中国东方蜜蜂资源遗传多样性分析
摘要
本研究首次系统地采集中国18个省、市、自治区的20个东方蜜蜂不同地理种群工蜂样本,建立了我国东方蜜蜂DNA基因库,并在此基础上应用形态标记、微卫星标记及线粒体序列变异对中国东方蜜蜂遗传多样性进行全面研究,分析群体间和群体内的遗传变异,评估群体遗传结构,并探讨彼此之间的亲缘关系,为进一步保护和利用我国东方蜜蜂遗传资源提供理论依据。主要试验结果如下:
1.本研究对工蜂的右前翅长与宽、吻长、肘脉指数、第3与4背板总长等形态指标进行测定,结果表明:20个东方蜜蜂种群之间在形态标记上存在显著或极显著差异(P <0.05或P <0.01)。西藏中蜂和迪庆中蜂右前翅长、宽及翅面积存在一个或多个指标的显著大于其它种群,表明它们具有更强的飞翔能力;从化中蜂除了与桐庐中蜂、西双版纳蜜蜂、阿坝中蜂、兴城中蜂、长白山中蜂之外,吻长均显著长于其它种群,表明其具有更强的花蜜觅食能力;长白山中蜂与西藏中蜂的第3、4背板总长显著长于其它种群,表明其具有更强的贮蜜能力。通过对形态指标相关性分析表明20个东方蜜蜂种群的右前翅长、前翅宽、前翅面积、吻长与第3、4背板总长均有极显著相关(P <0.01);肘脉指数与其它形态指标均不相关(P >0.05),表明肘脉指数是一个相对独立的形态标记。20个东方蜜蜂种群的6个形态标记和8项生态特征数据的主成分分析结果表明,不同东方蜜蜂种群形态指标的差异主要反映在包括形态标记和生态因素在内的7个主成分上,其累计贡献率达89.386%。主成分分析结果表明东方蜜蜂种群间差异分析除了考虑蜜蜂个体形态特征差异外,蜂群所生活地区的环境因素也应被纳入其中。形态学的结果表明中国东方蜜蜂形态指标存在丰富的遗传多样性。
2.本研究使用从西方蜜蜂上适用的56对微卫星引物中筛选出的21对微卫星引物对20个东方蜜蜂地理种群遗传多样性进行分析,共检测到502个等位基因,平均等位基因数为24.1430。21对微卫星位点的平均期望杂合度为0.8689±0.0525,PIC值为0.8564±0.0603,所有位点均具有较高的多态性,表明所筛选的微卫星标记能为分析东方蜜蜂不同地理种群遗传多样性提供充分的信息。20个地理种群的平均期望杂合度为0.8378±0.0136,显示出丰富的遗传多样性和较高的选择潜力。单个位点偏离Hardy-Weinberg平衡的群体数从6到20不等。群体间的平均遗传分化为42.3% (P <0.001),群体间存在着极显著的遗传分化。群体间的Reynolds'遗传距离从0.020(迪庆中蜂-武定中蜂)到1.085(北京中蜂-南昌中蜂)不等,而Nm值变异范围从0.128(南昌中蜂-北京中蜂)到12.376(迪庆中蜂-武定中蜂)。长白山中蜂、北京中蜂同其它种群的基因流动值总体很小,是相对隔绝的地理种群。
3.对中国20个东方蜜蜂地理种群mtDNA中tRNAleu-COⅡ片段的核苷酸序列进行分析,结果发现mtDNA COII基因部分序列中A、C、G、T这4种核苷酸的平均比例分别为38.2%、10.0%、4.6%和47.1%,序列中富含A+T,表现出碱基组成的偏倚性;测定非编码区序列A、C、G、T这4种核苷酸的平均比例分别为47.5%、8.3%、5.0%和39.2%,A+T含量也明显占优势,同样表现出碱基组成的偏倚性。COⅡ基因序列中共发现16个变异位点,约占分析位点总数的6.18%,没有检测到插入和缺失,颠换和转换之比为0.07。20个地理种群中只有武定中蜂、海南中蜂和南昌中蜂检测到颠换现象。非编码区序列中发现17个变异位点,约占分析位点总数的17.53%,其中插入/缺失位点2个,颠换和转换之比为0.25。20个地理种群中只有黄山中蜂、海南中蜂和费县中蜂检测到颠换现象。东方蜜蜂非编码区的变异位点比COⅡ基因多,表明东方蜜蜂非编码区变异程度高于COⅡ基因。
4.COⅡ基因序列中共检测到18种单倍型,其中5种为共享单倍型,其它13个单倍型均为各种群所特有,其中单倍型H2出现最多,为测试蜜蜂种群的主体单倍型。非编码区序列中共检测到22种单倍型,其中7种为共享单倍型,其它15个单倍型均为各种群所特有,其中单倍型H1包含的个体最多,为测试蜜蜂群体的主体单倍型。所有种群中,武定中蜂和醴县中蜂COⅡ基因和非编码区单倍型类型最多,均各为4个,长白山中蜂的单倍型类型最少,分别为1个和2个。20个东方蜜蜂群体内单倍型多样度差异较大,COⅡ基因的单倍型多样度从0到0.778不等,单倍型变异度总体为0.621±0.038;非编码区的单倍型多样度变异范围从0到0.778,单倍型变异度总体为0.699±0.035。20个东方蜜蜂种群COⅡ基因序列整体的平均核苷酸差异数为0.939,核苷酸多样度为0.851%;非编码区序列整体的平均核苷酸差异数为1.003,核苷酸多样度为1.034%。20个东方蜜蜂地理种群间mtDNA COⅡ基因序列核苷酸分歧度(Dxy)在0%～0.965%之间变化,非编码区序列核苷酸分歧度(Dxy)在0%～2.062%之间变化,核苷酸分歧度差异均很大;种群间mtDNA COⅡ基因序列核苷酸净遗传距离(Da)为-0.007%～0.781%,非编码区序列核苷酸净遗传距离(Da)为-0.037%～1.489%,核苷酸净遗传距离差异也很大。20个东方蜜蜂地理种群mtDNA COⅡ基因和非编码区序列群体间Fst分别为0.4978和0.4332,差异均极显著(P <0.001)。表明20个东方蜜蜂不同地理种群间存在显著的遗传分化。
