棉铃虫不同地理种群滞育及生物学特性的比较研究
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摘要
棉铃虫Helicoverpa armigera (Hubner)是以蛹越冬的兼性滞育昆虫,是全球最重要的农业害虫之一,主要为害棉花、玉米等作物,广泛分布于世界各气候带,是一种多化性害虫。滞育特性是反映棉铃虫生态适应性的重要指标,也是决定其分布地域的主要因素。本论文以棉铃虫为研究材料,系统比较了棉铃虫不同地理种群滞育及生物学特性的差异,研究结果分述如下：
1、棉铃虫不同地理种群滞育诱导及解除的比较
较系统比较了来自中国的5个地理种群[广东广州(简称GZ,23.08°N,113.14°E)、江西永修(简称YX,29.04°N,115.82°E)、山东泰安(简称TA,36.15°N,116.59°E)、河北廊坊(简称LF,39.31°N,116.42°E)和辽宁喀佐(简称KZ,41.34°N,120.27°E)]的滞育诱导的光周期反应、临界光周期和滞育持续期。结果表明：在恒温20-28℃下,棉铃虫滞育率随地理纬度的上升而逐渐增加,临界光周期随纬度的上升而逐渐延长,临界光周期与其栖息地纬度呈正相关关系,纬度每上升5°,临界光周期约增加1.2h左右。广州种群只在20℃、12-13h短日照下才有低的滞育率(26.53-21.05%)；永修种群在温度超过22℃时,任何光周期都不能诱导滞育。不同地理种群的滞育持续期存在显著差异,滞育持续期随栖息地纬度升高而延长,与纬度呈正相关关系。滞育的解除不需要低温的刺激,高温可以加速滞育的解除,但低温处理可使成虫羽化相对集中。低温处理对滞育的解除也存在地理变异,北方种群对低温的敏感性更弱些。这些结果进一步揭示了南方种群对温度的敏感性强于北方种群,而北方种群对光周期的敏感性强于南方种群。
2、棉铃虫不同地理种群滞育诱导的光敏感性的比较
系统调查了棉铃虫江西永修种群、山东泰安种群和辽宁喀佐种群幼虫在20℃,交替接受长光周期(L16：D8)和短光周期(L12：D12)下的滞育诱导的光敏感阶段。当用5d长光周期干扰生长在短光周期(L12：D12)背景条件下的幼虫时,最高光敏感期：永修种群出现在4龄第4d至5龄第3d(77.5%～93.7%个体发育),泰安种群出现在3龄第5d至5龄第1d(64.06%～89.55%个体发育),喀佐种群出现在第6龄(57.69%-60.78%个体发育)。当泰安和喀佐种群幼虫分别饲养在交替的短光周期(L12：D12)和长光周期(L16：D8)时,泰安种群幼虫光敏感龄期主要出现在5龄前,而喀佐种群幼虫光敏感龄期主要出现在4龄后。这些结果揭示了棉铃虫滞育诱导的光敏感性存在地理变异。
3、温周期对棉铃虫不同地理种群幼虫发育及蛹滞育的影响
系统调查了棉铃虫广东广州种群、江西永修种群、山东泰安种群和辽宁喀佐种群在滞育诱导的光周期(L12：D12)条件下,温周期(光期高温20-28℃,暗期低温5和9℃)对幼虫发育和蛹滞育的影响。结果表明：当光期温度相同时,各地理种群20T(温期thermophase,20℃):5C(低温期cryophase,5℃)和20T：9C,22T：5C和22T：9C,25T：5C和25T：9C,28T：5C和28T：9C之间幼虫的发育历期没有显著差异(P>0.05);而在平均温度相同或相近的情况下,各地理种群T20：9C(平均温度14.5℃)下的幼虫发育历期显著长于在22T：5C(平均温度13.5℃)下的发育历期(P<0.05)；20T：20C(恒温20℃)的幼虫发育历期也显著长于28T：9C(平均温度18.5℃)(P<0.05),表明幼虫发育主要受光期温度的影响。在平均温度相同或相近的情况下,各地理种群温周期20T：9C的滞育率高于22T：5C的滞育率,25T：20C的滞育率也高于28T：9C；各地理种群温周期20T：20C的滞育率显著高于28T：5C的滞育率(平均温度14.5℃)(P<0.05)。当光期温度相同时,20T：5C和20T：9C,22T：5C和22T：9C,25T：5C和25T：9C,28T：5C和28T：9C之间滞育率没有显著差异(P>0.05),表明光期温度是影响棉铃虫滞育的主要因素。在相同的温周期下,幼虫发育历期及滞育率存在显著的地理变异,幼虫历期和滞育率与其栖息地纬度均呈正相关关系,揭示了棉铃虫对温度的敏感性随着纬度的南移而逐渐增强。
4、暗期干扰对棉铃虫两个不同地理种群滞育抑制作用的比较
在研究了棉铃虫泰安种群和喀佐种群光周期反应的基础上,在滞育诱导的短光周期下(L12：D12和L9：D15),分别测试了暗期不同时段1h光脉冲对这两个不同地理种群滞育抑制的影响。25和22℃下的光周期反应显示了泰安种群在长暗期11-14h的滞育率均显著低于喀佐种群；泰安种群的临界暗长分别为11.7h和11.5h,喀佐种群分别为10.5h和10.3h,泰安种群均比喀佐种群长1.2h。在所测试的暗期干扰实验中,除了极少数光脉冲干扰点外,泰安种群蛹滞育率显著低于喀佐种群,但两者的滞育反应曲线基本相似。在短光周期L9：D15下,泰安种群和喀佐种群均显示了光脉冲落入暗期的第9-11h最有效地抑制了滞育的发生。在短光周期L12：D12下,泰安种群和喀佐种群在25℃时均显示了光脉冲落入暗期的第3-4h和第10h导致了最低的滞育发生；但在22℃时,喀佐种群只在暗期的第3-4h显示了最高的滞育抑制。这些结果揭示了偏南的泰安种群对暗期干扰的敏感性强于偏北的喀佐种群,但这两个地理种群的最高光敏感位点基本相同。
