Haplotype and genetic diversity analysis of Spodoptera frugiperda invading in three provinces of central and southern China
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摘要:目的
揭示入侵我国中南三省(区)的草地贪夜蛾Spodoptera frugiperda种群间可能存在的遗传变异，探讨其入侵来源、入侵路线和扩散方式，为制定有效控制和阻击草地贪夜蛾的方案提供理论依据和遗传信息。
方法利用分子标记技术从线粒体DNA和核DNA角度研究草地贪夜蛾种群的遗传多样性和种群结构。对采集自我国广东、广西和湖南3个省(区)的草地贪夜蛾，利用CO I和Tpi基因片段进行了单倍体型分析，利用9个微卫星标记分析种群间的遗传结构和遗传多样性。
结果供试样品中，除了湖南张家界群体有2个样品为CO I玉米型单倍体型、1个样品为杂合型，其他群体的个体均为CO I水稻型单倍体型。基于SSR标记的群体遗传分析结果表明，被检测的群体遗传多样性较巴西等国家的美洲群体低，各群体遗传分化不明显。广东东莞群体与湖南张家界群体的遗传距离最近，与广西南宁群体的遗传距离较远。
结论遗传距离与实际距离之间不一定呈正相关，通过遗传结构的分析有助于推测草地贪夜蛾扩散过程中，天气背景场、地理隔离和人为传播各自所起的作用。入侵湖南省张家界的草地贪夜蛾出现了低频率CO I玉米型和高频率的CO I水稻型单倍体型，说明虫源的非单一性；草地贪夜蛾在入侵地定殖时间较短，目前未形成明显的遗传分化。
Abstract:ObjectiveTo reveal possible genetic variation among Spodoptera frugiperda populations in three provinces of central and southern China, trace the invasion source and spread pathway of S. frugiperd, and provide the theoretical basis and genetic information for effectively controlling this insect.
MethodThe samples of S. frugiperda were collected from Guangdong, Guangxi and Hunan of China. Haplotype analysis was carried out using CO I and Tpigene markers. Nine microsatellite (SSR) markers were used to analyze the genetic diversity and genetic structure of S. frugiperda populations.
ResultsExpect that two samples of S. frugiperda populations from Zhangjiajie of Hunan were characterized to be CO I corn strain and one sample to be hybrid strain, other samples were CO I rice train. The results of SSR markers showed that lower genetic diversity was detected for each population compared with those in South America populations, and the genetic differentiation across populations was no significant. However, the genetic distance was the nearest between Guangdong Dongguan population and Hunan Zhangjiajie population, and the farthest between Guangdong Dongguan population and Guangxi Nanning population.
ConclusionThere is no significant relationship between genetic distance and geographic distance. The genetic structure analysis helps to speculate on the role of weather background field, geographical isolation and human transmission in the process of S. frugiperda spread. S. frugiperda invading in central-south China might be original from multiple locations. The short time of S. frugiperda colonization in the invasive lands leads to the low genetic diversity. No obvious genetic differentiation has formed at present.
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Keywords:
- Spodoptera frugiperda /
- haplotype /
- SSR marker /
- genetic diversity
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图 1 草地贪夜蛾群体间Nei’ s标准遗传距离的UPGMA法聚类图
Figure 1. UPGMA cluster analysis using the Nei’ s standard genetic distance indices for Spodoptera frugiperda
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图 2 基于SSR标记的草地贪夜蛾群体的主坐标分析
1：湖南张家界群体为主；2：广东三地群体为主；3：广西南宁群体为主
Figure 2. Principal coordinate analysi based on SSR markers for Spodoptera frugiperda populations
1：The populations were mainly from Zhangjiajie of Hunan；2：The populations were mainly from three cities (district) of Guangdong；3：The populations were mainly from Nanning of Guangxi
表 1 供试幼虫样本信息表
Table 1 Information of larva samples used in this study
样品编号
Sample code采集地点
Sampling site采集时间
Sampling date虫态
Stage寄主植物
Host数量/头
NumberGDHD 广东省广州市花都花山镇儒林村 2019−05−10 幼虫 玉米 15 HNZJJ 湖南省张家界市慈利县通津钱铺 2019−05−20 幼虫 玉米 11 GXNN 广西壮族自治区南宁市宾阳县古济 2019−05−21 幼虫 玉米 7 GDDG 广东省东莞市 2019−05−01 幼虫 玉米 3 GDQY 广东省清远市连山壮族瑶族自治县 2019−06−16 幼虫 玉米 1 GDSG 广东省韶关市翁源县 2019−06−19 幼虫 甘蔗 5 合计 Total 42 
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表 2 草地贪夜蛾2种单倍型CO I基因序列多态性位点比较
Table 2 Polymorphic site distribution of CO I gene sequence of haplotypes in Spodoptera frugiperda populations
单倍体型1)
