谷蠹不同地理种群对磷化氢抗性及其遗传分化
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摘要
谷蠹分布于世界各地,以热带和亚热带地区发生最重,幼虫和成虫均可在粮粒内为害,是我国主要的储粮害虫之一。目前,控制储粮害虫为害的主要熏蒸剂为磷化氢。由于仓库较差的密封性及多次重复无效熏蒸造成的选择性压力是造成储粮害虫对磷化氢产生抗性的重要原因。本研究主要针对中国储粮区内谷蠹不同地理种群对磷化氢的抗性进行了详细的调查,并对不同抗性的谷蠹种群瞬时增长率(ri)进行了探讨。利用AFLP及mtCOI基因进行了谷蠹种群的遗传变异研究,并讨论了谷蠹体内三种保护酶在磷化氢熏蒸前后的变化。具体研究结果如下：
1.谷蠹不同地理种群对磷化氢的抗性
采用FAO推荐的磷化氢抗性测定方法,测定了全国10个省及1个直辖市的16个谷蠹种群的抗性。试验种群的致死中浓度LC50范围为0.017mg/L-4.272mg/L,16个谷蠹种群均对磷化氢产生了不同程度的抗药性,其中,低抗性种群5个,中等抗性6个,高抗性种群5个。对谷蠹不同地理种群的磷化氢抗性水平研究表明,谷蠹对磷化氢的抗性在中国已经成为一个严重的问题。对谷蠹不同抗性种群的瞬时增长率研究表明,较高抗性的种群具有较低的瞬时增长率。把16个种群按照抗性系数大小进行排序,采集自粮食加工企业的种群聚在最顶端,接着是来自国家粮食储备库的种群,后面就是来自地方粮库的种群,采集自这三种粮库类型的谷蠹种群的平均RF值分别为5.465,61.490和277.643,这个结果说明,谷蠹种群的抗性跟仓库类型有关。
2.谷蠹不同地理种群的遗传变异AFLP分析
采用扩增片段长度多态性技术对这16个种群的110个个体进行种群遗传变异分析,结果表明：采用4对选择性扩增引物,在50-450bp范围内共得到129个多态性位点,在种群水平上和种的水平上的多态性位点的百分率分别为43.95%和84.87%。Nei's遗传多样性指数(h)和Shannon's指数(Ⅰ)与多态性比例具有一致性。这三个指数表明,谷蠹种群内存在高的遗传变异。
分子变异分析(AMOVA)结果表明,16个谷蠹种群的遗传结构存在显著性差异(P<0.0005)。在整个变异中,17.62%的变异发生在谷蠹种群间,82.38%的变异发生在种群内部的个体之间。这个结果说明谷蠹的遗传变异主要发生在种群内部,而不是种群之间。整个谷蠹种群的ΦST值为0.176,基因流(Nm)为1.170。基因流大于1表明在谷蠹种群之间还是有一定的基因交流,这在一定程度上防止了由于遗传漂变引起的种群间的遗传分化。
UPGMA聚类分析表明,谷蠹种群的遗传变异与种群的地理位置不相关。Mantel检测同样证明,谷蠹两两之间的Nei's遗传距离和地理距离(Lg值)之间没有显著的相关性(r=0.1054,P=0.8034>0.05),这也表明IBD (isolation by distance)并不能用于描述谷蠹种群的遗传变异与地理距离之间的关系。对谷蠹种群分布的仓库类型及储粮生态区对种群的遗传变异分析结果表明,仓库类型及储粮生态区并不是影响种群变异的显著因子。
3.谷蠹mtDNA-COI基因序列及种群遗传变异分析
对15个种群的100个个体进行mtCOI部分序列分析,得到691bp长度的片段,三个位点的密码子中,A+T的含量平均值为63.2%,在三个密码子中的含量分别是85.2%,53.3%和57.0%。这种A+T含量的偏向在第一位点较第二和第三位点强。序列中A, T, C, G的含量分别为33.2%,32%,18.7%,16.2%。691bp长度的碱基中,共发现26个多态位点,定义了25种单倍型,其中种群间共享单倍型有7个15个谷蠹种群之间存在一定的基因流(Nm=1.35),但是使用AMOVA分析种群的遗传结构表明,谷蠹种群的变异主要发生在种群内部的个体间。
4.磷化氢对不同地理种群谷蠹成虫体内三种保护酶的影响
对不同地理种群谷蠹成虫体内过氧化氢酶(CAT),超氧化物歧化酶(SOD),过氧化物酶(POD)的活性研究表明：熏蒸前,不同地理种群的谷蠹成虫体内三种保护酶的活性不同；使用磷化氢进行熏蒸处理后,不同地理种群谷蠹体内的三种保护酶活性的变化规律为：SOD活力升高,POD和CAT活性降低。采用SPSS16.0分别对熏蒸前试验种群的致死中浓度(LC50)及三种酶活性两两变量进行相关性分析,结果表明：试验种群的LC50与CAT之间存在显著简单相关性(P=0.031<0.05)；LC50与SOD,POD两种酶活性不存在显著相关性(P=0.545；0.053>0.05)；CAT及POD之间存在简单相关性(P=0.034<0.05)。
Rhyzopertha dominica (Fabricius) is one of the major coleopterous pests of stored products in China and around the world. Both larvae and adults are internal feeders, causing serious damages to grains, such as rice, maize, wheat and other stored commodities. It causes serious losses to stored grains in tropical and subtropical region. Currently, the main method for controlling stored-product insects in many countries, including China, is fumigation with phosphine. However, repeated ineffective fumigation in poorly sealed structures resulting in under-dosing could lead to the development of strong resistance in target pest. This research investigated the phosphine resistacne and the instantaneous growth rate (ri) among different geographical populations of R. dominica in China's grain region. AFLP and mtCOI gene also used to analyze the genetic differentiation among R. dominica population. The activities of catalase (CAT), superoxide dismutase (SOD) and peroxidase (POD) were also observed (CAT, SOD, POD). And the detail results showed as followed:
