Albian to earliest Cenomanian as well. This shows that parts of the central Kerguelen Plateau must have been subaerial at least until the late Albian and were covered with a diverse high-latitude flora, probably dense conifer forest with various fern taxa in the undergrowth. Early angiosperms are also present. The vegetational character represented in Unit VI did not change significantly through time. However, varying percentages of several sporomorph groups seem to show recurring abundance variations, which might possibly be cyclic, caused by Milankovitch-type cyclic events. Cores 183-1138A-67R through 69R, of open marine origin, contain medium- to high-diversity dinocyst assemblages. Based on previous stratigraphic zonation schemes, the ages of these strata range within the Heterosphaeridium Superzone, from the Palaeohystrichophora infusorioides Zone to the Conosphaeridium striatoconus Zone, which correlates to the latest Cenomanian to Coniacian.

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La Revolución Espiritual promovida por el Dalai Lama plantea una unión entre espiritualidad y política. El proyecto de una ética universal, que se inscribe dentro de dicha Revolución, busca impactar la manera en que las relaciones internacionales se desarrollan, dándole prevalencia a los valores humanos. Sin embargo, esa proposición se encuentra ligada al contexto de exilio en el marco del conflicto sino-tibetano que afecta al continente asiático. Por esto, en la presente monografía, haciendo uso de los conceptos de marco de acción colectiva e identidad inscritos en la corriente de los movimientos sociales en la disciplina de las Relaciones Internacionales, se pretende determinar la relación entre identidad tibetana, marco de acción colectiva y la propuesta de una ética universal. Para ello se recurre, metodológicamente, a textos y a trabajo de campo en Bogotá. Así, se pretende establecer la relación entre espiritualidad y política como propuesta tibetana atravesada por el conflicto sino-tibetano.

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Sichuanissa Tiibetin ylängön metsäkato on pysähtynyt mutta eroosio-ongelmat jatkuvat Viikin tropiikki-instituutin tutkija Ping ZHOU kartoitti trooppisen metsänhoidon alaan kuuluvassa väitöskirjatyössään maaperän eroosioalttiutta ja sen riippuvuutta metsäkasvillisuudesta Jangtsen tärkeää sivuhaaraa Min-jokea ympäröivällä n. 7400 neliökilometrin suuruisella valuma-alueella Sichuanin Aba-piirikunnassa. Aineistonaan hän käytti muun muassa satelliittikartoitustietoja ja mittaustuloksia yli 600 maastokoealalta. Tutkimuksen nimi suomeksi on "Maaperän eroosion mallinnus ja vuoristoisen valuma-alueen ekologinen ennallistaminen Sichuanissa Kiinassa". Aikaisempien tutkimusten perusteella oli tiedossa että metsien häviäminen tällä alueella pysähtyi jo 1980-luvun alussa. Sen jälkeen on metsien pinta-ala hitaasti kasvanut etupäässä sen vuoksi, että teollinen puunhakkuu luonnonmetsissä kiellettiin kokonaan v. 1998 ja 25 astetta jyrkemmillä rinteillä myös maatalouden harjoittaminen on saatu lopetetuksi viljelijöille tarjottujen taloudellisten houkuttimien avulla. Täten myös pelto- ja laidunmaata on voitu ennallistaa metsäksi. Ping Zhou pystyi jakamaan 5700 metrin korkeuteen saakka kohoavan vuoristoalueen eroosioalttiudeltaan erilaisiin vyöhykkeisiin rinteen kaltevuuden, sademäärän, kasvipeitteen ja maalajin perusteella. Noin 15 prosentilla tutkitun valuma-alueen pinta-alasta, lähinnä Min-joen pääuomaa ympäröivillä jyrkillä rinteillä, eroosioriski oli suuri tai erittäin suuri. Eri tyyppisellä kasvillisuudella oli hyvin erilainen vaikutus eroosioalttiuteen, ja myös alueen sijainti vuoriston eri korkeuksilla vaikutti eroosioon. Säästyneet lähes luonnontilaiset havumetsät, joita on etupäässä vuoriston ylimmissä osissa 2600-4000 metrin korkeudella, edistävät tehokkaasti metsän luontaista uudistumista ja levittäytymistä vaurioituneille alueille. Säilyneiden metsien puulajikoostumus antoi tutkimuksessa mahdollisuuden ennustaa metsien tulevaa kehitystä koko tutkitulla valuma-alueella sen eri korkeusvyöhykkeissä ja eri maaperätyypeillä. Ennallistamisen kannalta ongelmallisimpia olivat alueet joilta metsäpeite oli lähinnä puiden teollisen hakkuun vuoksi kokonaan hävinnyt ja joilla maaperä yleisesti oli eroosion pahoin kuluttama. Näillä alueilla ei ole tehty juuri mitään uudistamis- tai ennallistamistoimenpiteitä. Niillä metsien ennallistaminen vaatii myös puiden tai pensaiden istuttamista. Tähän sopivia ovat erityisesti ilmakehän typpeä sitovat lajit, joista alueella kasvaa luontaisena mm. sama tyrnilaji joka esiintyy myös Suomessa. Työssä tutkittiin yli kahdeksankymmenen paikallisen luontaisen puulajin (joista peräti noin kolmannes on havupuulajeja) ekologisia ominaisuuksia ja soveltuvuutta metsien ennallistamiseen. Avainasemassa työn onnistumisen kannalta ovat nyt paikalliset asukkaat, joiden maankäytön muutokset ovat jo selvästi edistänet luonnonmetsän ennalleen palautumista. Suomen Akatemia rahoitti vuosina 2004-2006 VITRI:n tutkimushanketta, josta Ping Zhou'n väitöskirjatyö muodosti keskeisen osan. Kenttätyö Sichuanissa avasi mahdollisuuden hedelmälliseen monitieteiseen yhteistyöhön ja tutkijavaihtoon Kiinan tiedeakatemian alaisen Chengdun biologiainstituutin (CIB) kanssa; tämä tieteellinen kanssakäyminen jatkuu edelleen.