5.利用21对微卫星标记对20个东方蜜蜂地理种群亲缘关系进行分析,NJ系统发生树和Sturcture程序运行结果显示20个种群中华东地区六个种群首先形成一大类,然后再与其它种群聚类,表明华东地区蜜蜂种群可能形成相对一致的遗传特征。黄山中蜂、阿坝中蜂、天水中蜂、海南中蜂、武定中蜂等多个种群遗传多样性较为丰富,可能与它们具有复杂的遗传基础有关。从化中蜂、南宁中蜂和醴县中蜂,西藏中蜂和西双版纳蜜蜂彼此之间的遗传基础十分类似,表明它们可能受到共同的起源影响。迪庆中蜂遗传多样性较低,是一个相对独立的类群,与西藏中蜂亲缘关系较远。西藏中蜂、海南中蜂与阿坝中蜂聚类分析时没有和中华蜜蜂的各类群区分开,表明三者可能不是独立于中华蜜蜂之外的亚种。黄山中蜂存在两个不同生态类群,并且与桐庐中蜂存在较大的基因流动。对20个种群mtDNACOⅡ基因和非编码区序列单倍型进行分子系统树和网络关系分析及利用Kimura双参数模型构建的分子系统树中,分析表明中国东方蜜蜂地理种群具有复杂的起源,利用MEGA3.1构建的与GENBANK中报道的东方蜜蜂各地理种群COⅡ及非编码区序列单倍型间的分子系统树也证实了这一点。综合微卫星DNA和mtDNA对中国东方蜜蜂20个种群的遗传多样性分析结果表明东方蜜蜂的群体遗传差异可能是由于某些种群在独特的生态环境下发生的,遗传分化仅仅是类群间的差异,而不是亚种水平上的分化,因而在东方蜜蜂资源保护中应充分根据不同地理种群的特征制定科学的保护措施。
6.利用微卫星标记和mtDNA标记分析20个东方蜜蜂种群间遗传距离和地理距离的关联性,20个地理种群遗传距离与地理距离回归公式:Fst/(1-Fst) = 0.209 +0.084ln (d)以及Mantel’s检验结果(P=0.055)并不能为遗传距离与地理距离之间的显著联系提供足够的证据。线粒体COⅡ基因和非编码区序列的差异与种群间地理距离和种群地理分布也没有相关。研究结果说明我国东方蜜蜂地理种群的形成过程中,各自的地理分布并不是影响其群体遗传结构的决定因素。
Genetic diversity of 20 Apis cerana populations from 18 provinces, cities and municipalities in China were established to DNA GenBank of Apis cerana in China and evaluated with morphology markers, microsatellite markers and mtDNA sequences firstly and roundly. Genetic variability within populations, population structure and the relationship of them were estimated to provide academic reference for farther protection and utilize of Apis cerana in China. The main results were summarized as follows:
1.The 6 main indexes of external morphology such as the length and width of the anterior wing, the cubital vein indexes, the length of proboscis and the length of terga 3+4 were detected. There were significant differences (P <0.05 and P <0.01) among 6 characteristics of 20 populations in China. Tibet bee and Diqing bee had differences in one or more characters in length of bee right forewing, width of bee right forewing, area of the anterior wing from other populations, indicating better collecting ability; there were no difference in length of proboscis among Conghua bee, Tonglu bee, Xishuangbanna bee, A-ba bee, Xingcheng bee and Changbaishan bee, and their values were higher than other ones; Changbaishan bee and Tibet bee had the longest length of terga 3+4, which indicated that they had the best honey storing capability. The research result showed that significant difference (P <0.01) existed in the length, width and area of the anterior wing ,the length of proboscis and the length of terga 3+4; there was no difference between the cubital vein indexes and other characters(P>0.05), which indicated that cubital vein indexes was an independent morphological features. The principal component analysis (PCA)of the 6 main indexes of external morphological and 8 ecological features of 20 populations showed that the difference among morphological features of 20 populations were performed by 7 principal components included both morphological features and ecological features, and contribution rate of the two was 89.386%. Not only induviduals’morphological features but also ecological features should be considered in principal component analysis.
