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资助项目
GST, P < 0.05) were exhibited by this species. The SAMOVA revealed seven diverging groups of related chlorotypes, six of them had distinct nonoverlapping geographical ranges: one in the northeast comprising 10 populations, a second with a southeast distribution comprising 22 populations, and the remaning four groups comprising 15 populations located in the west part of the species’ range along different river valleys. The genetic clustering of populations into three regions was also supported by analysis of molecular variance, which showed that most genetic variation (82.43%) was found among these three regions. Two clusters were distinguished by both phylogenetic analysis and genealogical analysis of chlorotypes, one consisting of chlorotypes from the western region and the second consisting of those from the eastern region. Significant genetic differences between the two regions might be attributed to vicariance and restricted gene flow, and this vicariance could be explained by the physical environmental heterogeneity on each side of the Tanaka-Kaiyong Line. Following the uplift of the Tibetan Plateau, the reorganization of the major river drainages was primarily caused by river separation and capture events. These historical events could change the distribution of S. davidii from fragmented to continuous (Upper/Lower Jinshajiang and Yalongjiang/Daduhe), and from continuous to fragmented (Nujiang and Jinshajiang/Honghe). However, spatial and temporal patterns of phylogeographic divergence are strongly associated with historical disjunction rather than modern drainage connections. Moreover, the following north-south split in the eastern region and effective isolation with their genetic diversity were essentially modelled by genetic drift. The higher chlorotype richness and genetic divergence for populations in western region compared with other two regions suggests that there were multipe refugia or in situ survival of S. davidii in the Himalayan-Hengduan Mountain region. Fixation of chlorotypes in the northeastern region and near fixation in the southeastern region suggest a recent colonization of these areas. We further found that this species underwent past range expansion around 37-303 thousand years ago (kya). The southeastern populations likely experienced a demographic expansion via unidirectional gene flow along rivers, while northeastern populations underwent a more northward expansion, both from initial populations (s) (21, 22, 23) preserved on eastern refugia (Jinshajiang). This process might have been accompanied with a series of founder effects or bottlenecks making populations genetically impoverished. 3. Phylogeographic analysisbased on nuclear sequence，We sequenced the nuclear (ncpGS) region in all populations sampled, recovering 23 nuclear haplotypes. Compared to cpDNA, both NST (0.470) and GST (0.338) were relatively lower, but NST was also significantly larger than GST. 37.10% of the total variation was distributed among regions which was much lower than that shown by chlorotypes. Thus, more extensive distribution of nuclear haplotypes was exhibited across the geographical range instead of the strong population subdivision observed in chlorotypes. Similarly to the chloroplast data, we found that genetic differentiation of nDNA was positively correlated with the geographical distance, but the increase in the geographical distance between populations did not increase the genetic differentiation of nDNA as rapidly as that of cpDNA. These contrasting levels between the chloroplast and nuclear genomes of S. davidii are likely due to limited gene flow of cpDNA by seeds vs. the extensive gene flow of nDNA by wind-mediated pollen in the population history. We also determined from nuclear markers that haplotype diversity was reduced in the southeastern and northeastern regions due to the loss of rare haplotypes in western region. This reduction of gene diversity is also a signature of founder events or recent bottleneck during post-glacial colonization. However, nuclear diversity within populations remains high. This provides evidence that regionally pollen flow might be sufficiently high to blur the genetic identity of founder populations over a reasonably large spatial scale.3. Relationships among three varieties，The phylogenetic analysis identified two phylogroups of chlorotypes, corresponding to S. davidii var. davidii and var. chuansinesis. The former was distinguished by the abscence of predonminant nuclear haplotype H1 of the latter. The monophyletic group of chlorotypes in var. davidii and var. liangshanesis showed their relatively close relationship. And their genetic divergence from the third variety appears to be relative to their slight morphological difference in leaf size and the divergent environmental niche spaces they occupy. Thus, the observed differences in morphological characters between var. chuansinesis and other two varieties can be