1. Phosphine resistance in R. dominica in China from different geographical populations in China
The present investigation was carried out to determine the level of phosphine resistance in16R. dominica (Fabricius) populations that were collected from ten provinces and one municipality in China following the Food and Agriculture Organization's (FAO) standard method. Results showed that the50%lethal concentration (LC5o) of phosphine to these R. dominica populations ranged from0.017mg/L to4.272mg/L. Of the16populations,5were of low resistance,6were moderately resistant, and5were high resistant. The present results have revealed the seriousness of the problems of phosphine resistance in China. R. dominica populations with high ri showed low resistance factor (RF for LC50), while low ri presented high RF value. The populations were ranked in order of RF values, the populations from processing facilities cluster near the top, then the national storage populations, then the populations from local storage. The average RF for processing facilities, national storages and local storage are5.465,61.490and277.643, respectively. This result indicated that resistance is connected with the type of grain storage.
2. Genetic diversity among different geographical populations of R. dominica in China as revealed by AFLP
In this study, the genetic diversity and structure of16populations (110individuals) of R. dominica were examined using Amplified Fragment Length Polymorphism (AFLP) analysis. Four primer combinations were selected to identify a total of128polymorphic loci ranging from50to450bp. The percentage of polymorphic loci (Pp) was43.95%at the population level and84.87%at the species level, respectively. Nei's gene diversity (h) and Shannon's Information index (I) had similar trend with the percentage of polymorphism. Thus, the estimated genetic diversity using these three indicators revealed a high level of genetic variation within each R. dominica population. At the species level, Analysis of Molecular Variance (AMOVA) revealed a significantly greater genetic variation within populations (82.38%) than among populations (17.62%)(P<0.0005). This result showed that the genetic diversity mainly occurred within the population, not among populations. The ΦST value of the specied was0.176and the gene flow (Nm) value was1.170at the species level. This indicated that, a certain level of gene flow was among the populations and prevented the genetic differentiation among populations caused by genetic drift.