2. 21 microsatellite markers suited for Apis cerana were selected from 56 microsatellite markers used in Apis mellifera to be used to analyze genetic diversity of 20 Apis cerana populations. 502 alleles were found, and the average value was 24.1430. The average value of gene heterozygosity (He) of 21 microsatellite loci was 0.8689±0.0525, The PIC was 0.8564±0.0603. All 21 microsatellite loci in this study showed high levels of polymorphism. The average value of gene heterozygosity (He) of all populations was 0.8378±0.0136, showed rich genetic diversity and higher selected potentiality. The number of populations deviated from Hardy-Weinberg equilibrium per locus ranged from 6 to 20. The average value of Fst was 42.3% (P <0.001), and all loci were contributed significantly (P <0.001) to this differentiation. There was significant genetic differentiation among 20 populations. Reynolds’distance values varied between 0.020 (Diqing bee-Wuding bee pair) and 1.085 (Beijing bee-Nanchang bee pair). The Nm value was ranged from 0.128 (between Nanchang bee and Beijing bee) to 12.376 (between Diqing bee and Wuding bee). The Nm value between Changbai bee, Beijing bee and the others were low, so they maybe separate populations from the others.
3. Seqence of mtDNA tRNAleu-COⅡo f 20 Apis cerana populations were sequenced and analyzed. The result showed the average ratio of A、C、G、T in the sequence of mtDNA COⅡwas 38.2%、10.0%、4.6%、47.1%, respectively. High A+T was contained in the sequence demonstrating the bias of base composition. The average ratio of A、C、G、T of noncoding area was 47.5%、8.3%、5.0%、39.2%. Similarly, High A+T showed the bias of base composition. There were 16 polymorphic sites represent 6.18 % of total analyzed sites in COⅡ, no insertion/deletion were found in this region, the ratio of transition and transversion in this study was 0.07.There were 17 polymorphic sites represent 17.53% of total analyzed sites in noncoding area, contained 2 insertion/deletion, and the ratio of transition and transversion in this study was 0.25. Transitions were only found in Huangshan bee, Hainan bee and Feixian bee. The variability measure of noncoding area of Apis cerana populations was more than COⅡ.
