新疆阜康地区重要荒漠植物自然种群生态遗传学研究
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摘要
本论文试图以新疆阜康绿洲沙漠过渡带分布的三种重要植物红砂(Reaumuria
soongorica)、无叶假木贼(Anabasis aphylla)和角果藜(Ceratocarpus arenarius)作为研究对象，就其各自的生态遗传学特点做了初步的研究，进而探讨三种荒漠植物自然种群在形态学、遗传学及生理生态学等方面的变异和分化与环境的关系，为阐释物种进化、物种形成的生态遗传学机制和制定生物多样性保护策略提供理论上的依据和遗传学背景。
首先，应用RAPD标记技术对分布于新疆阜康荒漠的重要植物红砂种群遗传结构和遗传多样性进行了分析。15条随机引物对红砂7个亚种群的136个个体进行扩增，共检测71个位点，其中多态位点69个。研究结果表明：红砂种群的多态位点比率(PPB)为97.18％，显示出分布于过渡带的红砂种群内存在较高的遗传多样性。Shannon多样性指数(0.3075)、Nei基因多样性指数(0.3127)和基因分化系数(Gst=0.3120)揭示了红砂种群遗传变异多存在于亚种群内，而亚种群间的遗传分化则较小。红砂亚种群间的基因流Nm=1.1028，Nm＞1但低于一般风媒传粉植物(Nm=5.24)的基因流水平，处于分化的临界状态。AMOVA分析说明红砂种群变异的61.58％存在于亚种群内，而亚种群间的变异占总变异的38.02％。另外，通过RAPD资料的聚类分析、主成分分析以及相关性分析研究，发现红砂自然种群的遗传结构与绿洲沙漠过渡带的微生境生态因子(主要是土壤因子)相关，其中红砂亚种群遗传多样性水平与土壤中全磷和Cl~-含量呈显著负相关，与CO_3~(2-)含量呈显著正相关。而其它土壤生态因子则与红砂的遗传多样性指数的相关性均不显著(P＞0.05)。表明红砂个体的分布可能与过渡带土壤的某些易溶性盐分相关。研究还发现，随着土壤中有机质、水分、全氮和全磷含量的减少，红砂种群的遗传多样性水平有上升的趋势。这可能与植物为了适应恶劣环境而在长期进化中产生更大的遗传变异潜力有关。本试验结果也进一步证实分布于较小范围内的红砂亚种群间的遗传距离、遗传多样性与亚种群的地理距离(经纬度差异)大小无显著的相关性。
同时，本文采用RAPD标记技术对分布于新疆阜康绿洲沙漠过渡带的重要藜科盐生半灌木无叶假木贼种群和一年生泌盐性草本角果藜种群的遗传多样性和遗传分化进行了研究。试验分别选择了16条随机引物，对同一生境中的无叶假木贼3个亚种群(58个个体)及角果藜4个亚种群(80个个体)进行RAPD扩增。研究结果表明：无叶假木贼种群的多态位点比率(PPB)为94.56％，角果藜种群的多态位点比率则为98％，显示出分布于过渡带的无叶假木贼和角果藜种群内均存在较高的遗传多样性，而角果藜种群遗传多样性水平高于无叶假木贼种群。经分析表明，无叶假木贼种群与角果藜种群的Shannon多样性指数分别为0.1403和0.1615，基因分化系数(G_(ST))为0.1469和0.1659，揭示了两物种种群遗传变异多存在于亚种群内，亚种群间的遗传分化则不明显。AMOVA分析说明无叶假木贼种群变异的87.59％存在于亚种群内，而亚种群间的变异占总变异的12.41％；而角果藜亚种群内与亚种群间的变异则分别占总变异的87.43％和12.57％。另外，通过RAPD资料的聚类分析及相关性分析研究，发现无叶假木贼和角果藜自然种群的遗传结构与绿洲沙漠过渡带的微生境生态因子(主要是土壤因子)相关，其中无叶假木贼亚种群遗传多样性水平不仅与土壤含水量(W)、钾钠离子浓度(K+Na)和氯离子浓度(Cl)呈显著的正相关(P＜0.05)，还与土壤中有机质(SOM)、全氮(Nt)和全磷(Pt)含量呈显著的负相关；同时，角果藜种群的遗传多样性水平与土壤中有机质(SOM)和全氮(Nt)含量呈显著的正相关，而与CO_3~(2-)浓度呈显著的负相关；除此之外，其它土壤生态因子与两物种遗传多样性水平的相关性均不显著
(卜0.05)。试验结果还进一步证实，分布于较小范围内的无叶假木贼和角果葵亚种群间的
遗传距离、遗传多样性与亚种群的地理距离(经纬度差异)大小无显著的相关性。研究通过
对同一生境、不同生活型的无叶假木贼和角果要两种盐生植物种群的遗传多样性进行分析比
较，探讨了两种植物的遗传多样性水平与生态生物学特点和进化过程的关系，为进一步全面
研究两种植物种群的遗传结构和进化适应提供参考。
另外，本文以构件理论为基础，运用灰色关联度分析技术，对新疆阜康荒漠红砂自然种
群构件结构及其与环境因子的关系进行了研究.结果表明:l)植株冠幅是表征红砂生长情
况的重要指标，随着红砂植株冠幅的增大，植株高度明显增加，而且植株冠幅与植株的一级
分枝系统和当年生枝系统的构件因子相关度较大.2)在某种意义上，一级分枝系统是红砂
植株的基础，三级分枝系统和当年生枝系统则是二级分枝系统的派生物，它们之间的关联性
显示了红砂种群构件组成间的整合作用。3)一级分枝系统内的分枝高度与分枝数、长度及
分枝角度有较大的关联度，但与径粗关联度相对较小，说明一级分枝系统内后者的变化与前
四者并不完全相似，表现出一定的异质性。4)在二级分枝系统中，一方面，各构件特征与
一级分枝系统存在较大的关联度，反映了红砂种群构件结构一定程度的整合作用;另一方面，
二级分枝构件系?