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The development and changes in the distribution of herbivorous mammal communities during the Neogene is complex. The Eurasian scale environmental patterns reflect the large scale geographical and climatic patterns. The reorganization of these affect the biome distribution throughout the continent. The distribution of mammal taxa was closely associated with the distribution of biomes. In Eurasia the Neogene development of environments was twofold. The early and middle Miocene that seemed to have been advantageous for mammals was followed by drying of environments during the late Neogene. The mid-latitude drying was the main trend, and it is the combined result of the retreat of Paratethys, the uplift of Tibetan Plateau and changes in the ocean currents and temperatures. The common mammals were "driving" the evolution of mammalian communities. During the late Miocene we see the drying affecting more and more regions, and we see changes in the composition of mammalian communities.

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Mountain waves in the stratosphere have been observed over elevated topographies using both nadir-looking and limb-viewing satellites. However, the characteristics of mountain waves generated over the Himalayan Mountain range and the adjacent Tibetan Plateau are relatively less explored. The present study reports on three-dimensional (3-D) properties of a mountain wave event that occurred over the western Himalayan region on 9 December 2008. Observations made by the Atmospheric Infrared Sounder on board the Aqua and Microwave Limb Sounder on board the Aura satellites are used to delineate the wave properties. The observed wave properties such as horizontal (lambda(x), lambda(y)) and vertical (lambda(z)) wavelengths are 276 km (zonal), 289 km (meridional), and 25 km, respectively. A good agreement is found between the observed and modeled/analyzed vertical wavelength for a stationary gravity wave determined using the Modern Era Retrospective Analysis for Research and Applications (MERRA) reanalysis winds. The analysis of both the National Centers for Environmental Prediction/National Center for Atmospheric Research reanalysis and MERRA winds shows that the waves are primarily forced by strong flow across the topography. Using the 3-D properties of waves and the corrected temperature amplitudes, we estimated wave momentum fluxes of the order of similar to 0.05 Pa, which is in agreement with large-amplitude mountain wave events reported elsewhere. In this regard, the present study is considered to be very much informative to the gravity wave drag schemes employed in current general circulation models for this region.