An Unweighted Pair Group Method with Arithmetic Mean (UPGMA) dendrogram based on Nei's unbiased genetic distance revealed four distinct groups that did not correspond to the geographical regions. The Mantel test of isolation by distance revealed that there was no significant correlation between Nei's genetic distance and geographical distance (Lg) among all populations (r=0.1054, P=0.8034>0.05). This result suggests that isolation by distance does not describe the pattern of gene flow among R. dominica populations. The ecology of grain storage area and the depot/storage facility type were not the main factors determining the genetic diversity of R. dominica.
3. The analysis of mtCOI gene and genetic variation of R. dominica population
The mtCOI gene partial sequences (691bp) of100individual from15R. dominica populations were amplified and sequenced. Nucleotides compositi on, transiti on and transversi on, genetic distance of this segment had been analyzed. The molecular volution characteristic of COI gene in different individuals was discussed. The results indicate that average A+T content of COI gene in R. dominica is63.2%, which shows strong A+T bias. The A+T content was85.2%,53.3%and57.0%respectively in the three points. A+T content of this bias showed stronger in the first than that of the second and third points. A, T, C, G content of33.2%,32%,18.7%,16.2%. Twenty-six nucleoide site were substituted and twenty-five haplotypes were identified in all sequences with seven shared. The Nm value was1.35showed that there was certain gene flow among R. dominica population. AMOVA analysis of genetic structure indicated that the gentic variation of R. dominica mainly occurred among individuals.
4. Effect of phosphine on three protective enzymes in R. dominica (Fabricius) adult in different geographical populations
The activities of catalase (CAT), superoxide dismutase (SOD) and peroxidase (POD) were observed in the R. dominica adult in seven different geographical population. The activities of CAT, POD and SOD showed difference among populations in R. dominica adults before fumigation. The results showed that the activities of CAT and POD in seven different population of adults decreased after fumigation treatment, While the activities of SOD enhanced. The activity of CAT in seven populations had expressed significant simple correlation with the50%lethal concentration (LC50) of R. dominica populations (P=0.031<0.05); Meanwhile, there were no significant correlation between the SOD, POD and the LC50three enzyme activities before fumigation (P=0.545;0.053>0.05), while the CAT activity showed simple correlation with the POD activity (P=0.034<0.05).
1.谷蠹不同地理种群对磷化氢的抗性
采用FAO推荐的磷化氢抗性测定方法,测定了全国10个省及1个直辖市的16个谷蠹种群的抗性。试验种群的致死中浓度LC50范围为0.017mg/L-4.272mg/L,16个谷蠹种群均对磷化氢产生了不同程度的抗药性,其中,低抗性种群5个,中等抗性6个,高抗性种群5个。对谷蠹不同地理种群的磷化氢抗性水平研究表明,谷蠹对磷化氢的抗性在中国已经成为一个严重的问题。对谷蠹不同抗性种群的瞬时增长率研究表明,较高抗性的种群具有较低的瞬时增长率。把16个种群按照抗性系数大小进行排序,采集自粮食加工企业的种群聚在最顶端,接着是来自国家粮食储备库的种群,后面就是来自地方粮库的种群,采集自这三种粮库类型的谷蠹种群的平均RF值分别为5.465,61.490和277.643,这个结果说明,谷蠹种群的抗性跟仓库类型有关。
2.谷蠹不同地理种群的遗传变异AFLP分析
采用扩增片段长度多态性技术对这16个种群的110个个体进行种群遗传变异分析,结果表明：采用4对选择性扩增引物,在50-450bp范围内共得到129个多态性位点,在种群水平上和种的水平上的多态性位点的百分率分别为43.95%和84.87%。Nei's遗传多样性指数(h)和Shannon's指数(Ⅰ)与多态性比例具有一致性。这三个指数表明,谷蠹种群内存在高的遗传变异。
