福寿螺繁殖生态与遗传多样性研究
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摘要
福寿螺(Pomacea canaliculata),原产于南美洲亚马逊河流域,属热带和亚热带种,是一种大型淡水食用螺类。1981年从台湾引入广东养殖,很快扩展到全国各地,并成为我国南方农作物的主要敌害生物,对水稻生产威胁很大。在我国北纬30°以南的省份均有福寿螺发生危害的报道。其危害程度与温度成正相关,与纬度成负相关,与其繁殖世代成正相关。福寿螺还是人畜共患的广州管圆线虫病传播中最为重要的中间宿主之一,如果生吃或者食用未煮熟的福寿螺螺肉,极易引起食源性“广州管圆线虫病”,引起人体嗜酸性粒细胞增多性脑膜炎。福寿螺给生态环境和经济活动造成了极大的负面影响。
本研究以江苏苏州( 120°62′E, 31°32′N)、福建漳州( 117°35′E, 24°52′N)和广东珠海( 113°52′E, 22°3′N)的福寿螺为材料,研究了不同地理群体、不同繁殖世代、不同壳色的福寿螺的繁殖基础和遗传变异,比较福寿螺的繁殖力、交配选择性、遗传多样性及遗传结构变异等特性,为控制福寿螺对水稻秧苗危害及防止其扩散提供科学依据。主要结果与结论如下:
1、不同地理群体福寿螺的繁殖力比较。应用单对交配的方法以福寿螺的平均个体产卵量、个体产卵块数和雌螺个体体重的相对产卵量为指标分析了不同地理群体、不同壳色福寿螺的繁殖力。结果显示福寿螺的繁殖力变异较大。在个体产卵量方面,福建螺与江苏螺有显著差异(P<0.05);在雌螺个体体重的相对产卵量方面,广东螺与福建螺、广东螺与江苏螺有显著差异(P<0.05);不同地区、不同交配个体产卵块数差异不显著(P>0.05);不同壳色的福寿螺繁殖力差异不明显(P>0.05);交配个体雌雄大小不同其交配产卵量与雌螺个体大小成正相关,与雄螺个体大小无关;雌螺个体越大产卵量越多,但相对产卵量反而越小;雌螺产卵量与卵块数成正相关。福寿螺的繁殖力与环境条件、食物种类、温度等因素有密切关系。
2、不同地理群体福寿螺的交配选择性研究。应用体外标记法对三个地理群体福寿螺的选择性交配情况进行观察,结果表明:广东福寿螺倾向于群体内相互交配,而福建螺与江苏螺倾向于不同地理群体间相互杂交;在福寿螺的交配选择性试验中发现,虽然福寿螺的交配是混交型没有固定配偶,但不同壳色福寿螺间的交配明显高于同壳色福寿螺间的交配(P<0.05);交配个体间雌螺大于雄螺的交配形式占绝大多数(P<0.05)。
3、不同壳色福寿螺的遗传规律分析。应用单对交配方法对江苏养殖群体中出现的不同壳色福寿螺的遗传规律进行研究,结果显示:福寿螺黄黑壳色是一对相对性状,受核基因控制,与性别无关,其遗传规律符合孟德尔单因子分离规律,野生群体调查发现,福寿螺壳色差异自然存在,但不同壳色在不同地区所占比例不同,表现为广东螺黑色为主,云南螺黄色为主,福建螺和江苏螺中黄螺多于黑螺。
4、不同地理群体福寿螺遗传多样性的AFLP分析。利用筛选出的8组引物(E32M54、E42M61、E42M54、E45M62、E32M61、E32M62、E32M59、E39M54)对福建、江苏和广东三个群体的福寿螺进行AFLP扩增,共得到382个清晰位点,多态位点比率为96.60%,不同引物组合产生的扩增位点为35-61个,平均47.75个。福建、江苏和广东三个群体的多态位点比率分别为85.08%、84.82%和79.06%;Nei's基因多样性指数分别为0.2860、0.2819和0.2710; Shannon's多样性信息指数分别为0.4308、0.4259和0.4079;各群体的遗传多样性丰富,其中福建螺遗传多样性最高,广东螺最低。各群体间遗传距离为0.0845-0.1017,其中福建螺与广东螺遗传距离较近(0.0845),广东螺与江苏螺的遗传距离较远(0.1017)。找到了三个群体的特征位点21个,其中福建螺11个,江苏螺6个,广东螺4个。
5、不同壳色福寿螺的AFLP分析。应用AFLP技术对不同壳色福寿螺进行分析,黄螺和黑螺的多态位点比率分别为92.41%和93.72%;Nei's基因多样性指数分别为0.3214和0.3327; Shannon's多样性信息指数分别为0.4819和0.4974;其遗传多样性丰富,黄螺与黑螺相比有7个位点发生缺失。
6、不同性别福寿螺的AFLP分析。应用AFLP技术对不同性别福寿螺进行分析,雌螺和雄螺的多态位点比率分别为94.76%和91.36%;Nei's基因多样性指数分别为0.3402和0.3222; Shannon's多样性信息指数分别为0.5068和0.4816;其遗传多样性丰富,雄螺与雌螺相比有7个位点发生缺失。
7、不同地理群体福寿螺遗传多样性的ISSR分析。利用经筛选的3个ISSR引物对福建、江苏和广东三个群体的福寿螺进行扩增,共得到33个清晰位点,多态位点比率为93.94%。福建、江苏和广东三个群体的多态位点比率分别为.69.70%、63.64%和48.48%;Nei's基因多样性指数分别为0.2940、0.2387和0.1903; Shannon's多样性信息指数分别为0.4247、0.3536和0.2796;各群体的遗传多样性都很丰富,其中福建螺遗传多样性最高,广东螺最低。各群体间遗传距离为0.1020-0.2158,其中福建螺与广东螺遗传距离较近(0.1020),广东螺与江苏螺的遗传距离较远(0.2158)。其趋势与AFLP分析结果一致。
8、不同地理群体福寿螺对水稻秧苗的嗜食性比较。福寿螺在我国南方主要危害的农作物是水稻秧苗,实验表明:福寿螺是杂食性螺类,喜欢摄食幼嫩的绿色水生植物,在有其它幼嫩的绿色水生植物的情况下对水稻秧苗摄食量很少,而对浮水植物满江红和紫背浮萍有较强的摄食选择性,因此,水稻秧苗期在稻田中投放适量的浮水植物(满江红和紫背浮萍等)可以作为防止福寿螺危害水稻秧苗的一种临时措施。