4. 18 haplotypes were found in COⅡsequence,among which 5 haplotypes were shared among some populations, 13 haplotypes were unique for one population. H2 appeared the most times which was the main haplotype of 20 populations. 22 haplotypes were found in noncoding aera,among which 7 haplotypes were shared among some populations, 15 haplotypes were unique for one population. H1 concluded the most individuals, was the main haplotype to test the population of honeybee. The result showed that, Wuding bee and fengxian bee had the most haplotypes in the noncoding area of COⅡ, and all were 4; Changbaishan bee had the fewest, 1 and 2 respectively. The distribution of all haplotypes among the populations was disequilibrium and the diversity of haplotypes in COⅡwas ranged from 0 to 0.778, the average diversity of haplotypes was 0.621±0.038; The diversity of haplotypes in noncoding area was ranged from 0 to 0.778.The average diversity of haplotypes was 0.699±0.035. The average number of nucleotide divergence (K) and average nucleotide diversity (Pi) of COⅡwere 0.939 and 0.851%, respectively. The average number of nucleotide divergence (K) and average nucleotide diversity (Pi) of noncoding area were 1.003 and 1.034%, respectively. Inter-population Nucleotide Divergence (Dxy) of COⅡin 20 Apis cerana populations was ranged from 0%～0.965%, Inter-population Nucleotide Divergence (Dxy) of noncoding area in 20 Apis cerana populations was ranged from 0%～2.062%. The results indicated that the genetic diversity of 20 Apis cerana populations was very abundant.Inter-population Net Nucleotide Divergence (Da) in COⅡin 20 Apis cerana populations was ranged from -0.007%～0.781%, Inter-population Net Nucleotide Divergence (Da) in COⅡin 20 Apis cerana populations was ranged from -0.037%～1.489%. Fst value of mtDNA COⅡand noncoding area of 20 Apis cerana populations was 0.4978 and 0.4332 respectively, which indicated the genetic variation was extremely significant within populations (P < 0.001). There was significant divergence among 20 Apis cerana populations.
5. The phylogenetic relationship among 20 Apis cerana populations in China was analyzed with 21 microsatellite loci. The 6 populations from Eastern China fell together in the first place by NJ phylogenetic tree and Sturcture procedure, and the reason may be that Apis cerana from Eastern China had formed the same genetic characters. nshui bee, Hainan bee, Wuding bee), related to their complex genetic basis. Populations from Conghua, Nanning and Fengxian, populations from Tibet and Xishuangbanna had similar genetic basis, and the results suggested that they were affected by the same phylogenetic origin. The genetic diversity of Diqing bee was low, it maybe a separate population, and had great far relationship with Tibet bee. Huangshan bee had two different ecological groups, and had more genetic exchanges. Analysis of haplotypes of mtDNA COⅡand noncoding area among 20 populations estimated with molecular phylogenetic trees and network and molecular phylogenetic trees based on Kimura-2 parameters indicated that there was a complex origin for Apis cerena cerena populations, which was validated by molecular phylogenetic trees of the haplotypes of mtDNA COⅡand noncoding area of Apis cerena both from GENBANK and our research. Analysis of polymorphism and genetic relationship among 20 Apis cerena cerena populations estimated with 21 microsatellite loci and mtDNA sequences indicated that the genetic diversity were occurred as a result of certain geographic populations. Genetic diversity was only difference among different populations, rather than differentiation of the subspecies. So more reasonable measures should be made according to the charaters of different geographical populations to protect and make use of the Apis cerena resource.
6. Analysis of genetic distance and geographical distance among 20 Apis cerena cerena populations were estimated with 21 microsatellite loci and mtDNA. The result suggested that the equation Fst/ (1-Fst) = 0.209 +0.084ln (d) and the result from Mantel’s test (P=0.055) did not provide enough support for a significant correlation between the genetic and geographical pair wise distances. There were no significant correlations between the genetic diversity of mtDNA D-loop and sequence of noncoding area and the distributing of these populations. The results concluded that the geographical distributing maybe not the determinant influence on the genetic structure of Apis cerena cerena populations during the course of their developed history.