The paper presented a comprehensive survey of ecological genetic characteristics of Reawnuria soongorica, Anabasis aphylla and Ceratocarpus arenarius distributed in oasis- desert transitional zone in Fukang, Xinjiang. The relationships between the morphological, genetic, physiological and ecological variations and differentiations and the environmental factors were discussed to explain the mechanism of the ecological genetics during the process of evolution and formation of species and provide the theoretical basis and genetic background for planning the scientific strategies of biodiversity conservation.
Firstly, we studied the genetic structure and genetic differentiation of Reaumwia soongorica distributed in Fukang, Xinjiang, by the means of random amplified polymorphic DNA (RAPD) markers. The results showed that the higher genetic diversity and the genetic differentiation existed in the population of Reaumuria soongorica, which was composed of 7 subpopulations and 136 individuals, generated by 15 primers. 71 loci had been detected and 69 were polymorphic. The mean proportion of polymorphic loci (PPB) of Reawnuria soongorica was 97.18%. Furthermore, the study discussed the Shannon information index (0.3075), Nei's gene diversity index (0.3127) and Gst (0.3120), which indicated that there were more genetic variations within the subpopulations than those among the subpopulations. The gene flow among the subpopulations of Reawnuria soongorica (Mn=1.1028, Nm>1) was much less than that of the common anemophytes (ATm=5.24), so the species was in the critical state of genetic differentiation to some extent The results of AMOVA showed that there were 61.58% of the total genetic variations existing among the subpopulations, and 38.02% within them. Additionally, through the use of clustering analysis, principal factor analysis and the correlation analysis, we found that the genetic structure of natural population Reawnuria soongorica was related to the ecological factors (soil factors mainly) of the oasis-desert transitional zone. The genetic diversity level ofReaumuria soongorica had the negative correlation with the content of soil total P and Cl" significantly (P<0.05), on the contrary, it had the positive correlation with CO32- significantly (PO.05) but there was no significant correlation between the genetic diversity and the other salt factors in soil, which showed that the distribution of the individuals of Reawnuria soongorica correlated with some certain soluble salt. Furthermore, the study indicated that the genetic diversity of the natural population of Reawnuria soongorica increased with the decreasing of the content of organic matter, water, total N and total P in soil, which had some relation with more potentiality of the genetic and variation of Reawnuria soongorica in its long-term evolution to adapt worse environment. Besides, the paper demonstrated that there was no significant correlation between the genetic distance, genetic diversity and the geographical difference on latitude and longitude respectively within the small range.
