两近缘种铃夜蛾与烟草互作机制的比较研究
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摘要
可同域发生的棉铃虫Helicoverpa armigera(H(u丨¨)bner)和烟夜蛾H.assulta(Guenee)是铃夜蛾属两近缘种昆虫,二者具有相似的生物学和生态学习性,甚至杂交可育,但它们的寄主范围却截然不同,棉铃虫是多食性昆虫,烟夜蛾则是典型的寡食性昆虫。在主要农作物中,棉花和番茄上极少发现烟夜蛾,而辣椒上几乎没有棉铃虫,只有烟草均为二者所嗜食,也唯独在烟草上它们可以稳定共存。本文从行为、生理和生态适应三个方面,对棉铃虫和烟夜蛾与烟草间的互作进行了比较研究,主要结果如下:
成虫对烟草挥发物的EAG反应:(1)两种夜蛾对测试的20种烟草挥发物均可产生EAG反应;低浓度下对绿叶气味、脂肪族、芳香族化合物的EAG反应显著高于单萜类、倍半萜类和杂环化合物;高浓度下的EAG反应也表现出相似的总体反应趋势,但多数反应值间的差异未达显著水平。(2)低浓度下,烟夜蛾对大多数烟草挥发物的EAG反应显著大于棉铃虫;而高浓度下,两种夜蛾的EAG反应却无显著的种间差异。(3)低浓度和高浓度下,两种夜蛾对多数烟草挥发物的EAG反应均无显著的种内性别间差异。(4)在1.0×10~(-5)mol·L~(-1)～1.0×10~(-1)mol·L~(-1)范围内,两种夜蛾的EAG反应与烟草挥发物的浓度呈正相关,且均未达到饱和。
成虫对烟草挥发物的行为反应:(1)两种夜蛾对绿叶气味、脂肪族、芳香族化合物的相对反应率,显著高于单萜类、倍半萜类和杂环化合物。(2)烟夜蛾交配雌蛾对单萜类[(α-蒎烯、(-)-里那醇]和杂环化合物(糠醛)的相对反应率显著高于棉铃虫;棉铃虫交配雌蛾对倍半萜类化合物(β-水芹烯、桧烯)的相对反应率显著高于烟夜蛾。除此之外,两种夜蛾的行为反应既无显著的种间差异,也无显著的种内性别间差异。
烟夜蛾3种气味结合蛋白基因的时空表达:(1)烟夜蛾Hass-PBP、Hass-GOBP_1、Hass-GOBP_2基因在卵期、幼虫期、蛹期均不表达,只在成虫期表达,且表达量与成虫的日龄无关。(2)上述基因在成虫触角上特异性表达,在成虫的喙、去掉触角的头、胸、腹、足、翅等部位均不表达,其中Hass-PBP基因只在雄蛾触角上表达,而Hass-GOBP_1和Hass-GOBP_2基因在雌雄蛾触角上均表达。
两种夜蛾GOBP_1、GOBP_2基因相对表达量的定量分析:棉铃虫和烟夜蛾成虫触角上GOBP_1和GOBP_2基因的相对表达量,既无显著的种间差异,也无显著的种内性别间差异。
幼虫在烟草上的取食行为:(1)两种夜蛾的初孵幼虫对普通烟草和黄花烟草无取食选择性,但随着虫龄的增加,二者在普通烟草上的出现百分率均显著大于黄花烟草,表现出很强的选择性。(2)在黄花烟草上,两种夜蛾5龄幼虫的取食时间、活动时间、休息时间、取食叶面积比率均无显著差异;但是在普通烟草上,烟夜蛾5龄幼虫的取食时间和取食叶面积比率却显著高于棉铃虫。
幼虫取食烟草后的营养效应:(1)棉铃虫6龄幼虫取食两种烟草后的近似消化率及其相对生长率无显著差异,对黄花烟草的利用率和转化率显著降低,但相对取食量却显著增加。(2)烟夜蛾6龄幼虫取食两种烟草后的相对取食量和近似消化率无显著差异,但是对黄花烟草的利用率、转化率及其相对生长率均显著下降。(3)两种夜蛾6龄幼虫对烟草的近似消化率无显著的种间差异,但烟夜蛾对烟草的相对取食量、利用率和转化率却显著高于棉铃虫,从而使其具有较高的相对生长率。
烟草对两种夜蛾实验种群增长的影响:(1)两种烟草上,棉铃虫的发育历期、存活率、产卵量等均存在显著差异,主要表现为黄花烟草延长了幼虫的发育历期,降低了1龄、2龄幼虫的存活率和成虫的产卵量,导致该种群较低的内禀增长率(r_m=0.0951)、较小的净增殖率(R_0=30.5374)和周限增长率(λ=1.0998)。(2)烟夜蛾在两种烟草上的生长发育、存活和繁殖情况与棉铃虫类似,也表现为黄花烟草延长了幼虫的发育历期,降低了1～3龄幼虫的存活率和成虫的产卵量,导致该种群的内禀增长率(r_m=0.0848)、净增殖率(R_0=16.1085)和周限增长率(λ=1.0885)均小于普通烟草上。(3)两种烟草上,烟夜蛾实验种群的生长发育和存活情况比棉铃虫好,但成虫的雌性比和产卵量却显著降低,致使种群的增长速率小于棉铃虫。
烟草对两种夜蛾自然种群增长的影响:(1)第二代棉铃虫在普通烟田的种群增长(I=1.9922＞1)比黄花烟田(I=1.1581＞1)快;第三代棉铃虫在普通烟田的种群增长有所减慢但仍呈上升趋势(I=1.5994＞1),在黄花烟田却呈下降趋势(I=0.6435＜1)。(2)第二代烟夜蛾在普通烟田的种群数量呈上升趋势(I=1.0325＞1),在黄花烟田呈下降趋势(I=0.6577＜1);第三代烟夜蛾的种群数量在两种烟田内均呈下降趋势,但黄花烟田(I=0.3592＜1)的种群下降速率大于普通烟田(I=0.4571＜1)。(3)两种烟田内,第二代和第三代棉铃虫自然种群的增长速率均大于烟夜蛾。(4)生物因子对两种夜蛾自然种群增长有重要的控制作用,“捕食性天敌”、“寄生性天敌”、“烟草和其他”主要对低龄幼虫起控制作用,而“病原微生物”对整个幼虫期均有一定的控制作用。
两种夜蛾在烟田内的种群消长动态:(1)普通烟田内,第二代和第三代棉铃虫的发生高峰期分别为6月24日、8月7日,盛期持续时间分别为7.09天、9.71天。烟夜蛾的发生高峰期分别比棉铃虫晚17天和13天;第二代烟夜蛾的盛期持续时间与棉铃虫基本一致,而第三代比棉铃虫短3.78天。(2)黄花烟田内,第二代和第三代棉铃虫的发生高峰期分别为6月23日、8月8日,盛期持续时间分别为6.78天、10.36天。烟夜蛾的发生高峰期分别比棉铃虫晚18天和15天;第二代烟夜蛾的盛期持续时间与棉铃虫基本一致,而第三代比棉铃虫短4.69天。(3)棉铃虫和烟夜蛾可通过时间生态位的分离,有效避免对烟草的种间资源竞争。
总之,两种夜蛾对烟草挥发物具有相似的嗅觉感受和行为反应,均能通过烟草挥发物准确找到烟草,从而构成了二者在烟草上共存的行为基础;两种夜蛾均能取食烟草,并能以烟草为食完成各自的生命周期,从而构成了二者在烟草上共存的生理基础;两种夜蛾的实验种群和自然种群均能在烟草上保持连续世代的种群繁衍,并可通过时间生态位的分离,有效避免对烟草的种间资源竞争,从而构成了二者在烟草上共存的生态基础。