5、光温条件对棉铃虫不同地理种群滞育强度的影响
系统地比较了光温条件对来自中国的4个地理种群(广东广州、江西永修、山东泰安和辽宁喀佐)棉铃虫滞育强度的影响。结果表明：光周期对棉铃虫的滞育强度有明显影响,在日长11-14h范围内,更长的日照能诱导更强的滞育,但不同的种群表现不一,北方种群对光周期的敏感性明显强于南方种群。滞育强度主要受温周期(光期高温,暗期低温)的光期温度的影响,南方GZ种群温周期比恒温有更强的诱导效应,北方KZ和TA种群则相反,温周期所诱导的滞育强度并不高于恒温,有时比恒温还低,而YX种群则温周期与恒温具有相同的滞育诱导强度。更高的温度(恒温)能诱导更强的滞育,纬度越高,这种趋势越明显。棉铃虫滞育强度也存在明显地理变异,在相同的光期温度(20-25℃)下,滞育强度与栖息地纬度呈负相关关系。
6、棉铃虫不同地理种群越冬蛹羽化节律和性比的比较
系统详细地比较了来自中国的4个地理种群(广东广州、江西永修、山东泰安和辽宁喀佐)的滞育蛹的羽化习性、羽化进度、雌雄虫羽化差异及其性比结构。结果表明：棉铃虫越冬蛹羽化节律存在明显地理变异,南方(亚热带型)种群多集中在16：00-23：00内羽化(约占94-97%),而北方(温带型)种群在17：00-6：00整个夜间均有不同程度的羽化(占98%),且北方种群羽化高峰比南方种群约晚1h。在相同条件下,北方种群均较南方种群提早9-13d羽化。另外,棉铃虫越冬代成虫的性比也存在明显地理变异,南方种群雌雄比为1：1,而北方种群雌雄比为1：1.8。各种群越冬代成虫均存在雌性先熟现象,雌虫比雄虫早3-7d羽化。
7、棉铃虫不同地理种群发育历期的比较
系统比较了来自中国的4个地理种群(广东广州、江西永修、山东泰安、辽宁喀佐)在20、22、25和28℃恒温下的幼虫、非滞育蛹和滞育蛹发育历期及其与栖息地纬度的关系。结果表明,棉铃虫幼虫发育历期随栖息地纬度的升高而缩短,在25℃下,与地理纬度存在明显的负相关关系,其它温度下呈不显著的负相关关系,雌雄幼虫发育历期相似,差异不显著；在20℃下,蛹历期(包括滞育蛹和非滞育蛹)与栖息地纬度呈显著负相关,其它温度下的蛹期与栖息地纬度呈正相关,雄蛹历期均要略长于雌蛹历期,但差异不显著。这些结果进一步揭示了即使在同一种类昆虫中,不同实验条件,其各个虫态发育历期的地理变异也可能不同。
8、棉铃虫不同地理种群发育起点温度和有效积温的比较
系统比较了来自中国的4个地理种群(广东广州、江西永修、山东泰安、辽宁喀佐)在20、22、25和28℃恒温下的幼虫、非滞育蛹和滞育蛹发育起点温度和有效积温及其与栖息地纬度的关系。结果表明,在实验室人工饲料饲养的条件下,棉铃虫幼虫、非滞育蛹和滞育蛹发育起点温度分别为10.18-12.19℃、10.60-13.50℃、15.56-17.96℃,有效积温分别为254.91-300.89日度、126.21-204.43日度、181.03～363.39日度,滞育蛹的发育起点温度比非滞育蛹高3.39-5.50℃(其中雌性高2.08～5.41℃,雄性高3.52-6.03℃),其有效积温比非滞育蛹高35.56～165.89日度(其中雌性高72.75-187.68日度,雄性高2.45-158.96日度)。棉铃虫发育起点温度和有效积温均存在明显地理变异,随栖息地纬度的升高,幼虫、非滞育蛹和滞育蛹的发育起点温度均呈下降趋势,呈负相关关系,而有效积温则相反,呈上升趋势,呈正相关关系。
The cotton bollworm, Helicoverpa armigera (Hubner) is one of the most important agricultural pests in the global and mainly damages cotton, corn and other crops. It widely distributes in different climatic zones of the world, and it is a kind of the multivoltine pests, with a facultative winter pupal diapause. Diapause characteristics are reflected in an important indicator of ecological adaptation of the cotton bollworm, but also are the main factors to determine their geographical distribution. In this thesis, we systematically investigated the differences of diapause and biological characteristics among different geographic populations of the cotton bollworm. The results are as follows.