The golden apple snails (Pomacea canaliculata) are big food fresh-water snail, and they live in tropical or Subtropical Region which are found in the Amazon Plain of South America. As a new breed source, the snails were introduced into Taiwan intentionally in 1981 and were transferred from Taiwan to Guangdong, and then it was spread to the national scale, in southern China it has become a serious pest of aquatic plants, rice specially. The report of the snails’damage came from the south of 30°N in China, which showed the harm extent had a positive correlation with temperature and propagation generation, a negative correlation with latitude. It deserves attention greatly because the snail is one of the most important intermediate host through infecting Angiostrongylus cantonensis both domestic animals and human as a cause of eosinophilic meningitis of human. The snails have made huge negative influence on ecological environment and economic activities.
In this study, the snails were collected from Zhangzhou (Fujian)、Zhuhai (Guangdong) and Suzhou (Jiangsu). Compare study on reproductive basis and genetic differentiation, including fecundity, mating selection, genetic diversity and genetic structure differentiation and so on, was carried out for different geographic populations, propagation generation and shell color of the snails. This study eventually provide theoretical basis for control the snails’harm and prevent from its spread. The main result and conclusion are as follows:
1. Compare study on fecundity of different geographic populations of P. canaliculata. The single-pair mating method was used to analyze the snails’fecundity of different geographic and shell-color populations. The results indicated that there was difference to fecundity of different geographic populations, and the difference was significant between Fujian and Jiangsu in individual fecundity, and the difference was also significant between Guangdong and Jiangsu、Guangdong and Fujian in relative fecundity of female. Fecundity has insignificant difference for different shell-color populations of the snails. The spawning number is independent on individual size of the male and is positively related with individual size of the female. The snails’fecundity has close relationship with the factor, including environmental condition, food species and temperature and so on.