Haplotype多态性位点 Polymorphic site 33 69 120 351 426 432 462 492 HM136592(CS) T T C T T C C T HNZJJ-30# T T C T T C C T HM136602(RS) C A T C C T T C GDHD-7# C A T C C T T C 1)HM136592(CS)：草地贪夜蛾玉米型；HNZJJ-30#：湖南张家界30#样本；HM136602(RS)：草地贪夜蛾水稻型；GDHD-7#：广州花都7#样本
1) HM136592(CS)：S. frugiperda corn strain；HNZJJ-30#：Sample 30# from Zhangjiajie of Hunan；HM136602(RS)：S. frugiperda rice strain；GDHD-7#：Sample 7# from Huadu of Guangdong
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表 3 CO I和Tpi标记鉴定的草地贪夜蛾42个样品单倍型样品数量
Table 3 The number of haplotype of 42 Spodoptera frugiperda samples identified by COI and Tpi markers
采样地点
Sampling siteCO I 标记 CO I marker Tpi 标记 Tpi marker 水稻型
Rice strain玉米型
Corn strain杂合型 (S. f/M. s)
Hybrid strain玉米型 (AT)
Rice strain水稻型 (GA)
Corn strain杂合型 (AT/GA)
Hybrid strain广东花都 GDHD 15 0 0 9 3 3 湖南张家界 HNZJJ 8 2 1 6 0 5 广西南宁 GXNN 7 0 0 6 0 1 广东东莞 GDDG 3 0 0 3 0 0 广东清远 GDQY 1 0 0 1 0 0 广东韶关 GDSG 5 0 0 2 0 3 合计 Total 39 2 1 27 3 12
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表 4 9个SSR位点的遗传变异参数、F-statistics统计值和基因流1)
Table 4 Indices of gene diversity，F-statistics values and gene flow deduced by nine SSR loci
SSR位点 Site NA NE I HO HE FIS FIT FST NM spf05 3.000 1.718 0.62 0.221 0.338 0.346 0.465 0.183 1.117 spf1502 3.500 1.832 0.647 0.323 0.334 0.035 0.271 0.244 0.773 spf1592 6.667 5.000 1.633 0.958 0.762 −0.258 −0.13 0.102 2.206 spf1706 2.167 1.283 0.337 0.217 0.186 −0.163 −0.048 0.099 2.269 spf343 4.667 3.524 1.291 0.92 0.686 −0.342 −0.188 0.115 1.929 spf670 5.500 3.887 1.376 0.824 0.681 −0.209 −0.015 0.160 1.312 spf789 5.500 4.340 1.437 0.93 0.707 −0.315 −0.111 0.156 1.357 spf918 4.167 2.737 1.125 0.824 0.608 −0.354 −0.227 0.094 2.407 spf997 4.500 2.856 1.135 0.536 0.612 0.125 0.303 0.204 0.976 平均值 Average 4.408 3.020 1.067 0.639 0.546 −0.126 0.036 0.151 1.594 1) NA: 观测等位基因数; NE: 有效等位基因数; I: Shannon信息指数; HO: 表观杂合度; HE: 预期杂合度; FIS: 种群内近交系数; FIT: 总近交系数; FST: 种群间分化系数；NM：基因流参数
1) NA : Allele number; NE : Effective allele number; I: Shannon index; HO : Observed heterozygosity; HE: Expected heterozygosity; FIS : Inbreeding coefficient within population; FIT : Total inbreeding coefficient; FST: Differentiation coefficient among populations; NM: Gene flow among populations
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表 5 草地贪夜蛾群体遗传多样性指数1)
Table 5 The genetic diversity indexes of Spodoptera frugiperda populations
地理种群 Population NA NE I HO HE F 广东花都 GDHD 8.444±0.784 4.374±0.728 1.594±0.188 0.629±0.085 0.685±0.074 0.087±0.069 湖南张家界 HNZJJ 5.222±0.76 3.467±0.518 1.300±0.166 0.657±0.100 0.640±0.066 −0.028±0.099 广西南宁 GXNN 4.778±0.572 3.412±0.479 1.285±0.145 0.683±0.111 0.651±0.058 −0.033±0.118 广东东莞 GDDG 2.333±0.441 2.071±0.370 0.649±0.183 0.556±0.157 0.389±0.101 −0.426±0.151 广东韶关 GDSG 4.000±0.601 3.126±0.631 1.111±0.174 0.644±0.093 0.578±0.072 −0.113±0.097 平均值 Average 4.407±0.554 3.020±0.482 1.067±10.162 0.640±0.119 0.546±0.076 −0.252±0.089 1) NA: 观测等位基因数; NE: 有效等位基因数; I: Shannon信息指数; HO: 表观杂合度; HE: 预期杂合度; F: 固定指数
1) NA: Allele number; NE: Effective allele number; I: Shannon index; HO: Observed heterozygosity; HE: Expected heterozygosity; F: Fixation index
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表 6 草地贪夜蛾不同地理种群的Nei’ s遗传距离(D)和遗传相似性(I)1)
Table 6 Genetic distance (D) and genetic identity (I) between different geo-populations of Spodoptera frugiperda
种群 Population 花都 Huadu 张家界 Zhangjiajie 南宁 Nanning 东莞 Dongguan 韶关 Shaoguan 花都 Huadu 0.895 0.863 0.831 0.866 张家界 Zhangjiajie 0.111 0.798 0.781 0.814 南宁 Nanning 0.147 0.225 0.670 0.771 东莞 Dongguan 0.185 0.247 0.400 0.740 韶关 Shaoguan 0.144 0.206 0.260 0.301 1)表中对角线以上为遗传相似性(I)，对角线以下为Nei’ s标准遗传距离(D)
1)The genetic identity data(I) are located above the diagonal line，and Nei’ s standard genetic distance(D) below the diagonal line
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