Two sibling species Helicoverpa armigera(H(u|¨)bner) and H.assulta(Guenee) are serious crop pests in China.They have the similar morphological,biological and ecological characterstics,as well as they can even hybridize with each other.Their host plant ranges, however,are quite different.The former is a typical polyhpagous species,and its host plant range including at least more than 60 crop species such as cotton,corn,wheat,soybean, tobacco,and tomato,and 67 wild plant species from about 30 plant families including Malvaceae,Solanaceae,Gramineae,Leguminosae,etc.The latter is an oligophagous species with a narrower host plant range,and only feeds on plant species in Solanaceae,such as tobacco and hot pepper.In crops,H.assulta was rarely found in cotton and tomato field,and there was almost no H.armigera existing on hot pepper,only tobacco was the host plant on which both of them preferred and coexisted.In this paper,the behavioral,physiological, ecological adaption mechanism of H.armigera and H.assulta to their common host plant tobacco were studied comparatively,and the main results were summarized as follows:
Electroantennogram(EAG) response of adult to tobacco volatiles:(1) H.armigera and H.assulta elicited similar EAG responses to the tested 20 tobacco volatiles;both of them elicited significantly more sensitive to green leaf volatiles,aliphatic and aromatic compounds, than to monoterpene,sesquiterpene,heterocyclic compounds under lower concentration; while the similar EAG responses under higher concentration were detected,but most of them were not significantly different with each other.(2) H.assulta elicited higher EAG responses to the most tested tobacco volatiles than H.armigera under lower concentration;however, there were no significant interspecfic differences of the two species under higher concentration.(3) There were no significant sexual differences of the two species to most of the tested tobacco volatiles under both lower and higher concentration,respectively.(4) The EAG responses of both species were positively dose-dependent to tobacco volatiles under the tested concentration range,and far from their maximum responses.
Behavioral response of adult to tobacco volatiles:(1) The results of behavioral response suggested that the two species elicited significantly higher relative behavioral response to green leaf volatiles,aliphatic and aromatic compounds,than to monoterpenes, sesquiterpenes and heterocyclic compounds.(2) The relative behavioral response of mated H. assulta females to monoterpenes[α-pinene and(-)-linalool]and heterocyclic compounds (furfural) was much higher than that of H.armigera,while the response to sesquiterpenes (β-phellandrene and sabinene hydrate) was quite in the opposite,in addition,there were no significant interspecific differences of the two species.