1.本研究对工蜂的右前翅长与宽、吻长、肘脉指数、第3与4背板总长等形态指标进行测定,结果表明:20个东方蜜蜂种群之间在形态标记上存在显著或极显著差异(P <0.05或P <0.01)。西藏中蜂和迪庆中蜂右前翅长、宽及翅面积存在一个或多个指标的显著大于其它种群,表明它们具有更强的飞翔能力;从化中蜂除了与桐庐中蜂、西双版纳蜜蜂、阿坝中蜂、兴城中蜂、长白山中蜂之外,吻长均显著长于其它种群,表明其具有更强的花蜜觅食能力;长白山中蜂与西藏中蜂的第3、4背板总长显著长于其它种群,表明其具有更强的贮蜜能力。通过对形态指标相关性分析表明20个东方蜜蜂种群的右前翅长、前翅宽、前翅面积、吻长与第3、4背板总长均有极显著相关(P <0.01);肘脉指数与其它形态指标均不相关(P >0.05),表明肘脉指数是一个相对独立的形态标记。20个东方蜜蜂种群的6个形态标记和8项生态特征数据的主成分分析结果表明,不同东方蜜蜂种群形态指标的差异主要反映在包括形态标记和生态因素在内的7个主成分上,其累计贡献率达89.386%。主成分分析结果表明东方蜜蜂种群间差异分析除了考虑蜜蜂个体形态特征差异外,蜂群所生活地区的环境因素也应被纳入其中。形态学的结果表明中国东方蜜蜂形态指标存在丰富的遗传多样性。
2.本研究使用从西方蜜蜂上适用的56对微卫星引物中筛选出的21对微卫星引物对20个东方蜜蜂地理种群遗传多样性进行分析,共检测到502个等位基因,平均等位基因数为24.1430。21对微卫星位点的平均期望杂合度为0.8689±0.0525,PIC值为0.8564±0.0603,所有位点均具有较高的多态性,表明所筛选的微卫星标记能为分析东方蜜蜂不同地理种群遗传多样性提供充分的信息。20个地理种群的平均期望杂合度为0.8378±0.0136,显示出丰富的遗传多样性和较高的选择潜力。单个位点偏离Hardy-Weinberg平衡的群体数从6到20不等。群体间的平均遗传分化为42.3% (P <0.001),群体间存在着极显著的遗传分化。群体间的Reynolds'遗传距离从0.020(迪庆中蜂-武定中蜂)到1.085(北京中蜂-南昌中蜂)不等,而Nm值变异范围从0.128(南昌中蜂-北京中蜂)到12.376(迪庆中蜂-武定中蜂)。长白山中蜂、北京中蜂同其它种群的基因流动值总体很小,是相对隔绝的地理种群。
3.对中国20个东方蜜蜂地理种群mtDNA中tRNAleu-COⅡ片段的核苷酸序列进行分析,结果发现mtDNA COII基因部分序列中A、C、G、T这4种核苷酸的平均比例分别为38.2%、10.0%、4.6%和47.1%,序列中富含A+T,表现出碱基组成的偏倚性;测定非编码区序列A、C、G、T这4种核苷酸的平均比例分别为47.5%、8.3%、5.0%和39.2%,A+T含量也明显占优势,同样表现出碱基组成的偏倚性。COⅡ基因序列中共发现16个变异位点,约占分析位点总数的6.18%,没有检测到插入和缺失,颠换和转换之比为0.07。20个地理种群中只有武定中蜂、海南中蜂和南昌中蜂检测到颠换现象。非编码区序列中发现17个变异位点,约占分析位点总数的17.53%,其中插入/缺失位点2个,颠换和转换之比为0.25。20个地理种群中只有黄山中蜂、海南中蜂和费县中蜂检测到颠换现象。东方蜜蜂非编码区的变异位点比COⅡ基因多,表明东方蜜蜂非编码区变异程度高于COⅡ基因。
4.COⅡ基因序列中共检测到18种单倍型,其中5种为共享单倍型,其它13个单倍型均为各种群所特有,其中单倍型H2出现最多,为测试蜜蜂种群的主体单倍型。非编码区序列中共检测到22种单倍型,其中7种为共享单倍型,其它15个单倍型均为各种群所特有,其中单倍型H1包含的个体最多,为测试蜜蜂群体的主体单倍型。所有种群中,武定中蜂和醴县中蜂COⅡ基因和非编码区单倍型类型最多,均各为4个,长白山中蜂的单倍型类型最少,分别为1个和2个。20个东方蜜蜂群体内单倍型多样度差异较大,COⅡ基因的单倍型多样度从0到0.778不等,单倍型变异度总体为0.621±0.038;非编码区的单倍型多样度变异范围从0到0.778,单倍型变异度总体为0.699±0.035。20个东方蜜蜂种群COⅡ基因序列整体的平均核苷酸差异数为0.939,核苷酸多样度为0.851%;非编码区序列整体的平均核苷酸差异数为1.003,核苷酸多样度为1.034%。20个东方蜜蜂地理种群间mtDNA COⅡ基因序列核苷酸分歧度(Dxy)在0%～0.965%之间变化,非编码区序列核苷酸分歧度(Dxy)在0%～2.062%之间变化,核苷酸分歧度差异均很大;种群间mtDNA COⅡ基因序列核苷酸净遗传距离(Da)为-0.007%～0.781%,非编码区序列核苷酸净遗传距离(Da)为-0.037%～1.489%,核苷酸净遗传距离差异也很大。20个东方蜜蜂地理种群mtDNA COⅡ基因和非编码区序列群体间Fst分别为0.4978和0.4332,差异均极显著(P <0.001)。表明20个东方蜜蜂不同地理种群间存在显著的遗传分化。
5.利用21对微卫星标记对20个东方蜜蜂地理种群亲缘关系进行分析,NJ系统发生树和Sturcture程序运行结果显示20个种群中华东地区六个种群首先形成一大类,然后再与其它种群聚类,表明华东地区蜜蜂种群可能形成相对一致的遗传特征。黄山中蜂、阿坝中蜂、天水中蜂、海南中蜂、武定中蜂等多个种群遗传多样性较为丰富,可能与它们具有复杂的遗传基础有关。从化中蜂、南宁中蜂和醴县中蜂,西藏中蜂和西双版纳蜜蜂彼此之间的遗传基础十分类似,表明它们可能受到共同的起源影响。迪庆中蜂遗传多样性较低,是一个相对独立的类群,与西藏中蜂亲缘关系较远。西藏中蜂、海南中蜂与阿坝中蜂聚类分析时没有和中华蜜蜂的各类群区分开,表明三者可能不是独立于中华蜜蜂之外的亚种。黄山中蜂存在两个不同生态类群,并且与桐庐中蜂存在较大的基因流动。对20个种群mtDNACOⅡ基因和非编码区序列单倍型进行分子系统树和网络关系分析及利用Kimura双参数模型构建的分子系统树中,分析表明中国东方蜜蜂地理种群具有复杂的起源,利用MEGA3.1构建的与GENBANK中报道的东方蜜蜂各地理种群COⅡ及非编码区序列单倍型间的分子系统树也证实了这一点。综合微卫星DNA和mtDNA对中国东方蜜蜂20个种群的遗传多样性分析结果表明东方蜜蜂的群体遗传差异可能是由于某些种群在独特的生态环境下发生的,遗传分化仅仅是类群间的差异,而不是亚种水平上的分化,因而在东方蜜蜂资源保护中应充分根据不同地理种群的特征制定科学的保护措施。