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Mountain waves in the stratosphere have been observed over elevated topographies using both nadir-looking and limb-viewing satellites. However, the characteristics of mountain waves generated over the Himalayan Mountain range and the adjacent Tibetan Plateau are relatively less explored. The present study reports on three-dimensional (3-D) properties of a mountain wave event that occurred over the western Himalayan region on 9 December 2008. Observations made by the Atmospheric Infrared Sounder on board the Aqua and Microwave Limb Sounder on board the Aura satellites are used to delineate the wave properties. The observed wave properties such as horizontal (lambda(x), lambda(y)) and vertical (lambda(z)) wavelengths are 276 km (zonal), 289 km (meridional), and 25 km, respectively. A good agreement is found between the observed and modeled/analyzed vertical wavelength for a stationary gravity wave determined using the Modern Era Retrospective Analysis for Research and Applications (MERRA) reanalysis winds. The analysis of both the National Centers for Environmental Prediction/National Center for Atmospheric Research reanalysis and MERRA winds shows that the waves are primarily forced by strong flow across the topography. Using the 3-D properties of waves and the corrected temperature amplitudes, we estimated wave momentum fluxes of the order of similar to 0.05 Pa, which is in agreement with large-amplitude mountain wave events reported elsewhere. In this regard, the present study is considered to be very much informative to the gravity wave drag schemes employed in current general circulation models for this region.

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The tropical easterly jet (TEJ) is a prominent atmospheric circulation feature observed during the Asian summer monsoon. It is generally assumed that sensible heating over the Tibetan Plateau directly influences the location of the TEJ. However, other studies have suggested the importance of latent heating in determining the jet location. In this paper, the relative importance of latent heating on the maintenance of the TEJ is explored through simulations with a general circulation model. The simulation of the TEJ by the Community Atmosphere Model, version 3.1 is discussed in detail. These simulations showed that the location of the TEJ is well correlated with the location of the precipitation. Significant zonal shifts in the location of the precipitation resulted in similar shifts in the zonal location of the TEJ. These zonal shifts had minimal effect on the large-scale structure of the jet. Further, provided that precipitation patterns were relatively unchanged, orography did not directly impact the location of the TEJ. These changes were robust even with changes in the cumulus parameterization. This suggests the potential important role of latent heating in determining the location and structure of the TEJ. These results were used to explain the significant differences in the zonal location of the TEJ in the years 1988 and 2002. To understand the contribution of the latitudinal location of latent heating on the strength of the TEJ, aqua-planet simulations were carried out. It has been shown that for similar amounts of net latent heating, the jet is stronger when heating is in the higher tropical latitudes. This may partly explain the reason for the jet to be very strong during the JJA monsoon season.

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This thesis aims at enhancing our fundamental understanding of the East Asian summer monsoon (EASM), and mechanisms implicated in its climatology in present-day and warmer climates. We focus on the most prominent feature of the EASM, i.e., the so-called Meiyu-Baiu (MB), which is characterized by a well-defined, southwest to northeast elongated quasi-stationary rainfall band, spanning from eastern China to Japan and into the northwestern Pacific Ocean in June and July.

We begin with an observational study of the energetics of the MB front in present-day climate. Analyses of the moist static energy (MSE) budget of the MB front indicate that horizontal advection of moist enthalpy, primarily of dry enthalpy, sustains the front in a region of otherwise negative net energy input into the atmospheric column. A decomposition of the horizontal dry enthalpy advection into mean, transient, and stationary eddy fluxes identifies the longitudinal thermal gradient due to zonal asymmetries and the meridional stationary eddy velocity as the most influential factors determining the pattern of horizontal moist enthalpy advection. Numerical simulations in which the Tibetan Plateau (TP) is either retained or removed show that the TP influences the stationary enthalpy flux, and hence the MB front, primarily by changing the meridional stationary eddy velocity, with reinforced southerly wind on the northwestern flank of the north Pacific subtropical high (NPSH) over the MB region and northerly wind to its north. Changes in the longitudinal thermal gradient are mainly confined to the near downstream of the TP, with the resulting changes in zonal warm air advection having a lesser impact on the rainfall in the extended MB region.