2. Study on the analysis for the mating selection of different geographic populations of P. canaliculata. The snails’mating selection of different geographic populations was observed by using the marker method in vitro. The result showed that the snails from Guangdong preferred to mate each other within their population, but the snails from Fujian and Jiangsu preferred to mate among different populations; the finding given by test of the mating selection was the snails’mating belong to mixed type (no fixed mate), the mating rate among different shell-color snail was obviously higher than that of same shell-color; in general case, the female body size is larger than that of the male.
3. The Analysis of the genetic law among different shell-color P. canaliculata. Study on the genetic law of the different color-shell snails by single-pair mating, which is existent in cultivating population in the Jiangsu province. The results showed that the yellow shell and black shell is a pair of relative properties, which is controlled by nuclear gene and have nothing to do with sex. The genetic law accord with Mendel’s single factor separating Law. The research of wild population found that the shell-color difference of the snails was existent in nature, but the proportion taken up in different area was different. The snails’shell from Guangdong was mostly black, that was mainly yellow from Yunnan. Number of the yellow snail was more than that of the black in Fujian and Jiangsu population.
4. Analysis of genetic diversity of different geographic populations of P. canaliculata by AFLP markers. Eight AFLP primer combinations, including E32M54, E42M61, E42M54, E45M62, E32M61, E32M62, E32M59 and E39M54, which were obtained by primer screening produced total 382 bands amplified (the percentage of polymorphic loci is 96.60%) from Fujian, Jiangsu and Guangdong populations, number of amplified loci produced by different primer combinations is 35-61 and their average value is 47.75. The proportion of polymorphic loci, Nei's index and Shannon’s Information index from Fujian, Jiangsu and Guangdong were 85.08%, 84.82% and 79.06%; 0.2860, 0.2819 and 0.2710; 0.4308, 0.4259 and 0.4079, respectively. The three populations of P. canaliculata were in a higher level of genetic diversity, there is the highest genetic diversity within Fujian population and the lowest that within Guangdong population. Genetic distance is 0.0845-0.1017 among three populations, and that is the nearest between Fujian and Guangdong and is the farthest between Guangdong and Jiangsu. 21 specific sites, including 11, 4 and 6 from Fujian, Jiangsu and Guangdong population, were found in this study, respectively.
5. Analysis of different shell-color of P. canaliculata. The proportion of polymorphic loci, Nei's index and Shannon’s Information index from yellow and black snail were 92.41% and 93.72%; 0.3214 and 0.3327; 0.4819 and 0.4974, respectively. Comparing with the black snail, the yellow lose seven sites but their genetic diversity was still abundant.
6. Analysis of t different genders of P. canaliculata. The proportion of polymorphic loci, Nei's index and Shannon’s Information index from female and male snail were 94.76% and 91.36%; 0.3402 and 0.3222; 0.5068 and 0.4816, respectively. Comparing with the female snail, the male lose seven sites but their genetic diversity was still abundant.
7. Study on genetic diversity of different geographic populations of P. canaliculata by ISSR markers. Three primers screened were made the amplification from three populations of P. canalicuata and 33 clear amplified loci were obtained, and their percentage of polymorphic loci is 93.94%. The proportion of polymorphic loci, Nei's index and Shannon’s Information index from Fujian, Jiangsu and Guangdong were 69.70%, 63.64% and 48.48%; 0.2940, 0.2387 and 0.1903; 0.4247, 0.3536 and 0.2796, respectively. There is abundant genetic diversity among three populations, Fujian population has the highest genetic diversity and Guangdong population is the lowest that. Genetic distance is 0.1020-0.2158 among three populations, and its trend is same as AFLP among three populations.