6.利用微卫星标记和mtDNA标记分析20个东方蜜蜂种群间遗传距离和地理距离的关联性,20个地理种群遗传距离与地理距离回归公式:Fst/(1-Fst) = 0.209 +0.084ln (d)以及Mantel’s检验结果(P=0.055)并不能为遗传距离与地理距离之间的显著联系提供足够的证据。线粒体COⅡ基因和非编码区序列的差异与种群间地理距离和种群地理分布也没有相关。研究结果说明我国东方蜜蜂地理种群的形成过程中,各自的地理分布并不是影响其群体遗传结构的决定因素。
Genetic diversity of 20 Apis cerana populations from 18 provinces, cities and municipalities in China were established to DNA GenBank of Apis cerana in China and evaluated with morphology markers, microsatellite markers and mtDNA sequences firstly and roundly. Genetic variability within populations, population structure and the relationship of them were estimated to provide academic reference for farther protection and utilize of Apis cerana in China. The main results were summarized as follows:
1.The 6 main indexes of external morphology such as the length and width of the anterior wing, the cubital vein indexes, the length of proboscis and the length of terga 3+4 were detected. There were significant differences (P <0.05 and P <0.01) among 6 characteristics of 20 populations in China. Tibet bee and Diqing bee had differences in one or more characters in length of bee right forewing, width of bee right forewing, area of the anterior wing from other populations, indicating better collecting ability; there were no difference in length of proboscis among Conghua bee, Tonglu bee, Xishuangbanna bee, A-ba bee, Xingcheng bee and Changbaishan bee, and their values were higher than other ones; Changbaishan bee and Tibet bee had the longest length of terga 3+4, which indicated that they had the best honey storing capability. The research result showed that significant difference (P <0.01) existed in the length, width and area of the anterior wing ,the length of proboscis and the length of terga 3+4; there was no difference between the cubital vein indexes and other characters(P>0.05), which indicated that cubital vein indexes was an independent morphological features. The principal component analysis (PCA)of the 6 main indexes of external morphological and 8 ecological features of 20 populations showed that the difference among morphological features of 20 populations were performed by 7 principal components included both morphological features and ecological features, and contribution rate of the two was 89.386%. Not only induviduals’morphological features but also ecological features should be considered in principal component analysis.