The expression of three odorant binding protein genes at different developmental stages and tissues in H.assulta:(1) Hass-PBP、Hass-GOBP_1、Hass-GOBP_2 genes were only expressed in adult stage,but not in the stages of egg,larva and pupa;and the expression quantity was irrelevant to the day-old of adult.(2) Hass-PBP、Hass-GOBP-1、Hass-GOBP_2 genes were only expressed in the antennae of adult,but not in the other parts of body,which including proboscises,head without antennae,thorax,abdomen,leg,wing;besides,Hass-PBP gene was only expressed in the antennae of male adult,while Hass-GOBP_1、Hass-GOBP_2 genes were expressed in the antennae both of male and female adult.
Quantitative analysis of relative expression of GOBP_1 and GOBP_2 genes:there were no significant interspecfic and sexual differences of the relative expression quantity of GOBP_1 and GOBP_2 genes in the antenna of H.armigera and H.assulta adult.
Feeding behaveor of larvae reared on two tobacco species:(1) 1~(st) instar of H. armigera and H.assulta larvae searched for food randomly,and did not showed any selectivity to plant leaves during the tested time scale;however,along with larvae growing up, they exhibited stronger selectivity to the tested tobacco species,and the percentage of appearance of H.armigera and H.assulta larvae on the leaves of Nicotiania tabacum were significantly higher than N.rustica.(2) Feeding time and area of H.assulta on N.tabacum leaves were significantly higher than H.armigera,but on N.rustica leaves,there were no significant interspecfic differences between H.armigera and H.assulta.
Nutritional function of larvae reared on two tobacco species:(1) There were no significant differences in the approximate digestibility(AD) and relative growth rate(RGR) of 6~(th) instar H.armigera larvae reared on the two tobacco species;compared to the larvae feeding on N.tabacum,both the efficiency conversation of ingestion(ECI) and digestion (ECD) to N.rustica significantly decreased,while the relative consumption rate(RCR) increased significantly.(2) There were no significant differences in the relative consumption rate and approximate digestibility of 6~(th) instar H.assulta larvae reared on two tobacco species; compared to the larvae feeding on N.tabacum,the relative growth rate,efficiency conversation of ingestion and digestion of larvae on N.rustica significantly decreased.(3) There were no significant differences in the approximate digestibility between H.armigera and H.assulta,but the relative consumption rate and growth rate,as well as the efficiency conversation of ingestion and digestion of H.assulta reared on the two tobacco species were significantly higher than H.armigera.