Secondly, the genetic diversity and genetic differentiation of the two important desert plants of Chenpodiaceae, Anabasis aphylla and Ceratocarpus arenarius, distributed in Fukang Desert, Xinjiang, were studied using RAPD markers. The results showed that the mean proportion of polymorphic loci (PPB) of Anabasis aphylla, which comprised 3 subpopulations (58 individuals sampled), generated by 16 primers was 94.56%, the value of PPB of Ceratocarpus arenarius, which had 4 subpopulations (80 individuals sampled), using 16 primers was 98.00%. The paper had proved that the higher genetic diversity and the genetic differentiation existed in the populations of boih Anabasis aphylla and Ceratocarpus arenarius, while the latter had much more
genetic diversity than the former. Moreover, the study discussed the Shannon information index and Nei's gene diversity index of the two species natural populations, which indicated that there were more genetic variations within the subpopu
soongorica)、无叶假木贼(Anabasis aphylla)和角果藜(Ceratocarpus arenarius)作为研究对象，就其各自的生态遗传学特点做了初步的研究，进而探讨三种荒漠植物自然种群在形态学、遗传学及生理生态学等方面的变异和分化与环境的关系，为阐释物种进化、物种形成的生态遗传学机制和制定生物多样性保护策略提供理论上的依据和遗传学背景。
首先，应用RAPD标记技术对分布于新疆阜康荒漠的重要植物红砂种群遗传结构和遗传多样性进行了分析。15条随机引物对红砂7个亚种群的136个个体进行扩增，共检测71个位点，其中多态位点69个。研究结果表明：红砂种群的多态位点比率(PPB)为97.18％，显示出分布于过渡带的红砂种群内存在较高的遗传多样性。Shannon多样性指数(0.3075)、Nei基因多样性指数(0.3127)和基因分化系数(Gst=0.3120)揭示了红砂种群遗传变异多存在于亚种群内，而亚种群间的遗传分化则较小。红砂亚种群间的基因流Nm=1.1028，Nm＞1但低于一般风媒传粉植物(Nm=5.24)的基因流水平，处于分化的临界状态。AMOVA分析说明红砂种群变异的61.58％存在于亚种群内，而亚种群间的变异占总变异的38.02％。另外，通过RAPD资料的聚类分析、主成分分析以及相关性分析研究，发现红砂自然种群的遗传结构与绿洲沙漠过渡带的微生境生态因子(主要是土壤因子)相关，其中红砂亚种群遗传多样性水平与土壤中全磷和Cl~-含量呈显著负相关，与CO_3~(2-)含量呈显著正相关。而其它土壤生态因子则与红砂的遗传多样性指数的相关性均不显著(P＞0.05)。表明红砂个体的分布可能与过渡带土壤的某些易溶性盐分相关。研究还发现，随着土壤中有机质、水分、全氮和全磷含量的减少，红砂种群的遗传多样性水平有上升的趋势。这可能与植物为了适应恶劣环境而在长期进化中产生更大的遗传变异潜力有关。本试验结果也进一步证实分布于较小范围内的红砂亚种群间的遗传距离、遗传多样性与亚种群的地理距离(经纬度差异)大小无显著的相关性。
同时，本文采用RAPD标记技术对分布于新疆阜康绿洲沙漠过渡带的重要藜科盐生半灌木无叶假木贼种群和一年生泌盐性草本角果藜种群的遗传多样性和遗传分化进行了研究。试验分别选择了16条随机引物，对同一生境中的无叶假木贼3个亚种群(58个个体)及角果藜4个亚种群(80个个体)进行RAPD扩增。研究结果表明：无叶假木贼种群的多态位点比率(PPB)为94.56％，角果藜种群的多态位点比率则为98％，显示出分布于过渡带的无叶假木贼和角果藜种群内均存在较高的遗传多样性，而角果藜种群遗传多样性水平高于无叶假木贼种群。经分析表明，无叶假木贼种群与角果藜种群的Shannon多样性指数分别为0.1403和0.1615，基因分化系数(G_(ST))为0.1469和0.1659，揭示了两物种种群遗传变异多存在于亚种群内，亚种群间的遗传分化则不明显。AMOVA分析说明无叶假木贼种群变异的87.59％存在于亚种群内，而亚种群间的变异占总变异的12.41％；而角果藜亚种群内与亚种群间的变异则分别占总变异的87.43％和12.57％。另外，通过RAPD资料的聚类分析及相关性分析研究，发现无叶假木贼和角果藜自然种群的遗传结构与绿洲沙漠过渡带的微生境生态因子(主要是土壤因子)相关，其中无叶假木贼亚种群遗传多样性水平不仅与土壤含水量(W)、钾钠离子浓度(K+Na)和氯离子浓度(Cl)呈显著的正相关(P＜0.05)，还与土壤中有机质(SOM)、全氮(Nt)和全磷(Pt)含量呈显著的负相关；同时，角果藜种群的遗传多样性水平与土壤中有机质(SOM)和全氮(Nt)含量呈显著的正相关，而与CO_3~(2-)浓度呈显著的负相关；除此之外，其它土壤生态因子与两物种遗传多样性水平的相关性均不显著
(卜0.05)。试验结果还进一步证实，分布于较小范围内的无叶假木贼和角果葵亚种群间的
遗传距离、遗传多样性与亚种群的地理距离(经纬度差异)大小无显著的相关性。研究通过
对同一生境、不同生活型的无叶假木贼和角果要两种盐生植物种群的遗传多样性进行分析比
较，探讨了两种植物的遗传多样性水平与生态生物学特点和进化过程的关系，为进一步全面
研究两种植物种群的遗传结构和进化适应提供参考。
另外，本文以构件理论为基础，运用灰色关联度分析技术，对新疆阜康荒漠红砂自然种
群构件结构及其与环境因子的关系进行了研究.结果表明:l)植株冠幅是表征红砂生长情
况的重要指标，随着红砂植株冠幅的增大，植株高度明显增加，而且植株冠幅与植株的一级
分枝系统和当年生枝系统的构件因子相关度较大.2)在某种意义上，一级分枝系统是红砂
植株的基础，三级分枝系统和当年生枝系统则是二级分枝系统的派生物，它们之间的关联性
显示了红砂种群构件组成间的整合作用。3)一级分枝系统内的分枝高度与分枝数、长度及
分枝角度有较大的关联度，但与径粗关联度相对较小，说明一级分枝系统内后者的变化与前
四者并不完全相似，表现出一定的异质性。4)在二级分枝系统中，一方面，各构件特征与
一级分枝系统存在较大的关联度，反映了红砂种群构件结构一定程度的整合作用;另一方面，
二级分枝构件系?