Similar mechanisms are shown to be implicated in present climate simulations in the Couple Model Intercomparison Project - Phase 5 (CMIP5) models. We find that the spatial distribution of the EASM precipitation simulated by different models is highly correlated with the meridional stationary eddy velocity. The correlation becomes more robust when energy fluxes into the atmospheric column are considered, consistent with the observational analyses. The spread in the area-averaged rainfall amount can be partially explained by the spread in the simulated globally-averaged precipitation, with the rest primarily due to the lower-level meridional wind convergence. Clear relationships between precipitation and zonal and meridional eddy velocities are observed.

Finally, the response of the EASM to greenhouse gas forcing is investigated at different time scales in CMIP5 model simulations. The reduction of radiative cooling and the increase in continental surface temperature occur much more rapidly than changes in sea surface temperatures (SSTs). Without changes in SSTs, the rainfall in the monsoon region decreases (increases) over ocean (land) in most models. On longer time scales, as SSTs increase, rainfall changes are opposite. The total response to atmospheric CO^2 forcing and subsequent SST warming is a large (modest) increase in rainfall over ocean (land) in the EASM region. Dynamic changes, in spite of significant contributions from the thermodynamic component, play an important role in setting up the spatial pattern of precipitation changes. Rainfall anomalies over East China are a direct consequence of local land-sea contrast, while changes in the larger-scale oceanic rainfall band are closely associated with the displacement of the larger-scale NPSH. Numerical simulations show that topography and SST patterns play an important role in rainfall changes in the EASM region.

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EXTRACT (SEE PDF FOR FULL ABSTRACT): The 1000 year records of particulate deposition (soluble and insoluble), oxygen isotopic ratios, and net accumulation from the Quelccaya ice cap are presented. The net accumulation record from Quelccaya is shown to serve as a reasonable proxy for the water levels in Lake Titicaca. ... The ice core record from the Dunde ice cap offers the potential to reconstruct a very detailed history of environmental conditions on the Tibetan Plateau for the last 3000 years.

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利用松科植物特殊的遗传体系(叶绿体基因组一父系遗传、线粒体基因组—母系遗传、核基因组一双亲遗传),我们对高山松及其两个亲本种进行了广泛的群体取样,通过线粒体基因nadl、叶绿体基因rbcL和trnL-F基因间区以及低拷贝核基因4CL的序列分析或PCR-RFLP分析,为高山松同倍体杂种起源假说提供了翔实的遗传学证据,同时在个体水平上探讨了高山松不同群体的遗传组成、群体遗传结构、基因交流方向、群体建立过程以及杂种基因组的进化。具体结果如下: 1.细胞质基因组分析 1)线粒体基因nudl分析 本研究对油松、高山松和云南松的19个群体、295个个体的线粒体基因nadl的一个内含子进行了序列分析或PCR-RFLP分析,共检测到3种线粒体DNA单倍型-A、B和C。油松所有的取样群体仅含单倍型A;除BX群体外,所有的云南松群体仅含单倍型B; 10个高山松群体中,5个群体固定单倍型A,4个群体固定单倍型B,1个群体(ZD)分布有A和B两种单倍型。2)叶绿体rbcL基因分析 对同一组群体的rbcL基因进行序列分析或PCR-RFLP分析,共检测到两个变异位点和三种叶绿体单倍型(TT、TC和GC)。TT和GC分别是油松和云南松种特异性叶绿体单倍型,而在高山松群体里则三种单倍型均有分布,而且TC单倍型广泛地分布在7个杂种群体中,该单倍型很可能来源于点突变或第三个已灭绝的亲本。rbcL基因检测到的高山松群体分化系数很高(Gst=0.533)。 3)叶绿体trn L-F区序列分析 叶绿体trnL-F分子标记检测到的不同单倍型的差异主要是由引物“e”下游120碱基处一个多聚T结构的长度变异所致(叶绿体SSR位点)。10个高山松群体中共检测到5种叶绿体单倍型,其中两种主要的单倍型(9T和11T)分别为油松和云南松的种特异性单倍型,其他单倍型均为非典型单倍型。群体遗传结构分析表明:杂种群体表现最高的遗传多样性,而且trnL-F分析得到的高山松群体的分化系数也很高( Gst=0.443)。 总之,对高山松、油松和云南松的同一组群体取样进行的细胞质基因组分析表明:高山松群体分布有油松和云南松种特异性的线粒体和叶绿体单倍型,该细胞质DNA单倍型的地理分布为假说“高山松为油松和云南松的的二倍体杂种”提供了翔实的遗传学证据。油松和云南松在不同的杂种群体中分别做父本和母本,即两亲本在杂交过程中发生了双向基因交流。群体遗传结构分析发现高山松群体表现最高的遗传多样性,而且群体间的分化系数很高。不同的杂种群体在遗传组成上的差异表明他们经历过不同的建立和进化历史。从线粒体和叶绿体单倍型的地理分布可以看出杂种群体的建立曾经历强烈的奠基者效应和回交。青藏高原的隆升对高山松的起源、杂种群体的适应辐射以及保持产生了重要的影响。川西南和滇西北作为青藏高原的东边边界,很可能是当初云南松和油松分布的重叠区及杂交地带,即高山松的起源地。 2.核基因4CL分析 对高山松、油松和云南松的19个群体、32个个体的低拷贝核基因4CL进行了克隆及序列分析,获得的78条序列可分为两种类型(类型A和类型B)。这两种类型明显的差别是类型A相对于类型B在内含子区有- 20bp的缺失。以华山松的3条序列为外类群,对得到的78条序列进行基因谱系分析,发现所有的序列分成明显的两支,分别对应于类型A和类型B,而且每一支均包含三个种的部分序列,表明4CL基因在这三个种分化之前就已发生重复。另一个明显的特点是某个种的一条序列与另一个种的序列比其与同种的其他序列关系更近,可能因基因交流(杂交和渐渗)、非共祖、致同进化和重组等进化事件所致。三种松树中共检测到4CL基因序列的两种类型和六个亚类型,高山松群体中没有发现杂种独特的类型或亚类型。高山松和云南松共享三种序列亚类型以及最多的序列多态性,表明这两个种之间曾存在广泛的基因交流。