The effects of tobacco on the experimental population increase of H.armigera and H.assulta:(1) The development duration of H.armigera larvae were significantly prolonged when reared on N.rustica,while the survival rate of both 1~(st) and 2~(nd) instar larvae,as well as the fecundity of female adults reduced notably,so the net reproductive rate(R_0=30.5374) and innate capacity of increase(r_m=0.0951) decreased compared with those reared on N.tabacum. (2) When reared on N.rustica,the development duration of H.assulta larvae significantly prolonged,while the survival rate of each stage of larvae from 1~(st) to 3~(rd) instar and the fecundity of female adults reduced notablely,and the net reproductive rate(R_0=16.1086) and innate capacity of increase(r_m=0.0848) decreased.(3) The development and survivorship of H. assulta reared on two tobacco species were much better than H.armigera,but the ratio and fecundity of female in the H.assulta population were significantly decreased,leading to the growth rate of population lower than H.armigera.
The effects of tobacco on the natural population increase of H.armigera and H. assulta:(1) The 2n~(nd) generation population of H.armigera in N.tabacum field(I=1.9922＞1) increased faster than that in N.rustica field;while the increase of the 3~(rd) generation population slowed down in N.tabacum field(I=1.5994＞1),and declined in yellow flower tobacco field(I=0.6435＜1).(2) The 2~(nd) generation population of H.assulta in N.tabacum tobacco field(I=1.9922＞1) increased,but declined in N.rustica tobacco field(I=0.6577＜1), the increase of the 3~(rd) generation population slowed down in both two tobacco fields,while the population decreasing rate was faster in N.rustica field(I=0.3592＜1)than in N.tabacum field(I=0.4571＜1).(3) The 2~(nd) and 3~(rd) generation of H.armigera increased faster than H. assulta in both N.tabacum and N.rustica tobacco field.(4) Biotic factors were important to control the population quantities of H.armigera and H.assulta;the early stage larvae were mainly affected by the factors of "predators","parasitoids","tobacco and others";while "pathogenic microorganism" had the control effects on the whole larval stage.
Occurrence dynamics of population quantities of H.armigera and H.assuita in two tobacco fields:(1) The peak of occurrence with 2~(nd) and 3~(rd) generation of H.armigera in N. tabacum field was June 24 and August 7,respectively;while the duration of high occurrence period of 3~(rd) generation was 2.62 days longer than 2~(nd) generation.The peak of occurrence with 2~(nd) and 3~(rd) generation of H.assulta in N.tabacum field was seventeen and thirteen days later than H.armigera,respectively;the duration of high occurrence period of 2~(nd) generation of H.assulta was similar to H.armigera,but 3~(rd) generation of H.assulta was 3.78 days shorter than H.armigera.(2) The peak of occurrence with 2~(nd) and 3~(rd) generation of H. armigera in N.rustica tobacco field were much similar to the N.tabacum field,the duration of high occurrence period of 3~(rd) generation was 3.85 days longer than 2~(nd) generation;the peak of occurrence with 2~(nd) and 3~(rd) generation of H.assulta in N.tabacum field was eighteen and fifteen days later than H.armigera,respectively;the duration of high occurrence period of 2~(nd) generation of H.assulta was similar to H.armigera,but 3~(rd) generation of H.assulta was 4.69 days shorter than H.armigera.(3) H.armigera and H.assulta could through the separation of temporal niche to effectively avoid the interspecific competition in tobacco feld.
