Belokon' M M, Belokon' Iu S, Politov D V, Altukhov Iu P
Genetika. 2005 Nov;41(11):1538-51.
Swiss stone pine Pinus cembra L. is a species with fragmented range, occurring in the Alpine-East Carpathian mountain system. Seeds of P. cembra are dispersed by nutcrackers, which offers potential possibilities for gene exchange among populations. Using isozyme analysis, we have examined five samples from two parts of the Swiss stone pine range: the Alps (Switzerland and Austria) and the Carpathians (two samples from the northern macroslope of the Gorgany Ridge, Eastern Carpathians, Ivano-Frankovsk oblast and one sample from Trans-Carpathian oblast of Ukraine). The allele frequencies of 30 isozyme loci, coding for enzymes ADH, FDH, FEST, GDH, GOT, IDH, LAP, MNR, MDH, PEPCA, 6PGD, PGI, PGM, SDH, SKDH, SOD, were analyzed using cluster analysis and methods of principal components. Two clusters, corresponding to the isolated Alpine and Carpathian parts of the range, were found. The main contribution to these differences were made by loci Adh-1, Adh-2, Fest-2, Lap-3, Mdh-4, and Sod-4. The interpopulation differentiation proved to be somewhat higher than that typical for pines (F(ST) = 7.4%), but within the limits characteristic for taxonomically close species. Thus, isolation of the populations did not lead to their marked differentiation, which may be explained by gene flow and balancing selection, which equalizes gene frequencies across the fragmented species area. Interlocus F(ST) heterogeneity (from 0.003 to 0.173) suggests adaptive significance of some of the allozyme polymorphisms or linkage of some loci with adaptive genes. The Carpathian populations were shown to have higher gene diversity than the Alpine ones (expected heterozygosities 0.095-0.114 and 0.060-0.080, respectively). A deficit of heterozygotes (as compared to the Hardy-Weinberg proportions), observed in the embryo sample, was probably explained by inbreeding. The reduction in the area of Carpathian pine forests in Holocene, caused by the global climatic changes and the anthropogenic impact, is hazardous for the gene pool of the species. The maintenance of genetic uniqueness of both Carpathian populations of P. cembra in general, and individual stands in particular, requires special measures for protection of Swiss stone pine in the Eastern Carpathians.
瑞士石松(Pinus cembra L.)是一种分布范围破碎化的物种,生长在阿尔卑斯 - 东喀尔巴阡山脉系统中。瑞士石松的种子由坚果雀传播,这为种群间的基因交流提供了潜在可能性。我们利用同工酶分析,检测了来自瑞士石松分布范围两个区域的五个样本:阿尔卑斯山地区(瑞士和奥地利)以及喀尔巴阡山脉地区(两个样本来自东喀尔巴阡山脉伊万诺 - 弗兰科夫斯克州戈尔加尼岭北宏坡,一个样本来自乌克兰外喀尔巴阡州)。使用聚类分析和主成分分析法,分析了编码乙醇脱氢酶(ADH)、甲酸脱氢酶(FDH)、酯酶(FEST)、谷氨酸脱氢酶(GDH)、谷草转氨酶(GOT)、异柠檬酸脱氢酶(IDH)、亮氨酸氨肽酶(LAP)、锰依赖的核酸酶(MNR)、苹果酸脱氢酶(MDH)、磷酸烯醇式丙酮酸羧化酶(PEPCA)、6 - 磷酸葡萄糖脱氢酶(6PGD)、磷酸葡萄糖异构酶(PGI)、磷酸葡萄糖变位酶(PGM)、琥珀酸脱氢酶(SDH)、琥珀酰 - 辅酶A合成酶(SKDH)、超氧化物歧化酶(SOD)的30个同工酶位点的等位基因频率。发现了两个聚类,分别对应该分布范围中孤立的阿尔卑斯山和喀尔巴阡山脉区域。造成这些差异的主要位点是Adh - 1、Adh - 2、Fest - 2、Lap - 3、Mdh - 4和Sod - 4。种群间分化程度略高于松树的典型分化程度(F(ST) = 7.4%),但仍在分类学上相近物种的特征范围内。因此,种群隔离并未导致其显著分化,这可能是由于基因流和平衡选择,使破碎化物种区域内的基因频率趋于均衡。位点间F(ST)异质性(从0.003到0.173)表明一些等位酶多态性具有适应性意义,或者某些位点与适应性基因存在连锁关系。结果表明,喀尔巴阡山脉种群的基因多样性高于阿尔卑斯山种群(预期杂合度分别为0.095 - 0.114和0.06