2. 21 microsatellite markers suited for Apis cerana were selected from 56 microsatellite markers used in Apis mellifera to be used to analyze genetic diversity of 20 Apis cerana populations. 502 alleles were found, and the average value was 24.1430. The average value of gene heterozygosity (He) of 21 microsatellite loci was 0.8689±0.0525, The PIC was 0.8564±0.0603. All 21 microsatellite loci in this study showed high levels of polymorphism. The average value of gene heterozygosity (He) of all populations was 0.8378±0.0136, showed rich genetic diversity and higher selected potentiality. The number of populations deviated from Hardy-Weinberg equilibrium per locus ranged from 6 to 20. The average value of Fst was 42.3% (P <0.001), and all loci were contributed significantly (P <0.001) to this differentiation. There was significant genetic differentiation among 20 populations. Reynolds’distance values varied between 0.020 (Diqing bee-Wuding bee pair) and 1.085 (Beijing bee-Nanchang bee pair). The Nm value was ranged from 0.128 (between Nanchang bee and Beijing bee) to 12.376 (between Diqing bee and Wuding bee). The Nm value between Changbai bee, Beijing bee and the others were low, so they maybe separate populations from the others.
3. Seqence of mtDNA tRNAleu-COⅡo f 20 Apis cerana populations were sequenced and analyzed. The result showed the average ratio of A、C、G、T in the sequence of mtDNA COⅡwas 38.2%、10.0%、4.6%、47.1%, respectively. High A+T was contained in the sequence demonstrating the bias of base composition. The average ratio of A、C、G、T of noncoding area was 47.5%、8.3%、5.0%、39.2%. Similarly, High A+T showed the bias of base composition. There were 16 polymorphic sites represent 6.18 % of total analyzed sites in COⅡ, no insertion/deletion were found in this region, the ratio of transition and transversion in this study was 0.07.There were 17 polymorphic sites represent 17.53% of total analyzed sites in noncoding area, contained 2 insertion/deletion, and the ratio of transition and transversion in this study was 0.25. Transitions were only found in Huangshan bee, Hainan bee and Feixian bee. The variability measure of noncoding area of Apis cerana populations was more than COⅡ.