In conclusion,the tested insects exhibited the similar olfactory sensitivity and behavioral response to tobacco volatiles between H.armigera and H.assulta,both of them could find tobacco plant accurately through tobacco volatiles,which constituting the behavioral foundations of H.armigera and H.assulta coexisted in tobacco;both of the two Helicoverpa species could feed on tobacco,and they could also complete their life cycles on tobacco, respectively,which constituting the physiological foundations of H.armigera and H.assulta coexisted in tobacco;both of the two Helicoverpa species could keep population increasing with continuous generations in tobacco,and they could effectively avoided interspecific competition to tobacco resources through the separation of temporal niche,which constituting the ecological foundations of H.armigera and H.assulta coexisted in tobacco.
成虫对烟草挥发物的EAG反应:(1)两种夜蛾对测试的20种烟草挥发物均可产生EAG反应;低浓度下对绿叶气味、脂肪族、芳香族化合物的EAG反应显著高于单萜类、倍半萜类和杂环化合物;高浓度下的EAG反应也表现出相似的总体反应趋势,但多数反应值间的差异未达显著水平。(2)低浓度下,烟夜蛾对大多数烟草挥发物的EAG反应显著大于棉铃虫;而高浓度下,两种夜蛾的EAG反应却无显著的种间差异。(3)低浓度和高浓度下,两种夜蛾对多数烟草挥发物的EAG反应均无显著的种内性别间差异。(4)在1.0×10~(-5)mol·L~(-1)～1.0×10~(-1)mol·L~(-1)范围内,两种夜蛾的EAG反应与烟草挥发物的浓度呈正相关,且均未达到饱和。
成虫对烟草挥发物的行为反应:(1)两种夜蛾对绿叶气味、脂肪族、芳香族化合物的相对反应率,显著高于单萜类、倍半萜类和杂环化合物。(2)烟夜蛾交配雌蛾对单萜类[(α-蒎烯、(-)-里那醇]和杂环化合物(糠醛)的相对反应率显著高于棉铃虫;棉铃虫交配雌蛾对倍半萜类化合物(β-水芹烯、桧烯)的相对反应率显著高于烟夜蛾。除此之外,两种夜蛾的行为反应既无显著的种间差异,也无显著的种内性别间差异。
烟夜蛾3种气味结合蛋白基因的时空表达:(1)烟夜蛾Hass-PBP、Hass-GOBP_1、Hass-GOBP_2基因在卵期、幼虫期、蛹期均不表达,只在成虫期表达,且表达量与成虫的日龄无关。(2)上述基因在成虫触角上特异性表达,在成虫的喙、去掉触角的头、胸、腹、足、翅等部位均不表达,其中Hass-PBP基因只在雄蛾触角上表达,而Hass-GOBP_1和Hass-GOBP_2基因在雌雄蛾触角上均表达。
两种夜蛾GOBP_1、GOBP_2基因相对表达量的定量分析:棉铃虫和烟夜蛾成虫触角上GOBP_1和GOBP_2基因的相对表达量,既无显著的种间差异,也无显著的种内性别间差异。
幼虫在烟草上的取食行为:(1)两种夜蛾的初孵幼虫对普通烟草和黄花烟草无取食选择性,但随着虫龄的增加,二者在普通烟草上的出现百分率均显著大于黄花烟草,表现出很强的选择性。(2)在黄花烟草上,两种夜蛾5龄幼虫的取食时间、活动时间、休息时间、取食叶面积比率均无显著差异;但是在普通烟草上,烟夜蛾5龄幼虫的取食时间和取食叶面积比率却显著高于棉铃虫。
幼虫取食烟草后的营养效应:(1)棉铃虫6龄幼虫取食两种烟草后的近似消化率及其相对生长率无显著差异,对黄花烟草的利用率和转化率显著降低,但相对取食量却显著增加。(2)烟夜蛾6龄幼虫取食两种烟草后的相对取食量和近似消化率无显著差异,但是对黄花烟草的利用率、转化率及其相对生长率均显著下降。(3)两种夜蛾6龄幼虫对烟草的近似消化率无显著的种间差异,但烟夜蛾对烟草的相对取食量、利用率和转化率却显著高于棉铃虫,从而使其具有较高的相对生长率。
烟草对两种夜蛾实验种群增长的影响:(1)两种烟草上,棉铃虫的发育历期、存活率、产卵量等均存在显著差异,主要表现为黄花烟草延长了幼虫的发育历期,降低了1龄、2龄幼虫的存活率和成虫的产卵量,导致该种群较低的内禀增长率(r_m=0.0951)、较小的净增殖率(R_0=30.5374)和周限增长率(λ=1.0998)。(2)烟夜蛾在两种烟草上的生长发育、存活和繁殖情况与棉铃虫类似,也表现为黄花烟草延长了幼虫的发育历期,降低了1～3龄幼虫的存活率和成虫的产卵量,导致该种群的内禀增长率(r_m=0.0848)、净增殖率(R_0=16.1085)和周限增长率(λ=1.0885)均小于普通烟草上。(3)两种烟草上,烟夜蛾实验种群的生长发育和存活情况比棉铃虫好,但成虫的雌性比和产卵量却显著降低,致使种群的增长速率小于棉铃虫。
烟草对两种夜蛾自然种群增长的影响:(1)第二代棉铃虫在普通烟田的种群增长(I=1.9922＞1)比黄花烟田(I=1.1581＞1)快;第三代棉铃虫在普通烟田的种群增长有所减慢但仍呈上升趋势(I=1.5994＞1),在黄花烟田却呈下降趋势(I=0.6435＜1)。(2)第二代烟夜蛾在普通烟田的种群数量呈上升趋势(I=1.0325＞1),在黄花烟田呈下降趋势(I=0.6577＜1);第三代烟夜蛾的种群数量在两种烟田内均呈下降趋势,但黄花烟田(I=0.3592＜1)的种群下降速率大于普通烟田(I=0.4571＜1)。(3)两种烟田内,第二代和第三代棉铃虫自然种群的增长速率均大于烟夜蛾。(4)生物因子对两种夜蛾自然种群增长有重要的控制作用,“捕食性天敌”、“寄生性天敌”、“烟草和其他”主要对低龄幼虫起控制作用,而“病原微生物”对整个幼虫期均有一定的控制作用。