The paper presented a comprehensive survey of ecological genetic characteristics of Reawnuria soongorica, Anabasis aphylla and Ceratocarpus arenarius distributed in oasis- desert transitional zone in Fukang, Xinjiang. The relationships between the morphological, genetic, physiological and ecological variations and differentiations and the environmental factors were discussed to explain the mechanism of the ecological genetics during the process of evolution and formation of species and provide the theoretical basis and genetic background for planning the scientific strategies of biodiversity conservation.
Firstly, we studied the genetic structure and genetic differentiation of Reaumwia soongorica distributed in Fukang, Xinjiang, by the means of random amplified polymorphic DNA (RAPD) markers. The results showed that the higher genetic diversity and the genetic differentiation existed in the population of Reaumuria soongorica, which was composed of 7 subpopulations and 136 individuals, generated by 15 primers. 71 loci had been detected and 69 were polymorphic. The mean proportion of polymorphic loci (PPB) of Reawnuria soongorica was 97.18%. Furthermore, the study discussed the Shannon information index (0.3075), Nei's gene diversity index (0.3127) and Gst (0.3120), which indicated that there were more genetic variations within the subpopulations than those among the subpopulations. The gene flow among the subpopulations of Reawnuria soongorica (Mn=1.1028, Nm>1) was much less than that of the common anemophytes (ATm=5.24), so the species was in the critical state of genetic differentiation to some extent The results of AMOVA showed that there were 61.58% of the total genetic variations existing among the subpopulations, and 38.02% within them. Additionally, through the use of clustering analysis, principal factor analysis and the correlation analysis, we found that the genetic structure of natural population Reawnuria soongorica was related to the ecological factors (soil factors mainly) of the oasis-desert transitional zone. The genetic diversity level ofReaumuria soongorica had the negative correlation with the content of soil total P and Cl" significantly (P<0.05), on the contrary, it had the positive correlation with CO32- significantly (PO.05) but there was no significant correlation between the genetic diversity and the other salt factors in soil, which showed that the distribution of the individuals of Reawnuria soongorica correlated with some certain soluble salt. Furthermore, the study indicated that the genetic diversity of the natural population of Reawnuria soongorica increased with the decreasing of the content of organic matter, water, total N and total P in soil, which had some relation with more potentiality of the genetic and variation of Reawnuria soongorica in its long-term evolution to adapt worse environment. Besides, the paper demonstrated that there was no significant correlation between the genetic distance, genetic diversity and the geographical difference on latitude and longitude respectively within the small range.
Secondly, the genetic diversity and genetic differentiation of the two important desert plants of Chenpodiaceae, Anabasis aphylla and Ceratocarpus arenarius, distributed in Fukang Desert, Xinjiang, were studied using RAPD markers. The results showed that the mean proportion of polymorphic loci (PPB) of Anabasis aphylla, which comprised 3 subpopulations (58 individuals sampled), generated by 16 primers was 94.56%, the value of PPB of Ceratocarpus arenarius, which had 4 subpopulations (80 individuals sampled), using 16 primers was 98.00%. The paper had proved that the higher genetic diversity and the genetic differentiation existed in the populations of boih Anabasis aphylla and Ceratocarpus arenarius, while the latter had much more
genetic diversity than the former. Moreover, the study discussed the Shannon information index and Nei's gene diversity index of the two species natural populations, which indicated that there were more genetic variations within the subpopu
引文
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