分子变异分析(AMOVA)结果表明,16个谷蠹种群的遗传结构存在显著性差异(P<0.0005)。在整个变异中,17.62%的变异发生在谷蠹种群间,82.38%的变异发生在种群内部的个体之间。这个结果说明谷蠹的遗传变异主要发生在种群内部,而不是种群之间。整个谷蠹种群的ΦST值为0.176,基因流(Nm)为1.170。基因流大于1表明在谷蠹种群之间还是有一定的基因交流,这在一定程度上防止了由于遗传漂变引起的种群间的遗传分化。
UPGMA聚类分析表明,谷蠹种群的遗传变异与种群的地理位置不相关。Mantel检测同样证明,谷蠹两两之间的Nei's遗传距离和地理距离(Lg值)之间没有显著的相关性(r=0.1054,P=0.8034>0.05),这也表明IBD (isolation by distance)并不能用于描述谷蠹种群的遗传变异与地理距离之间的关系。对谷蠹种群分布的仓库类型及储粮生态区对种群的遗传变异分析结果表明,仓库类型及储粮生态区并不是影响种群变异的显著因子。
3.谷蠹mtDNA-COI基因序列及种群遗传变异分析
对15个种群的100个个体进行mtCOI部分序列分析,得到691bp长度的片段,三个位点的密码子中,A+T的含量平均值为63.2%,在三个密码子中的含量分别是85.2%,53.3%和57.0%。这种A+T含量的偏向在第一位点较第二和第三位点强。序列中A, T, C, G的含量分别为33.2%,32%,18.7%,16.2%。691bp长度的碱基中,共发现26个多态位点,定义了25种单倍型,其中种群间共享单倍型有7个15个谷蠹种群之间存在一定的基因流(Nm=1.35),但是使用AMOVA分析种群的遗传结构表明,谷蠹种群的变异主要发生在种群内部的个体间。
4.磷化氢对不同地理种群谷蠹成虫体内三种保护酶的影响
对不同地理种群谷蠹成虫体内过氧化氢酶(CAT),超氧化物歧化酶(SOD),过氧化物酶(POD)的活性研究表明：熏蒸前,不同地理种群的谷蠹成虫体内三种保护酶的活性不同；使用磷化氢进行熏蒸处理后,不同地理种群谷蠹体内的三种保护酶活性的变化规律为：SOD活力升高,POD和CAT活性降低。采用SPSS16.0分别对熏蒸前试验种群的致死中浓度(LC50)及三种酶活性两两变量进行相关性分析,结果表明：试验种群的LC50与CAT之间存在显著简单相关性(P=0.031<0.05)；LC50与SOD,POD两种酶活性不存在显著相关性(P=0.545；0.053>0.05)；CAT及POD之间存在简单相关性(P=0.034<0.05)。
Rhyzopertha dominica (Fabricius) is one of the major coleopterous pests of stored products in China and around the world. Both larvae and adults are internal feeders, causing serious damages to grains, such as rice, maize, wheat and other stored commodities. It causes serious losses to stored grains in tropical and subtropical region. Currently, the main method for controlling stored-product insects in many countries, including China, is fumigation with phosphine. However, repeated ineffective fumigation in poorly sealed structures resulting in under-dosing could lead to the development of strong resistance in target pest. This research investigated the phosphine resistacne and the instantaneous growth rate (ri) among different geographical populations of R. dominica in China's grain region. AFLP and mtCOI gene also used to analyze the genetic differentiation among R. dominica population. The activities of catalase (CAT), superoxide dismutase (SOD) and peroxidase (POD) were also observed (CAT, SOD, POD). And the detail results showed as followed:
1. Phosphine resistance in R. dominica in China from different geographical populations in China
The present investigation was carried out to determine the level of phosphine resistance in16R. dominica (Fabricius) populations that were collected from ten provinces and one municipality in China following the Food and Agriculture Organization's (FAO) standard method. Results showed that the50%lethal concentration (LC5o) of phosphine to these R. dominica populations ranged from0.017mg/L to4.272mg/L. Of the16populations,5were of low resistance,6were moderately resistant, and5were high resistant. The present results have revealed the seriousness of the problems of phosphine resistance in China. R. dominica populations with high ri showed low resistance factor (RF for LC50), while low ri presented high RF value. The populations were ranked in order of RF values, the populations from processing facilities cluster near the top, then the national storage populations, then the populations from local storage. The average RF for processing facilities, national storages and local storage are5.465,61.490and277.643, respectively. This result indicated that resistance is connected with the type of grain storage.