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青藏高原是我国植物多样性和特有性最高的地区,其中横断山区由于复杂的地质变迁历史和独特的地貌已成为研究物种多样性的一个热点。全缘叶绿绒蒿是分布于青藏高原及其周边地区的高山植物,该物种的谱系生物地理学研究不仅有助于揭示横断山区的物种分化机制及地理分布格局的形成过程,而且有助于探讨青藏高原隆升及新生代晚期气候变化对横断山区物种遗传结构的影响。 本研究对全缘叶绿绒蒿的12 个群体、153 个个体的叶绿体DNA trnS-trnfM片段进行了序列测定,比对校正后该片段的长度为877-962 bp,其中含6 个突变位点和5 个插入缺失(两个7 bp,两个39 bp,一个6 bp)。根据这些变异,可分为12 种叶绿体单倍型(H1-H12)。全缘叶绿绒蒿在物种水平上的核苷酸多态性分别为π = 0.00152 和θ = 0.00122,单倍型多样性为HE 为0.791,群体之间的遗传分化系数FST 为0.46579。中性检测结果显示无论在物种水平还是在群体水平,都没有显著偏离中性模式,说明该物种近期没有经历明显的瓶颈效应或奠基者效应等历史事件。我们从地理上将全缘叶绿绒蒿的群体分为四个地区,即云南、川西、川北和青海地区。AMOVA 分析显示群体间的遗传分化系数FST为0.46579,大多数的核苷酸多态性来自群体内(53.42%),其次来自于地区内的群体间(40.44%),而四个地区之间的分化非常小,仅占6.14%。失配分析显示在整个物种的分布范围内呈现单峰式样(r = 0.0475, P = 0.07924),这种分布虽然是不显著的,但P 值接近显著的临界,说明可能存在不明显的扩散,但在各个地区内的群体都没有扩散的迹象。 比较单倍型的地理分布和单倍型之间的进化关系,发现川西和川北地区存在明显差异,这两个地区都拥有比较古老的单倍型,也具有地区特异的稀有单倍型,因此很可能是全缘叶绿绒蒿在冰期时的避难所。云南地区只具有一个古老单倍型H1,这种分布可能是由川西和川北避难所的南迁造成的,也可能该地区自身就是古老单倍型H1 的发源地。青海地区只有两种古老的单倍型,由于冰期后物种有往高纬度迁移的趋势,因此这个地区的单倍型可能是在冰期后由川西和川北迁移过去的。全缘叶绿绒蒿在横断山区单倍型的片段化分布及多个避难所的存在可能与该地区独特的高山低谷相间地貌以及受多次冰期影响有密切关系。