1. Comparisons of diapause-induction and-termination among different geographic populations
The geographic variations in the induction and termination of diapause were systematically investigated in the cotton bollworm, Helicoverpa armigera from5different geographic populations (Guangzhou population,23.08°N,113.14°E; Yongxiu population,29.04°N,115.82°N; Taian population,36.15°N,116.59°E; Langfang population,39.31°N,116.42°E; Kazuo population,41.34°N,120.27°E) in China. The photoperiodic response cueves indicated that the incidence of diapause and the critical photoperiod for pupal diapause induction increased with increasing geographical latitude at20-28℃. The critical daylength changed by about1.2h with every5°of latitude. Guangzhou population exhibited lower diapause rates (26.53～21.05%) under the short days of12-13h at20℃. Yongxiu population developed without diapause regardless of photoperiods when the temperature exceeds22℃. The duration of diapause was significantly different among different populations, and was positively correlated with latitude. The termination of diapause did not require cold stimulation, and high temperature could enhance the termination of diapause. However, chilling processing allows the relative concentration of adult emergence. The northern populations were lower sensitive to chilling than the sourthern populations. These results further revealed that the sensitivity to temperature is stronger in southern populations than in the northern populations, while the sensitivity to photoperiod is stronger in northern populations than in the southern population.
2. Comparisons of photosensitivity of diapause during larval development among different geographic populations
The geographic variation of photosensitivity was investigated by transferring larvae between long photoperiod (L16:D8) and short photoperiod (L12:D12) under the20℃conditions in3different geographic populations (Yongxiu population, Taian population, Kazuo population). When the photoperiodic background of L12:D12was interrupted by5×L16:D8at different larval stages, the most photosensitive stage occurred in the L4/4to the L5/3stages for Yongxiu population, the L3/5to the L5/1stages for Taian population and the L6stage for Kazuo population. When Taian population and Kazuo populations were exposed to alternating short photoperiod and long photoperiod, respectively, Taian population showed the most photosensitive stage before the fifth instar, Kazuo population showed the most photosensitive stage after the fourth instar. These results revealed that the most photosensitive stage is different in different geographic populations in H. armigera.
3. Effects of thermoperiod on larval development and pupal diapause in different geographic populations
The effects of thermoperiod on larval development and pupal diapause in4different geographic populations (Guangzhou population, Yongxiu population, Taian population and Kazuo population) were systematically investigated under the photoperiod of L12:D12. The results showed that, at the same photophase temperature, there were no significant differences in the larval duration(P>0.05) between20T (thermophase,20℃):5C(cryophase,5℃)(12h photophase temperature:12h scotophase temperature)(the rest in the same analogy) and20T:9C,22T:5C and22T:9C,25T:5C and25T:9C,28T:5C and28T:9C in4different populations. However, at the same or near average temperature, the larval duration at20T:9C (an average temperature of14.5℃) was significantly higher than that at22T:5C (an average temperature of13.5℃)(P<0.05); the larval duration at20T:20C (a constant temperature of20℃) was significantly higher than that at28T:9C (an average temperature of18.5℃)(P<0.05), indicating that the larval development was influenced strongly by the photophase temperature. At the same or near average temperature, the diapause rate in4different populations at20T:9C was higher than that at22T:5C, the diapause rate at25T:20C (an average temperature of22.5℃) was higher than that at28T:9C (an average temperature of18.5℃); diapause rate in4different populations at20T:20C was significantly higher than that at28T:5C (an average temperature of16.5℃)(P<0.05). However, there were significant differences at the same photophase temperature between20T:5C and20T:9C,22T:5C and22T:9C,25T:5C and25T:9C,28T:5C and28T:9C (P>0.05). This indicates the diapause induction is influenced strongly by the photophase temperature. At the same thermoperiod, the larval duration and diapause rate are significantly different among different geographic populations. The larval duration and diapause rate is positively correlated with latitude, suggesting that sensitivity to temperature is gradually enhanced with a decrease in latitude.