4. 18 haplotypes were found in COⅡsequence,among which 5 haplotypes were shared among some populations, 13 haplotypes were unique for one population. H2 appeared the most times which was the main haplotype of 20 populations. 22 haplotypes were found in noncoding aera,among which 7 haplotypes were shared among some populations, 15 haplotypes were unique for one population. H1 concluded the most individuals, was the main haplotype to test the population of honeybee. The result showed that, Wuding bee and fengxian bee had the most haplotypes in the noncoding area of COⅡ, and all were 4; Changbaishan bee had the fewest, 1 and 2 respectively. The distribution of all haplotypes among the populations was disequilibrium and the diversity of haplotypes in COⅡwas ranged from 0 to 0.778, the average diversity of haplotypes was 0.621±0.038; The diversity of haplotypes in noncoding area was ranged from 0 to 0.778.The average diversity of haplotypes was 0.699±0.035. The average number of nucleotide divergence (K) and average nucleotide diversity (Pi) of COⅡwere 0.939 and 0.851%, respectively. The average number of nucleotide divergence (K) and average nucleotide diversity (Pi) of noncoding area were 1.003 and 1.034%, respectively. Inter-population Nucleotide Divergence (Dxy) of COⅡin 20 Apis cerana populations was ranged from 0%～0.965%, Inter-population Nucleotide Divergence (Dxy) of noncoding area in 20 Apis cerana populations was ranged from 0%～2.062%. The results indicated that the genetic diversity of 20 Apis cerana populations was very abundant.Inter-population Net Nucleotide Divergence (Da) in COⅡin 20 Apis cerana populations was ranged from -0.007%～0.781%, Inter-population Net Nucleotide Divergence (Da) in COⅡin 20 Apis cerana populations was ranged from -0.037%～1.489%. Fst value of mtDNA COⅡand noncoding area of 20 Apis cerana populations was 0.4978 and 0.4332 respectively, which indicated the genetic variation was extremely significant within populations (P < 0.001). There was significant divergence among 20 Apis cerana populations.
5. The phylogenetic relationship among 20 Apis cerana populations in China was analyzed with 21 microsatellite loci. The 6 populations from Eastern China fell together in the first place by NJ phylogenetic tree and Sturcture procedure, and the reason may be that Apis cerana from Eastern China had formed the same genetic characters. nshui bee, Hainan bee, Wuding bee), related to their complex genetic basis. Populations from Conghua, Nanning and Fengxian, populations from Tibet and Xishuangbanna had similar genetic basis, and the results suggested that they were affected by the same phylogenetic origin. The genetic diversity of Diqing bee was low, it maybe a separate population, and had great far relationship with Tibet bee. Huangshan bee had two different ecological groups, and had more genetic exchanges. Analysis of haplotypes of mtDNA COⅡand noncoding area among 20 populations estimated with molecular phylogenetic trees and network and molecular phylogenetic trees based on Kimura-2 parameters indicated that there was a complex origin for Apis cerena cerena populations, which was validated by molecular phylogenetic trees of the haplotypes of mtDNA COⅡand noncoding area of Apis cerena both from GENBANK and our research. Analysis of polymorphism and genetic relationship among 20 Apis cerena cerena populations estimated with 21 microsatellite loci and mtDNA sequences indicated that the genetic diversity were occurred as a result of certain geographic populations. Genetic diversity was only difference among different populations, rather than differentiation of the subspecies. So more reasonable measures should be made according to the charaters of different geographical populations to protect and make use of the Apis cerena resource.
6. Analysis of genetic distance and geographical distance among 20 Apis cerena cerena populations were estimated with 21 microsatellite loci and mtDNA. The result suggested that the equation Fst/ (1-Fst) = 0.209 +0.084ln (d) and the result from Mantel’s test (P=0.055) did not provide enough support for a significant correlation between the genetic and geographical pair wise distances. There were no significant correlations between the genetic diversity of mtDNA D-loop and sequence of noncoding area and the distributing of these populations. The results concluded that the geographical distributing maybe not the determinant influence on the genetic structure of Apis cerena cerena populations during the course of their developed history.
引文
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