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种间杂交在自然界中普遍存在。杂交可以为群体带来多方面的遗传改变,具有重要的进化生物学意义。越来越多的证据表明,同倍性杂交物种形成是一种重要物种形成途径,在动、植物物种进化中扮演着重要的角色。同倍性杂交物种形成意味着一个稳定的、可育的、与亲本种间形成生殖隔离的杂种群体建立起来,但不改变染色体的数目。这种生殖隔离被认为是通过快速的染色体重组、生态分化、或空间分离形成的。对于现存杂交种的对比分析和对同倍性杂交物种形成的理论模拟都表明,适应性分化和生态转变在同倍性杂交物种形成中起着至关重要的作用。来自于生态成种的研究也表明生态选择自身就可以导致生殖隔离的形成和新物种的形成。尽管如此,我们对同倍性杂交物种形成过程的研究仍然较多地关注于染色体重组及相应的遗传效应;对于自然选择,特别是生态选择在同倍性杂交成种过程中的作用模式缺乏全面地了解。 本研究以同倍性杂交种-高山松为研究对象,利用地理信息学技术、基于天然群体取样的性状对比分析和人工杂交手段,全面地定量描述了高山松及其亲本种间的生态分化、生态转变下的天然群体生殖适应性、以及生态选择下的形态变异和种间杂交可配性。以此探讨了高山松的杂交成种过程,生态转变和自然选择的重要性以及它们与杂交种适应性、形态变异和生殖隔离之间的关系。利用地理信息学手段,对高山松及其亲本种间的生态分化对比分析表明:高山松与其亲本种间形成了明显的生态转变;在自然选择的作用下,高山松已经具备了对于青藏高原东南缘横断山区高山地带独特的生态适应性;相对于亲本种,它的适应性特征体现为混合型,在某些性状上它高出亲本种的适应性,在另一些性状上,它体现出居中、较低或者类似于亲本种适应性;高山松适生区由于地形地貌的复杂性,往往不连续,呈间断状。基于四个反映结实效率的性状、小孢子母细胞减数分裂异常现象及花粉萌发率在种间的对比分析表明,这些反映生殖状态的指标,在三个种及群体间并没有显著差异;相对于亲本种的天然群体,居于青藏高原的高山松群体,表现出了正常的授粉效率和生殖适应性。高山松天然群体正常的生殖效率表明,它经历了生态转变过程中的选择作用,已经具备了在新生境下,繁殖发展下去的生殖适应性。 在大规模群体取样的基础上,通过对反映植物个体发育和营养状态的球果和针叶形态性状的对比分析,我们发现,高山松及其亲本种间的形态变异存在以下特征:第一,绝大多数性状,在种间和种内群体间都存在着显著的差异,并且大部分性状的差异主要体现在种间;第二,高山松在大部分形态性状上介于两个亲本种之间,少数性状表现为超亲分离、类似于某个亲本或无种间差别;第三,在多数性状上,和亲本种相比,高山松群体都拥有较大的变异幅度。第四,在临近亲本种的群体中,高山松的形态特征更类似于所临近的亲本种。上述形态变异特征及其与气象、土壤、经纬度、海拔等因子间普遍的相关性,意味着生态选择在高山松成种过程中起了重要作用。总的来说,形态性状变异的总体模式揭示出:高山松在选择作用下的遗传和表型的精炼过程中已经形成了自身适应于高原环境的、稳定而有效的形态发育模式和资源利用方式,甚至是恰当的防御机制。 本研究通过云南松×油松的单交实验和油松×高山松部分双列杂交实验,检验了高山松及其亲本种间的杂交可配性。结果表明,就平均水平而言,高山松及其亲本种间的种间杂交结实可配性低于种内杂交,但个别种间杂交组合的结实可配性可能高于种间杂交;高山松及其亲本种间正在或已经形成了部分的合子后生殖障碍;现存的高山松群体及其亲本种间的生态地理隔离可能是很有效的生殖隔离形式,一方面做为合子前生殖隔离,它能避免种间相互传粉、阻碍种间杂交的发生,另一方面做为合子后生殖隔离,它能清除掉可能存在的劣势回交产物。 综合已有的证据,我们提出了高山松同倍性杂交物种形成模型:高山松的同倍性杂交成种是与青藏高原隆升过程以及隆升过程中所产生的全新生态空间紧密相关的;青藏高原隆升导致了生态环境的剧变,创造了全新的生态位,在选择的作用下,拥有综合了双亲遗传基础的杂种基因型开始占领这个全新的生态位;在分化的生态选择的持续作用下,杂种基因型在新的生境下稳定下来,形成了独立于亲本种的资源利用方式、形态特征和繁育系统,具备了完善的生殖和生态适应性;此时,一个与亲本种间以生态隔离存在的同倍性杂交种-高山松形成了;随着分化的生态选择的持续进行,高山松与亲本种间的内在生殖隔离不断加强。