4. A comparison of night-interruption on diapause-averting among two different geographic populations
So far, all the photoperiodic responses in insects are highly sensitive to night-interruption, and the long night effect is reversed by a light break. In the present study, the photoperiodic response and effects of night-interruption on diapause-averting, in which the scotophases of L9:D15and L12:D12were interrupted by1h light pulse, were investigated at25℃and22℃in Taian (TA,36.15°N,116.59°E) and Kazuo (KZ,41.34°N,120.27°E) populations of the cotton bollworm Helicoverpa armigera. The photoperiodic response curves showed that the incidences of diapause in the scotophases of11-14h were significantly lower in Taian population than in Kazuo population; the critical night length in Taian population at25℃and22℃was11.7h and11.5h, respectively, and1h20min longer than that in Kazuo population (10.5h and10.3h). At all night-interruption experiments, the incidences of diapause in almost all positions of light pulse in Taian population were significantly lower than that in Kazuo population, however, the diapause response curves was similar in the two populations. Under L9:D15, the incidence of diapause was inhibited most effectively in both Taian population and Kazuo population when the light pulses were placed9-11in the darkness. Under L12:D12, the incidence of diapause was lowest at25℃in both Taian population and Kazuo population when the light pulses were placed3-4h and10h in the darkness; however, Kazuo population only showed a peak of diapaus-averting occurring in3-4h in the darkness at22℃. Our results reveal that southern Taian population is more sensitive to night-interruption than northern Kazuo population, but the most highly photosensitive position is similar.
5. Effects of photoperiodic and temperature on the diapause intensity in different geographic populations
The effects of photoperiod and temperature on diapause intensity of Helicoverpa armigera from4different geographic populations (Guangzhou population, Yongxiu population, Taian population, and Kazuo population) were systematically investigated. The results indicated that photoperiod has a significant influence on diapause intensity in the range from11to14h daylengths. Generally, longer daylength can induce stronger diapause intensity. The diapause intensity was influenced strongly by the photophase temperature of thermoperiod. The diapause intensity induced at thermoperiod temperature was stronger than that at constant temperature in Guangzhuo population. On the contrary, the diapause intensity induced at thermoperiod temperature was not stronger than that at constant temperature in northern Kazuo and Taian population. However, there was the same diapause intensity in Yongxiu population regardless of thermoperiod or constant temperature. Higher constant temperature induced a stronger diapause, the more northern latitudes, and this trend more evident. The diapause intensity was negatively correlated with latitude under the same photophase temperature (20-25℃).
6. Comparisons of eclosion rhythm and sex ratio in overwintering pupae among different geographic populations
The eclosion rhythm and sex ratio of overwintering pupae from4different geographic populations (Guangzhou population, Yongxiu population, Taian population and Kazuo population) were systematically investigated. The results indicated that the eclosion behavior of the southern populations (subtropical geotypes) mainly occurred at16:00～23:00, about accounted for94～97%, but that of the northern populations (temperate geotypes) occurred at17:00～6:00, all night, and about accounted for98%. The eclosion peak times of the northern populations were one hour later than that of the southern populations. Under same conditions, the time of eclosion in the northern populations was suspended for9～13days in contrast to the time in the southern populations. In addition, there was distinctly geographic variation in the sex ratio of adults emerged from overwintering pupae, the sex ratio of the southern population females and males was1:1, but that of the northern population was1:1.8. There was a phenomenon of protogyny in overwinter generation adults; the time of female eclosion were3～7days earlier than that of male eclosion.
7. Comparisons of the developmental duration among different geographic populations
The developmental durations of larvae, non-diapause pupae and diapause pupae from4different geographic populations (Guangzhou population, Yongxiu population, Taian population and Kazuo population) were systematically investigated under20,22,25and28℃. The results showed that the developmental duration of larvae increased with increasing geographical latitude, Larval duration was significantly negatively correlated with latitude at25℃, but at the other temperature, the larval duration was not significantly negatively correlated with latitude, and the development duration of female and male larvae is similar without showing significant differences. At20℃, there was significant negative correlation between pupal duration and latitude, but at the other temperature, there was positive correlation, and male pupal duration was slightly longer than female pupal duration, but the difference was not significant. These results further suggest that the geographic variation in developmental duration may be different in different stage and different experiment conditions in the same specie of insects.
8. Comparisons of lower development threshold (LDT) and sum of effective temperatures (SET) among different geographic populations
The LDT and SET of larvae, non-diapause pupae and diapause pupae from4different geographic populations (Guangzhou population, Yongxiu population, Taian population and Kazuo population) were systematically investigated under20,22,25and28℃. The results showed that the LDT of larvae, non-diapause pupae and diapause pupae were10.18-12.19,10.60～13.50and15.56～17.96℃, respectively, their SET were254.91～300.89,126.21～204.43and181.03～363.39day-degrees, respectively. The LDT of diapause pupae was3.39～5.50℃more than that of non-diapause pupae, and its SET was35.56～165.89day-degrees more than that non-diapause pupae. The LDT of larvae, non-diapause pupae and diapause pupae decreased with increasing geographical latitude; whereas the SET increased with increasing geographical latitude.