8. Comparison on preference to rice seedling among different geographic populations of P. canaliculata. Mainly crops endangered by snails is the rice seedling in south of china, the experiment indicated that the snails is omnivory, prefer to eat fresh and tender aquatic plants. In four different feeds, the rice seedling is negative selection; Azolla imbricata and Spirodela polyrrhiza, which are floating plants, is positive selection. Therefore, floating plants can be one kind of interim measures preventing rice seedling from the snails’endangering.
本研究以江苏苏州( 120°62′E, 31°32′N)、福建漳州( 117°35′E, 24°52′N)和广东珠海( 113°52′E, 22°3′N)的福寿螺为材料,研究了不同地理群体、不同繁殖世代、不同壳色的福寿螺的繁殖基础和遗传变异,比较福寿螺的繁殖力、交配选择性、遗传多样性及遗传结构变异等特性,为控制福寿螺对水稻秧苗危害及防止其扩散提供科学依据。主要结果与结论如下:
1、不同地理群体福寿螺的繁殖力比较。应用单对交配的方法以福寿螺的平均个体产卵量、个体产卵块数和雌螺个体体重的相对产卵量为指标分析了不同地理群体、不同壳色福寿螺的繁殖力。结果显示福寿螺的繁殖力变异较大。在个体产卵量方面,福建螺与江苏螺有显著差异(P<0.05);在雌螺个体体重的相对产卵量方面,广东螺与福建螺、广东螺与江苏螺有显著差异(P<0.05);不同地区、不同交配个体产卵块数差异不显著(P>0.05);不同壳色的福寿螺繁殖力差异不明显(P>0.05);交配个体雌雄大小不同其交配产卵量与雌螺个体大小成正相关,与雄螺个体大小无关;雌螺个体越大产卵量越多,但相对产卵量反而越小;雌螺产卵量与卵块数成正相关。福寿螺的繁殖力与环境条件、食物种类、温度等因素有密切关系。
2、不同地理群体福寿螺的交配选择性研究。应用体外标记法对三个地理群体福寿螺的选择性交配情况进行观察,结果表明:广东福寿螺倾向于群体内相互交配,而福建螺与江苏螺倾向于不同地理群体间相互杂交;在福寿螺的交配选择性试验中发现,虽然福寿螺的交配是混交型没有固定配偶,但不同壳色福寿螺间的交配明显高于同壳色福寿螺间的交配(P<0.05);交配个体间雌螺大于雄螺的交配形式占绝大多数(P<0.05)。
3、不同壳色福寿螺的遗传规律分析。应用单对交配方法对江苏养殖群体中出现的不同壳色福寿螺的遗传规律进行研究,结果显示:福寿螺黄黑壳色是一对相对性状,受核基因控制,与性别无关,其遗传规律符合孟德尔单因子分离规律,野生群体调查发现,福寿螺壳色差异自然存在,但不同壳色在不同地区所占比例不同,表现为广东螺黑色为主,云南螺黄色为主,福建螺和江苏螺中黄螺多于黑螺。
4、不同地理群体福寿螺遗传多样性的AFLP分析。利用筛选出的8组引物(E32M54、E42M61、E42M54、E45M62、E32M61、E32M62、E32M59、E39M54)对福建、江苏和广东三个群体的福寿螺进行AFLP扩增,共得到382个清晰位点,多态位点比率为96.60%,不同引物组合产生的扩增位点为35-61个,平均47.75个。福建、江苏和广东三个群体的多态位点比率分别为85.08%、84.82%和79.06%;Nei's基因多样性指数分别为0.2860、0.2819和0.2710; Shannon's多样性信息指数分别为0.4308、0.4259和0.4079;各群体的遗传多样性丰富,其中福建螺遗传多样性最高,广东螺最低。各群体间遗传距离为0.0845-0.1017,其中福建螺与广东螺遗传距离较近(0.0845),广东螺与江苏螺的遗传距离较远(0.1017)。找到了三个群体的特征位点21个,其中福建螺11个,江苏螺6个,广东螺4个。
5、不同壳色福寿螺的AFLP分析。应用AFLP技术对不同壳色福寿螺进行分析,黄螺和黑螺的多态位点比率分别为92.41%和93.72%;Nei's基因多样性指数分别为0.3214和0.3327; Shannon's多样性信息指数分别为0.4819和0.4974;其遗传多样性丰富,黄螺与黑螺相比有7个位点发生缺失。
6、不同性别福寿螺的AFLP分析。应用AFLP技术对不同性别福寿螺进行分析,雌螺和雄螺的多态位点比率分别为94.76%和91.36%;Nei's基因多样性指数分别为0.3402和0.3222; Shannon's多样性信息指数分别为0.5068和0.4816;其遗传多样性丰富,雄螺与雌螺相比有7个位点发生缺失。
7、不同地理群体福寿螺遗传多样性的ISSR分析。利用经筛选的3个ISSR引物对福建、江苏和广东三个群体的福寿螺进行扩增,共得到33个清晰位点,多态位点比率为93.94%。福建、江苏和广东三个群体的多态位点比率分别为.69.70%、63.64%和48.48%;Nei's基因多样性指数分别为0.2940、0.2387和0.1903; Shannon's多样性信息指数分别为0.4247、0.3536和0.2796;各群体的遗传多样性都很丰富,其中福建螺遗传多样性最高,广东螺最低。各群体间遗传距离为0.1020-0.2158,其中福建螺与广东螺遗传距离较近(0.1020),广东螺与江苏螺的遗传距离较远(0.2158)。其趋势与AFLP分析结果一致。