两种夜蛾在烟田内的种群消长动态:(1)普通烟田内,第二代和第三代棉铃虫的发生高峰期分别为6月24日、8月7日,盛期持续时间分别为7.09天、9.71天。烟夜蛾的发生高峰期分别比棉铃虫晚17天和13天;第二代烟夜蛾的盛期持续时间与棉铃虫基本一致,而第三代比棉铃虫短3.78天。(2)黄花烟田内,第二代和第三代棉铃虫的发生高峰期分别为6月23日、8月8日,盛期持续时间分别为6.78天、10.36天。烟夜蛾的发生高峰期分别比棉铃虫晚18天和15天;第二代烟夜蛾的盛期持续时间与棉铃虫基本一致,而第三代比棉铃虫短4.69天。(3)棉铃虫和烟夜蛾可通过时间生态位的分离,有效避免对烟草的种间资源竞争。
总之,两种夜蛾对烟草挥发物具有相似的嗅觉感受和行为反应,均能通过烟草挥发物准确找到烟草,从而构成了二者在烟草上共存的行为基础;两种夜蛾均能取食烟草,并能以烟草为食完成各自的生命周期,从而构成了二者在烟草上共存的生理基础;两种夜蛾的实验种群和自然种群均能在烟草上保持连续世代的种群繁衍,并可通过时间生态位的分离,有效避免对烟草的种间资源竞争,从而构成了二者在烟草上共存的生态基础。
Two sibling species Helicoverpa armigera(H(u|¨)bner) and H.assulta(Guenee) are serious crop pests in China.They have the similar morphological,biological and ecological characterstics,as well as they can even hybridize with each other.Their host plant ranges, however,are quite different.The former is a typical polyhpagous species,and its host plant range including at least more than 60 crop species such as cotton,corn,wheat,soybean, tobacco,and tomato,and 67 wild plant species from about 30 plant families including Malvaceae,Solanaceae,Gramineae,Leguminosae,etc.The latter is an oligophagous species with a narrower host plant range,and only feeds on plant species in Solanaceae,such as tobacco and hot pepper.In crops,H.assulta was rarely found in cotton and tomato field,and there was almost no H.armigera existing on hot pepper,only tobacco was the host plant on which both of them preferred and coexisted.In this paper,the behavioral,physiological, ecological adaption mechanism of H.armigera and H.assulta to their common host plant tobacco were studied comparatively,and the main results were summarized as follows:
Electroantennogram(EAG) response of adult to tobacco volatiles:(1) H.armigera and H.assulta elicited similar EAG responses to the tested 20 tobacco volatiles;both of them elicited significantly more sensitive to green leaf volatiles,aliphatic and aromatic compounds, than to monoterpene,sesquiterpene,heterocyclic compounds under lower concentration; while the similar EAG responses under higher concentration were detected,but most of them were not significantly different with each other.(2) H.assulta elicited higher EAG responses to the most tested tobacco volatiles than H.armigera under lower concentration;however, there were no significant interspecfic differences of the two species under higher concentration.(3) There were no significant sexual differences of the two species to most of the tested tobacco volatiles under both lower and higher concentration,respectively.(4) The EAG responses of both species were positively dose-dependent to tobacco volatiles under the tested concentration range,and far from their maximum responses.