1、棉铃虫不同地理种群滞育诱导及解除的比较
较系统比较了来自中国的5个地理种群[广东广州(简称GZ,23.08°N,113.14°E)、江西永修(简称YX,29.04°N,115.82°E)、山东泰安(简称TA,36.15°N,116.59°E)、河北廊坊(简称LF,39.31°N,116.42°E)和辽宁喀佐(简称KZ,41.34°N,120.27°E)]的滞育诱导的光周期反应、临界光周期和滞育持续期。结果表明：在恒温20-28℃下,棉铃虫滞育率随地理纬度的上升而逐渐增加,临界光周期随纬度的上升而逐渐延长,临界光周期与其栖息地纬度呈正相关关系,纬度每上升5°,临界光周期约增加1.2h左右。广州种群只在20℃、12-13h短日照下才有低的滞育率(26.53-21.05%)；永修种群在温度超过22℃时,任何光周期都不能诱导滞育。不同地理种群的滞育持续期存在显著差异,滞育持续期随栖息地纬度升高而延长,与纬度呈正相关关系。滞育的解除不需要低温的刺激,高温可以加速滞育的解除,但低温处理可使成虫羽化相对集中。低温处理对滞育的解除也存在地理变异,北方种群对低温的敏感性更弱些。这些结果进一步揭示了南方种群对温度的敏感性强于北方种群,而北方种群对光周期的敏感性强于南方种群。
2、棉铃虫不同地理种群滞育诱导的光敏感性的比较
系统调查了棉铃虫江西永修种群、山东泰安种群和辽宁喀佐种群幼虫在20℃,交替接受长光周期(L16：D8)和短光周期(L12：D12)下的滞育诱导的光敏感阶段。当用5d长光周期干扰生长在短光周期(L12：D12)背景条件下的幼虫时,最高光敏感期：永修种群出现在4龄第4d至5龄第3d(77.5%～93.7%个体发育),泰安种群出现在3龄第5d至5龄第1d(64.06%～89.55%个体发育),喀佐种群出现在第6龄(57.69%-60.78%个体发育)。当泰安和喀佐种群幼虫分别饲养在交替的短光周期(L12：D12)和长光周期(L16：D8)时,泰安种群幼虫光敏感龄期主要出现在5龄前,而喀佐种群幼虫光敏感龄期主要出现在4龄后。这些结果揭示了棉铃虫滞育诱导的光敏感性存在地理变异。
3、温周期对棉铃虫不同地理种群幼虫发育及蛹滞育的影响
系统调查了棉铃虫广东广州种群、江西永修种群、山东泰安种群和辽宁喀佐种群在滞育诱导的光周期(L12：D12)条件下,温周期(光期高温20-28℃,暗期低温5和9℃)对幼虫发育和蛹滞育的影响。结果表明：当光期温度相同时,各地理种群20T(温期thermophase,20℃):5C(低温期cryophase,5℃)和20T：9C,22T：5C和22T：9C,25T：5C和25T：9C,28T：5C和28T：9C之间幼虫的发育历期没有显著差异(P>0.05);而在平均温度相同或相近的情况下,各地理种群T20：9C(平均温度14.5℃)下的幼虫发育历期显著长于在22T：5C(平均温度13.5℃)下的发育历期(P<0.05)；20T：20C(恒温20℃)的幼虫发育历期也显著长于28T：9C(平均温度18.5℃)(P<0.05),表明幼虫发育主要受光期温度的影响。在平均温度相同或相近的情况下,各地理种群温周期20T：9C的滞育率高于22T：5C的滞育率,25T：20C的滞育率也高于28T：9C；各地理种群温周期20T：20C的滞育率显著高于28T：5C的滞育率(平均温度14.5℃)(P<0.05)。当光期温度相同时,20T：5C和20T：9C,22T：5C和22T：9C,25T：5C和25T：9C,28T：5C和28T：9C之间滞育率没有显著差异(P>0.05),表明光期温度是影响棉铃虫滞育的主要因素。在相同的温周期下,幼虫发育历期及滞育率存在显著的地理变异,幼虫历期和滞育率与其栖息地纬度均呈正相关关系,揭示了棉铃虫对温度的敏感性随着纬度的南移而逐渐增强。
4、暗期干扰对棉铃虫两个不同地理种群滞育抑制作用的比较
在研究了棉铃虫泰安种群和喀佐种群光周期反应的基础上,在滞育诱导的短光周期下(L12：D12和L9：D15),分别测试了暗期不同时段1h光脉冲对这两个不同地理种群滞育抑制的影响。25和22℃下的光周期反应显示了泰安种群在长暗期11-14h的滞育率均显著低于喀佐种群；泰安种群的临界暗长分别为11.7h和11.5h,喀佐种群分别为10.5h和10.3h,泰安种群均比喀佐种群长1.2h。在所测试的暗期干扰实验中,除了极少数光脉冲干扰点外,泰安种群蛹滞育率显著低于喀佐种群,但两者的滞育反应曲线基本相似。在短光周期L9：D15下,泰安种群和喀佐种群均显示了光脉冲落入暗期的第9-11h最有效地抑制了滞育的发生。在短光周期L12：D12下,泰安种群和喀佐种群在25℃时均显示了光脉冲落入暗期的第3-4h和第10h导致了最低的滞育发生；但在22℃时,喀佐种群只在暗期的第3-4h显示了最高的滞育抑制。这些结果揭示了偏南的泰安种群对暗期干扰的敏感性强于偏北的喀佐种群,但这两个地理种群的最高光敏感位点基本相同。
5、光温条件对棉铃虫不同地理种群滞育强度的影响