8、不同地理群体福寿螺对水稻秧苗的嗜食性比较。福寿螺在我国南方主要危害的农作物是水稻秧苗,实验表明:福寿螺是杂食性螺类,喜欢摄食幼嫩的绿色水生植物,在有其它幼嫩的绿色水生植物的情况下对水稻秧苗摄食量很少,而对浮水植物满江红和紫背浮萍有较强的摄食选择性,因此,水稻秧苗期在稻田中投放适量的浮水植物(满江红和紫背浮萍等)可以作为防止福寿螺危害水稻秧苗的一种临时措施。
The golden apple snails (Pomacea canaliculata) are big food fresh-water snail, and they live in tropical or Subtropical Region which are found in the Amazon Plain of South America. As a new breed source, the snails were introduced into Taiwan intentionally in 1981 and were transferred from Taiwan to Guangdong, and then it was spread to the national scale, in southern China it has become a serious pest of aquatic plants, rice specially. The report of the snails’damage came from the south of 30°N in China, which showed the harm extent had a positive correlation with temperature and propagation generation, a negative correlation with latitude. It deserves attention greatly because the snail is one of the most important intermediate host through infecting Angiostrongylus cantonensis both domestic animals and human as a cause of eosinophilic meningitis of human. The snails have made huge negative influence on ecological environment and economic activities.
In this study, the snails were collected from Zhangzhou (Fujian)、Zhuhai (Guangdong) and Suzhou (Jiangsu). Compare study on reproductive basis and genetic differentiation, including fecundity, mating selection, genetic diversity and genetic structure differentiation and so on, was carried out for different geographic populations, propagation generation and shell color of the snails. This study eventually provide theoretical basis for control the snails’harm and prevent from its spread. The main result and conclusion are as follows:
1. Compare study on fecundity of different geographic populations of P. canaliculata. The single-pair mating method was used to analyze the snails’fecundity of different geographic and shell-color populations. The results indicated that there was difference to fecundity of different geographic populations, and the difference was significant between Fujian and Jiangsu in individual fecundity, and the difference was also significant between Guangdong and Jiangsu、Guangdong and Fujian in relative fecundity of female. Fecundity has insignificant difference for different shell-color populations of the snails. The spawning number is independent on individual size of the male and is positively related with individual size of the female. The snails’fecundity has close relationship with the factor, including environmental condition, food species and temperature and so on.