Behavioral response of adult to tobacco volatiles:(1) The results of behavioral response suggested that the two species elicited significantly higher relative behavioral response to green leaf volatiles,aliphatic and aromatic compounds,than to monoterpenes, sesquiterpenes and heterocyclic compounds.(2) The relative behavioral response of mated H. assulta females to monoterpenes[α-pinene and(-)-linalool]and heterocyclic compounds (furfural) was much higher than that of H.armigera,while the response to sesquiterpenes (β-phellandrene and sabinene hydrate) was quite in the opposite,in addition,there were no significant interspecific differences of the two species.
The expression of three odorant binding protein genes at different developmental stages and tissues in H.assulta:(1) Hass-PBP、Hass-GOBP_1、Hass-GOBP_2 genes were only expressed in adult stage,but not in the stages of egg,larva and pupa;and the expression quantity was irrelevant to the day-old of adult.(2) Hass-PBP、Hass-GOBP-1、Hass-GOBP_2 genes were only expressed in the antennae of adult,but not in the other parts of body,which including proboscises,head without antennae,thorax,abdomen,leg,wing;besides,Hass-PBP gene was only expressed in the antennae of male adult,while Hass-GOBP_1、Hass-GOBP_2 genes were expressed in the antennae both of male and female adult.
Quantitative analysis of relative expression of GOBP_1 and GOBP_2 genes:there were no significant interspecfic and sexual differences of the relative expression quantity of GOBP_1 and GOBP_2 genes in the antenna of H.armigera and H.assulta adult.
Feeding behaveor of larvae reared on two tobacco species:(1) 1~(st) instar of H. armigera and H.assulta larvae searched for food randomly,and did not showed any selectivity to plant leaves during the tested time scale;however,along with larvae growing up, they exhibited stronger selectivity to the tested tobacco species,and the percentage of appearance of H.armigera and H.assulta larvae on the leaves of Nicotiania tabacum were significantly higher than N.rustica.(2) Feeding time and area of H.assulta on N.tabacum leaves were significantly higher than H.armigera,but on N.rustica leaves,there were no significant interspecfic differences between H.armigera and H.assulta.
Nutritional function of larvae reared on two tobacco species:(1) There were no significant differences in the approximate digestibility(AD) and relative growth rate(RGR) of 6~(th) instar H.armigera larvae reared on the two tobacco species;compared to the larvae feeding on N.tabacum,both the efficiency conversation of ingestion(ECI) and digestion (ECD) to N.rustica significantly decreased,while the relative consumption rate(RCR) increased significantly.(2) There were no significant differences in the relative consumption rate and approximate digestibility of 6~(th) instar H.assulta larvae reared on two tobacco species; compared to the larvae feeding on N.tabacum,the relative growth rate,efficiency conversation of ingestion and digestion of larvae on N.rustica significantly decreased.(3) There were no significant differences in the approximate digestibility between H.armigera and H.assulta,but the relative consumption rate and growth rate,as well as the efficiency conversation of ingestion and digestion of H.assulta reared on the two tobacco species were significantly higher than H.armigera.