explained by the seed dispersal limitation illustrated above (as inferred by geographical separation) and by environmental heterogeneity (as inferred by precipitation or elevation) or by a combination of both. After all, the geological changes, drainage reorganization, and floristic differences following the Himalayan uplift have been suggested to affect the genetic structure of S. davidii. These results provide new insights into the phylogeographic pattern of plants in China. In addition, the unique population genetic structure found in S. davidii has provided important insights into the evolutionary history of this species. The genetic profile uncovered in this study is also critical for its conservation management. Our study has uncovered the existence of at least two ‘evolutionary significant units’ independent units within S. davidii, corresponding to var. davidii from eastern region and var. chuansinensis from western region. The conservation efforts should first focus on most western populations and on the southeastern ones exhibiting high levels of genetic diversity, while the genetically homogeneous northeastern populations located in the degraded Loess Plateau should require much greater conservation 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reaction of transphosphatidylation is which phospholipase D catalyzes hydrolytic cleavage of the terminal phosphate diester bond of glycerophosphatides, transfer the phosphatidyl moiety of a phospholipids to a primary alcohol or water, producing phosphatidyl alcohol or phosphatidic acid (PtdOH). Although this reaction plays important role in regulating physical process in plants and anminals, the pathway involved in and detailed regulation mechanism are still unknown or not clear. There are three factors which can affect transphosphatidylation, substrates, enzymes and products. This paper intended to uncover the physical effect of the reaction in detail by changing two indispensable factors, the type of substrate and the PLDs. This paper consist of three researches, the first one, changing the accept substrate of the reaction aims to study the relationship between the structure of alcohols and the physical effect, to reveal the significance of the reaction; secondly, using genetic methods to regulate the expression of PLDs mediating transphosphatidylation intends to study its roles in regulating senescence; thirdly, reducing mostly PA derived from PLDs by addition 1-Butanol try to study the effect of PLD-derived PA on phosphate starvation. 1. Initial research of the effect of alcohols on the palnt growth. In this study, we study on the effect of alcohols on Arabidopsis seed germination, seeding growth and membrane lipids molecules, the metabolic procees and signaling pathway invoved in plant responding to 1-Butanol. According to the results we observed, the toxicity of alcohols and its polarity is positively correlated; the toxicity of alcohols to plants may be related with the butanoate and propanoate metabolic pathway rather than transphosphatidylation, the increase of free IAA is contributed to the resistance to 1-Butanol for WS ecotype Arabidopsis thaliana. Futhermore, five 1-Butanol insensitive mutants have obtained and the mutant sites has identified. Under the same concentration 1-Butanol stress, the five mutants are in better condition no matter physical and chemical level or cellular levels compared to wild-type Arabidopsis thaliana. It is believed that the five mutants definitely provide direct evidence for us to uncover the mechanism of the toxicity of alcohols to plants. 2. The change pattern of the membrane lipid molecules responding to the phytohormones-promoted detached Arabidopsis leaves and the role of PLDα1 and PLDδ in regulating this process. The reason for PLDα1-antisense mutant and PLDδ-knockout mutant delaying phytohormone-promoted senescence may be PLDα1 and PLDδ are response to the ABA and ethylene sinaling; PLDα1-or PLDδ-derived PA contributes to reactive oxygen species (ROS) accumulation; PLDα1-antisense mutant and PLDδ-knockout mutant have higher level of indole-3-acrtic acid (IAA) and zeatin riboside (ZR) level but lower level of abscisic acid (ABA) and jasmonic acid (JA) compared to wild-type Col and wild-type WS, respectively, which is favorable to retard the phytohormone-promoted detached leaves senescence. 3. Application tert-butanol increase the efficiency of phosphate utilization and absord when the phosphate-limited. The seedinges phosphate-starved or not exposed to 1-Butanol was subjected to lethal stress. However, in low phosphate condition, application of tert-butanol lower anthocyanin accumulation, increase the phosphate levels and the siliques number, in a word, tert-butanol attenuates the symptom of phosphate-starved induced. It is suggested that tert-Butanol can increase the utilization and absord efficiency of the limited phosphate, however, more evidences are needed to prove the 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