2. Study on the analysis for the mating selection of different geographic populations of P. canaliculata. The snails’mating selection of different geographic populations was observed by using the marker method in vitro. The result showed that the snails from Guangdong preferred to mate each other within their population, but the snails from Fujian and Jiangsu preferred to mate among different populations; the finding given by test of the mating selection was the snails’mating belong to mixed type (no fixed mate), the mating rate among different shell-color snail was obviously higher than that of same shell-color; in general case, the female body size is larger than that of the male.
3. The Analysis of the genetic law among different shell-color P. canaliculata. Study on the genetic law of the different color-shell snails by single-pair mating, which is existent in cultivating population in the Jiangsu province. The results showed that the yellow shell and black shell is a pair of relative properties, which is controlled by nuclear gene and have nothing to do with sex. The genetic law accord with Mendel’s single factor separating Law. The research of wild population found that the shell-color difference of the snails was existent in nature, but the proportion taken up in different area was different. The snails’shell from Guangdong was mostly black, that was mainly yellow from Yunnan. Number of the yellow snail was more than that of the black in Fujian and Jiangsu population.
4. Analysis of genetic diversity of different geographic populations of P. canaliculata by AFLP markers. Eight AFLP primer combinations, including E32M54, E42M61, E42M54, E45M62, E32M61, E32M62, E32M59 and E39M54, which were obtained by primer screening produced total 382 bands amplified (the percentage of polymorphic loci is 96.60%) from Fujian, Jiangsu and Guangdong populations, number of amplified loci produced by different primer combinations is 35-61 and their average value is 47.75. The proportion of polymorphic loci, Nei's index and Shannon’s Information index from Fujian, Jiangsu and Guangdong were 85.08%, 84.82% and 79.06%; 0.2860, 0.2819 and 0.2710; 0.4308, 0.4259 and 0.4079, respectively. The three populations of P. canaliculata were in a higher level of genetic diversity, there is the highest genetic diversity within Fujian population and the lowest that within Guangdong population. Genetic distance is 0.0845-0.1017 among three populations, and that is the nearest between Fujian and Guangdong and is the farthest between Guangdong and Jiangsu. 21 specific sites, including 11, 4 and 6 from Fujian, Jiangsu and Guangdong population, were found in this study, respectively.
5. Analysis of different shell-color of P. canaliculata. The proportion of polymorphic loci, Nei's index and Shannon’s Information index from yellow and black snail were 92.41% and 93.72%; 0.3214 and 0.3327; 0.4819 and 0.4974, respectively. Comparing with the black snail, the yellow lose seven sites but their genetic diversity was still abundant.
6. Analysis of t different genders of P. canaliculata. The proportion of polymorphic loci, Nei's index and Shannon’s Information index from female and male snail were 94.76% and 91.36%; 0.3402 and 0.3222; 0.5068 and 0.4816, respectively. Comparing with the female snail, the male lose seven sites but their genetic diversity was still abundant.
7. Study on genetic diversity of different geographic populations of P. canaliculata by ISSR markers. Three primers screened were made the amplification from three populations of P. canalicuata and 33 clear amplified loci were obtained, and their percentage of polymorphic loci is 93.94%. The proportion of polymorphic loci, Nei's index and Shannon’s Information index from Fujian, Jiangsu and Guangdong were 69.70%, 63.64% and 48.48%; 0.2940, 0.2387 and 0.1903; 0.4247, 0.3536 and 0.2796, respectively. There is abundant genetic diversity among three populations, Fujian population has the highest genetic diversity and Guangdong population is the lowest that. Genetic distance is 0.1020-0.2158 among three populations, and its trend is same as AFLP among three populations.
8. Comparison on preference to rice seedling among different geographic populations of P. canaliculata. Mainly crops endangered by snails is the rice seedling in south of china, the experiment indicated that the snails is omnivory, prefer to eat fresh and tender aquatic plants. In four different feeds, the rice seedling is negative selection; Azolla imbricata and Spirodela polyrrhiza, which are floating plants, is positive selection. Therefore, floating plants can be one kind of interim measures preventing rice seedling from the snails’endangering.
引文
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