系统地比较了光温条件对来自中国的4个地理种群(广东广州、江西永修、山东泰安和辽宁喀佐)棉铃虫滞育强度的影响。结果表明：光周期对棉铃虫的滞育强度有明显影响,在日长11-14h范围内,更长的日照能诱导更强的滞育,但不同的种群表现不一,北方种群对光周期的敏感性明显强于南方种群。滞育强度主要受温周期(光期高温,暗期低温)的光期温度的影响,南方GZ种群温周期比恒温有更强的诱导效应,北方KZ和TA种群则相反,温周期所诱导的滞育强度并不高于恒温,有时比恒温还低,而YX种群则温周期与恒温具有相同的滞育诱导强度。更高的温度(恒温)能诱导更强的滞育,纬度越高,这种趋势越明显。棉铃虫滞育强度也存在明显地理变异,在相同的光期温度(20-25℃)下,滞育强度与栖息地纬度呈负相关关系。
6、棉铃虫不同地理种群越冬蛹羽化节律和性比的比较
系统详细地比较了来自中国的4个地理种群(广东广州、江西永修、山东泰安和辽宁喀佐)的滞育蛹的羽化习性、羽化进度、雌雄虫羽化差异及其性比结构。结果表明：棉铃虫越冬蛹羽化节律存在明显地理变异,南方(亚热带型)种群多集中在16：00-23：00内羽化(约占94-97%),而北方(温带型)种群在17：00-6：00整个夜间均有不同程度的羽化(占98%),且北方种群羽化高峰比南方种群约晚1h。在相同条件下,北方种群均较南方种群提早9-13d羽化。另外,棉铃虫越冬代成虫的性比也存在明显地理变异,南方种群雌雄比为1：1,而北方种群雌雄比为1：1.8。各种群越冬代成虫均存在雌性先熟现象,雌虫比雄虫早3-7d羽化。
7、棉铃虫不同地理种群发育历期的比较
系统比较了来自中国的4个地理种群(广东广州、江西永修、山东泰安、辽宁喀佐)在20、22、25和28℃恒温下的幼虫、非滞育蛹和滞育蛹发育历期及其与栖息地纬度的关系。结果表明,棉铃虫幼虫发育历期随栖息地纬度的升高而缩短,在25℃下,与地理纬度存在明显的负相关关系,其它温度下呈不显著的负相关关系,雌雄幼虫发育历期相似,差异不显著；在20℃下,蛹历期(包括滞育蛹和非滞育蛹)与栖息地纬度呈显著负相关,其它温度下的蛹期与栖息地纬度呈正相关,雄蛹历期均要略长于雌蛹历期,但差异不显著。这些结果进一步揭示了即使在同一种类昆虫中,不同实验条件,其各个虫态发育历期的地理变异也可能不同。
8、棉铃虫不同地理种群发育起点温度和有效积温的比较
系统比较了来自中国的4个地理种群(广东广州、江西永修、山东泰安、辽宁喀佐)在20、22、25和28℃恒温下的幼虫、非滞育蛹和滞育蛹发育起点温度和有效积温及其与栖息地纬度的关系。结果表明,在实验室人工饲料饲养的条件下,棉铃虫幼虫、非滞育蛹和滞育蛹发育起点温度分别为10.18-12.19℃、10.60-13.50℃、15.56-17.96℃,有效积温分别为254.91-300.89日度、126.21-204.43日度、181.03～363.39日度,滞育蛹的发育起点温度比非滞育蛹高3.39-5.50℃(其中雌性高2.08～5.41℃,雄性高3.52-6.03℃),其有效积温比非滞育蛹高35.56～165.89日度(其中雌性高72.75-187.68日度,雄性高2.45-158.96日度)。棉铃虫发育起点温度和有效积温均存在明显地理变异,随栖息地纬度的升高,幼虫、非滞育蛹和滞育蛹的发育起点温度均呈下降趋势,呈负相关关系,而有效积温则相反,呈上升趋势,呈正相关关系。
The cotton bollworm, Helicoverpa armigera (Hubner) is one of the most important agricultural pests in the global and mainly damages cotton, corn and other crops. It widely distributes in different climatic zones of the world, and it is a kind of the multivoltine pests, with a facultative winter pupal diapause. Diapause characteristics are reflected in an important indicator of ecological adaptation of the cotton bollworm, but also are the main factors to determine their geographical distribution. In this thesis, we systematically investigated the differences of diapause and biological characteristics among different geographic populations of the cotton bollworm. The results are as follows.