2. Genetic diversity among different geographical populations of R. dominica in China as revealed by AFLP
In this study, the genetic diversity and structure of16populations (110individuals) of R. dominica were examined using Amplified Fragment Length Polymorphism (AFLP) analysis. Four primer combinations were selected to identify a total of128polymorphic loci ranging from50to450bp. The percentage of polymorphic loci (Pp) was43.95%at the population level and84.87%at the species level, respectively. Nei's gene diversity (h) and Shannon's Information index (I) had similar trend with the percentage of polymorphism. Thus, the estimated genetic diversity using these three indicators revealed a high level of genetic variation within each R. dominica population. At the species level, Analysis of Molecular Variance (AMOVA) revealed a significantly greater genetic variation within populations (82.38%) than among populations (17.62%)(P<0.0005). This result showed that the genetic diversity mainly occurred within the population, not among populations. The ΦST value of the specied was0.176and the gene flow (Nm) value was1.170at the species level. This indicated that, a certain level of gene flow was among the populations and prevented the genetic differentiation among populations caused by genetic drift.
An Unweighted Pair Group Method with Arithmetic Mean (UPGMA) dendrogram based on Nei's unbiased genetic distance revealed four distinct groups that did not correspond to the geographical regions. The Mantel test of isolation by distance revealed that there was no significant correlation between Nei's genetic distance and geographical distance (Lg) among all populations (r=0.1054, P=0.8034>0.05). This result suggests that isolation by distance does not describe the pattern of gene flow among R. dominica populations. The ecology of grain storage area and the depot/storage facility type were not the main factors determining the genetic diversity of R. dominica.
3. The analysis of mtCOI gene and genetic variation of R. dominica population
The mtCOI gene partial sequences (691bp) of100individual from15R. dominica populations were amplified and sequenced. Nucleotides compositi on, transiti on and transversi on, genetic distance of this segment had been analyzed. The molecular volution characteristic of COI gene in different individuals was discussed. The results indicate that average A+T content of COI gene in R. dominica is63.2%, which shows strong A+T bias. The A+T content was85.2%,53.3%and57.0%respectively in the three points. A+T content of this bias showed stronger in the first than that of the second and third points. A, T, C, G content of33.2%,32%,18.7%,16.2%. Twenty-six nucleoide site were substituted and twenty-five haplotypes were identified in all sequences with seven shared. The Nm value was1.35showed that there was certain gene flow among R. dominica population. AMOVA analysis of genetic structure indicated that the gentic variation of R. dominica mainly occurred among individuals.
4. Effect of phosphine on three protective enzymes in R. dominica (Fabricius) adult in different geographical populations
The activities of catalase (CAT), superoxide dismutase (SOD) and peroxidase (POD) were observed in the R. dominica adult in seven different geographical population. The activities of CAT, POD and SOD showed difference among populations in R. dominica adults before fumigation. The results showed that the activities of CAT and POD in seven different population of adults decreased after fumigation treatment, While the activities of SOD enhanced. The activity of CAT in seven populations had expressed significant simple correlation with the50%lethal concentration (LC50) of R. dominica populations (P=0.031<0.05); Meanwhile, there were no significant correlation between the SOD, POD and the LC50three enzyme activities before fumigation (P=0.545;0.053>0.05), while the CAT activity showed simple correlation with the POD activity (P=0.034<0.05).
引文
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