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克隆植物,尤其是丛生禾草(如紫花针茅(Stipa purpurea Griseb)、丝颖针茅(S. capillacea Keng))和根茎苔草(如青藏苔草(Carex moorcroftii Falc. ex Boott)、窄叶苔草(C. montis-everestii Kük.)),在环境条件非常特殊的青藏高原高寒草原生态系统中,作为建群种或优势种而存在。本文利用控制实验研究不同构型克隆植物对资源供应的表型反应。 源于典型草地和灌木地的窄叶苔草和丝颖针茅对养分添加和光照处理产生广泛的表型反应。植物生长型影响分株数、比叶面积和叶鞘长度对光照强度的可塑性,源生境改变分株数、植株生物量和根茎总长对光照强度的可塑性。各性状对养分水平的可塑性反应均不受生长型和源生境的影响。游击型克隆植物窄叶苔草对高度异质性的灌木生境有较强的适应能力,而密集型克隆植物丝颖针茅的生长受生境的影响较小。 青藏苔草和紫花针茅对养分供应表现出了高度的表型可塑性。种群类型对少数性状有显著影响;除生物量分配和光合特征外,其它许多性状在基因型间有显著分化;除根茎生物量分配外,没有检测出表型可塑性的遗传分化;表型性状间有着广泛的表型整合及可塑性整合,并且整合格局随养分添加范围而变化,在种间也存在整合格局的差异。青藏高原高寒草原上强烈、稳定而持续的选择压力,以及一些与适合度相关的性状间的表型整合和可塑性整合可能对表型可塑性的进化有一定的限制作用。反应规范分析显示,高寒生境的克隆植物可能具有既能利用高养分环境,又能在低养分环境下维持生长的能力,这是植物适应养分环境变化的重要对策。 青藏苔草和紫花针茅的表型反应主要受养分供应量的影响,对由等量养分不同供应频次组成的实验处理的反应并不显著。青藏苔草的分株均重、叶片长度、比根长、根茎生物量分配和紫花针茅的比根长等性状在基因型间存在遗传分化;青藏苔草的分株数和总干重,以及紫花针茅的总干重和总根长表现出表型可塑性的遗传分化。比根长、根茎生物量分配等性状的遗传变异,以及这些性状的表型可塑性和可塑性的遗传变异对于不同基因型适应不同养分供应格局可能具有适应意义。 总之,青藏高原克隆植物对资源供应表现出了高度的表型可塑性,多数表型性状在不同基因型间存在显著的遗传分化,但表型可塑性的遗传分化相对有限。克隆植物可能通过遗传变异和表型可塑性适应异质性资源分布环境。表型可塑性及其遗传变异仍需要更多的研究,以便深入理解其适应性进化。