1. Comparisons of diapause-induction and-termination among different geographic populations
The geographic variations in the induction and termination of diapause were systematically investigated in the cotton bollworm, Helicoverpa armigera from5different geographic populations (Guangzhou population,23.08°N,113.14°E; Yongxiu population,29.04°N,115.82°N; Taian population,36.15°N,116.59°E; Langfang population,39.31°N,116.42°E; Kazuo population,41.34°N,120.27°E) in China. The photoperiodic response cueves indicated that the incidence of diapause and the critical photoperiod for pupal diapause induction increased with increasing geographical latitude at20-28℃. The critical daylength changed by about1.2h with every5°of latitude. Guangzhou population exhibited lower diapause rates (26.53～21.05%) under the short days of12-13h at20℃. Yongxiu population developed without diapause regardless of photoperiods when the temperature exceeds22℃. The duration of diapause was significantly different among different populations, and was positively correlated with latitude. The termination of diapause did not require cold stimulation, and high temperature could enhance the termination of diapause. However, chilling processing allows the relative concentration of adult emergence. The northern populations were lower sensitive to chilling than the sourthern populations. These results further revealed that the sensitivity to temperature is stronger in southern populations than in the northern populations, while the sensitivity to photoperiod is stronger in northern populations than in the southern population.
2. Comparisons of photosensitivity of diapause during larval development among different geographic populations
The geographic variation of photosensitivity was investigated by transferring larvae between long photoperiod (L16:D8) and short photoperiod (L12:D12) under the20℃conditions in3different geographic populations (Yongxiu population, Taian population, Kazuo population). When the photoperiodic background of L12:D12was interrupted by5×L16:D8at different larval stages, the most photosensitive stage occurred in the L4/4to the L5/3stages for Yongxiu population, the L3/5to the L5/1stages for Taian population and the L6stage for Kazuo population. When Taian population and Kazuo populations were exposed to alternating short photoperiod and long photoperiod, respectively, Taian population showed the most photosensitive stage before the fifth instar, Kazuo population showed the most photosensitive stage after the fourth instar. These results revealed that the most photosensitive stage is different in different geographic populations in H. armigera.
3. Effects of thermoperiod on larval development and pupal diapause in different geographic populations
The effects of thermoperiod on larval development and pupal diapause in4different geographic populations (Guangzhou population, Yongxiu population, Taian population and Kazuo population) were systematically investigated under the photoperiod of L12:D12. The results showed that, at the same photophase temperature, there were no significant differences in the larval duration(P>0.05) between20T (thermophase,20℃):5C(cryophase,5℃)(12h photophase temperature:12h scotophase temperature)(the rest in the same analogy) and20T:9C,22T:5C and22T:9C,25T:5C and25T:9C,28T:5C and28T:9C in4different populations. However, at the same or near average temperature, the larval duration at20T:9C (an average temperature of14.5℃) was significantly higher than that at22T:5C (an average temperature of13.5℃)(P<0.05); the larval duration at20T:20C (a constant temperature of20℃) was significantly higher than that at28T:9C (an average temperature of18.5℃)(P<0.05), indicating that the larval development was influenced strongly by the photophase temperature. At the same or near average temperature, the diapause rate in4different populations at20T:9C was higher than that at22T:5C, the diapause rate at25T:20C (an average temperature of22.5℃) was higher than that at28T:9C (an average temperature of18.5℃); diapause rate in4different populations at20T:20C was significantly higher than that at28T:5C (an average temperature of16.5℃)(P<0.05). However, there were significant differences at the same photophase temperature between20T:5C and20T:9C,22T:5C and22T:9C,25T:5C and25T:9C,28T:5C and28T:9C (P>0.05). This indicates the diapause induction is influenced strongly by the photophase temperature. At the same thermoperiod, the larval duration and diapause rate are significantly different among different geographic populations. The larval duration and diapause rate is positively correlated with latitude, suggesting that sensitivity to temperature is gradually enhanced with a decrease in latitude.