The effects of tobacco on the experimental population increase of H.armigera and H.assulta:(1) The development duration of H.armigera larvae were significantly prolonged when reared on N.rustica,while the survival rate of both 1~(st) and 2~(nd) instar larvae,as well as the fecundity of female adults reduced notably,so the net reproductive rate(R_0=30.5374) and innate capacity of increase(r_m=0.0951) decreased compared with those reared on N.tabacum. (2) When reared on N.rustica,the development duration of H.assulta larvae significantly prolonged,while the survival rate of each stage of larvae from 1~(st) to 3~(rd) instar and the fecundity of female adults reduced notablely,and the net reproductive rate(R_0=16.1086) and innate capacity of increase(r_m=0.0848) decreased.(3) The development and survivorship of H. assulta reared on two tobacco species were much better than H.armigera,but the ratio and fecundity of female in the H.assulta population were significantly decreased,leading to the growth rate of population lower than H.armigera.
The effects of tobacco on the natural population increase of H.armigera and H. assulta:(1) The 2n~(nd) generation population of H.armigera in N.tabacum field(I=1.9922＞1) increased faster than that in N.rustica field;while the increase of the 3~(rd) generation population slowed down in N.tabacum field(I=1.5994＞1),and declined in yellow flower tobacco field(I=0.6435＜1).(2) The 2~(nd) generation population of H.assulta in N.tabacum tobacco field(I=1.9922＞1) increased,but declined in N.rustica tobacco field(I=0.6577＜1), the increase of the 3~(rd) generation population slowed down in both two tobacco fields,while the population decreasing rate was faster in N.rustica field(I=0.3592＜1)than in N.tabacum field(I=0.4571＜1).(3) The 2~(nd) and 3~(rd) generation of H.armigera increased faster than H. assulta in both N.tabacum and N.rustica tobacco field.(4) Biotic factors were important to control the population quantities of H.armigera and H.assulta;the early stage larvae were mainly affected by the factors of "predators","parasitoids","tobacco and others";while "pathogenic microorganism" had the control effects on the whole larval stage.
Occurrence dynamics of population quantities of H.armigera and H.assuita in two tobacco fields:(1) The peak of occurrence with 2~(nd) and 3~(rd) generation of H.armigera in N. tabacum field was June 24 and August 7,respectively;while the duration of high occurrence period of 3~(rd) generation was 2.62 days longer than 2~(nd) generation.The peak of occurrence with 2~(nd) and 3~(rd) generation of H.assulta in N.tabacum field was seventeen and thirteen days later than H.armigera,respectively;the duration of high occurrence period of 2~(nd) generation of H.assulta was similar to H.armigera,but 3~(rd) generation of H.assulta was 3.78 days shorter than H.armigera.(2) The peak of occurrence with 2~(nd) and 3~(rd) generation of H. armigera in N.rustica tobacco field were much similar to the N.tabacum field,the duration of high occurrence period of 3~(rd) generation was 3.85 days longer than 2~(nd) generation;the peak of occurrence with 2~(nd) and 3~(rd) generation of H.assulta in N.tabacum field was eighteen and fifteen days later than H.armigera,respectively;the duration of high occurrence period of 2~(nd) generation of H.assulta was similar to H.armigera,but 3~(rd) generation of H.assulta was 4.69 days shorter than H.armigera.(3) H.armigera and H.assulta could through the separation of temporal niche to effectively avoid the interspecific competition in tobacco feld.
In conclusion,the tested insects exhibited the similar olfactory sensitivity and behavioral response to tobacco volatiles between H.armigera and H.assulta,both of them could find tobacco plant accurately through tobacco volatiles,which constituting the behavioral foundations of H.armigera and H.assulta coexisted in tobacco;both of the two Helicoverpa species could feed on tobacco,and they could also complete their life cycles on tobacco, respectively,which constituting the physiological foundations of H.armigera and H.assulta coexisted in tobacco;both of the two Helicoverpa species could keep population increasing with continuous generations in tobacco,and they could effectively avoided interspecific competition to tobacco resources through the separation of temporal niche,which constituting the ecological foundations of H.armigera and H.assulta coexisted in tobacco.
引文
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