4. A comparison of night-interruption on diapause-averting among two different geographic populations
So far, all the photoperiodic responses in insects are highly sensitive to night-interruption, and the long night effect is reversed by a light break. In the present study, the photoperiodic response and effects of night-interruption on diapause-averting, in which the scotophases of L9:D15and L12:D12were interrupted by1h light pulse, were investigated at25℃and22℃in Taian (TA,36.15°N,116.59°E) and Kazuo (KZ,41.34°N,120.27°E) populations of the cotton bollworm Helicoverpa armigera. The photoperiodic response curves showed that the incidences of diapause in the scotophases of11-14h were significantly lower in Taian population than in Kazuo population; the critical night length in Taian population at25℃and22℃was11.7h and11.5h, respectively, and1h20min longer than that in Kazuo population (10.5h and10.3h). At all night-interruption experiments, the incidences of diapause in almost all positions of light pulse in Taian population were significantly lower than that in Kazuo population, however, the diapause response curves was similar in the two populations. Under L9:D15, the incidence of diapause was inhibited most effectively in both Taian population and Kazuo population when the light pulses were placed9-11in the darkness. Under L12:D12, the incidence of diapause was lowest at25℃in both Taian population and Kazuo population when the light pulses were placed3-4h and10h in the darkness; however, Kazuo population only showed a peak of diapaus-averting occurring in3-4h in the darkness at22℃. Our results reveal that southern Taian population is more sensitive to night-interruption than northern Kazuo population, but the most highly photosensitive position is similar.
5. Effects of photoperiodic and temperature on the diapause intensity in different geographic populations
The effects of photoperiod and temperature on diapause intensity of Helicoverpa armigera from4different geographic populations (Guangzhou population, Yongxiu population, Taian population, and Kazuo population) were systematically investigated. The results indicated that photoperiod has a significant influence on diapause intensity in the range from11to14h daylengths. Generally, longer daylength can induce stronger diapause intensity. The diapause intensity was influenced strongly by the photophase temperature of thermoperiod. The diapause intensity induced at thermoperiod temperature was stronger than that at constant temperature in Guangzhuo population. On the contrary, the diapause intensity induced at thermoperiod temperature was not stronger than that at constant temperature in northern Kazuo and Taian population. However, there was the same diapause intensity in Yongxiu population regardless of thermoperiod or constant temperature. Higher constant temperature induced a stronger diapause, the more northern latitudes, and this trend more evident. The diapause intensity was negatively correlated with latitude under the same photophase temperature (20-25℃).
6. Comparisons of eclosion rhythm and sex ratio in overwintering pupae among different geographic populations
The eclosion rhythm and sex ratio of overwintering pupae from4different geographic populations (Guangzhou population, Yongxiu population, Taian population and Kazuo population) were systematically investigated. The results indicated that the eclosion behavior of the southern populations (subtropical geotypes) mainly occurred at16:00～23:00, about accounted for94～97%, but that of the northern populations (temperate geotypes) occurred at17:00～6:00, all night, and about accounted for98%. The eclosion peak times of the northern populations were one hour later than that of the southern populations. Under same conditions, the time of eclosion in the northern populations was suspended for9～13days in contrast to the time in the southern populations. In addition, there was distinctly geographic variation in the sex ratio of adults emerged from overwintering pupae, the sex ratio of the southern population females and males was1:1, but that of the northern population was1:1.8. There was a phenomenon of protogyny in overwinter generation adults; the time of female eclosion were3～7days earlier than that of male eclosion.
7. Comparisons of the developmental duration among different geographic populations
The developmental durations of larvae, non-diapause pupae and diapause pupae from4different geographic populations (Guangzhou population, Yongxiu population, Taian population and Kazuo population) were systematically investigated under20,22,25and28℃. The results showed that the developmental duration of larvae increased with increasing geographical latitude, Larval duration was significantly negatively correlated with latitude at25℃, but at the other temperature, the larval duration was not significantly negatively correlated with latitude, and the development duration of female and male larvae is similar without showing significant differences. At20℃, there was significant negative correlation between pupal duration and latitude, but at the other temperature, there was positive correlation, and male pupal duration was slightly longer than female pupal duration, but the difference was not significant. These results further suggest that the geographic variation in developmental duration may be different in different stage and different experiment conditions in the same specie of insects.
8. Comparisons of lower development threshold (LDT) and sum of effective temperatures (SET) among different geographic populations
The LDT and SET of larvae, non-diapause pupae and diapause pupae from4different geographic populations (Guangzhou population, Yongxiu population, Taian population and Kazuo population) were systematically investigated under20,22,25and28℃. The results showed that the LDT of larvae, non-diapause pupae and diapause pupae were10.18-12.19,10.60～13.50and15.56～17.96℃, respectively, their SET were254.91～300.89,126.21～204.43and181.03～363.39day-degrees, respectively. The LDT of diapause pupae was3.39～5.50℃more than that of non-diapause pupae, and its SET was35.56～165.89day-degrees more than that non-diapause pupae. The LDT of larvae, non-diapause pupae and diapause pupae decreased with increasing geographical latitude; whereas the SET increased with increasing geographical latitude.
引文
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