亚洲玉米螟滞育和生物学特性地理变异的研究
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摘要
亚洲玉米螟Ostrinia furnacalis (Guenee)在中国从南到北均有分布,是中国玉米主要害虫,以幼虫蛀食茎秆和穗等,导致减产,并以老熟幼虫在秸秆内滞育越冬。本研究以亚洲玉米螟为材料,研究了其滞育及生物学特性的地理变异,结果分述如下:
     1、亚洲玉米螟的滞育诱导和解除的地理变异
     详细比较了来自6个不同地理种群亚洲玉米螟(海南乐东18.8°N,109.2°E,广西阳朔24.8°N,110.5°E,江西南昌28.8°N,115.9°E,山东泰安36.2°N,117.1°E,河北廊坊39.5°N,116.7°E和黑龙江哈尔滨44.9°N,127.2°E)的滞育诱导的光周期反应,临界光周期、敏感期和滞育持续期。结果表明:在所测试的温度,阳朔、南昌、泰安、廊坊和哈尔滨种群的光周期反应曲线均显示了典型的长日照型,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。乐东种群在25-31℃几乎所有个体发育,没有光周期反应,仅在22℃显示了较弱的光周期反应。在25℃下,以4个短光照(12 h)在不同时期干扰以长光照(16 h)为背景或以4个长光照(16 h)在不同时期干扰以短光照(12 h)为背景的实验表明,诱导和抑制的滞育比率在不同地理种群间存在显著差异;但最高光敏感时期均出现在第3-4龄幼虫期。滞育持续期在不同地理种群间存在显著差异,最北的哈尔滨种群滞育持续期最长,其次泰安、廊坊和南昌,滞育持续期与栖息地纬度呈正相关关系
     2、亚洲玉米螟不同地理种群暗期干扰的滞育反应
     在研究了亚洲玉米螟南昌种群(28°41'N,115°53'E)和哈尔滨种群(44°56'N,127°10'E)光周期反应的基础上,在25和28℃测试了1h的光脉冲干扰光周期L9:D15和L12:D12对这两个不同地理种群幼虫滞育抑制的影响。结果表明:在25和28℃下,哈尔滨种群的临界日长分别比南昌种群延长了1h40min和2h;在所有测试的暗期干扰实验中,除了极少数光脉冲干扰点外,南昌种群幼虫滞育的发生率显著低于哈尔滨种群;1h光脉冲干扰光周期L9:D15的滞育发生率明显低于干扰光周期L12:D12,且前者表现了暗期的中间对光脉冲最敏感,而后者显示了暗期的初期对光脉冲最敏感;28°C下光脉冲对滞育的抑制效果强于25℃。这些结果进一步提示了,即使在同种昆虫中,如果地理种群和实验条件不同,暗期干扰对滞育抑制的效果也可能不同。
     3、亚洲玉米螟发育历期的地理变异
     详细比较了来自5个不同地理种群亚洲玉米螟(广西阳朔24.8°N,110.5°E,江西南昌28.8°N,115.9°E,山东泰安36.2°N,117.1°E,河北廊坊39.5°N,116.7°E和黑龙江哈尔滨44.9°N,127.2°E)在20-31℃下卵、幼虫、蛹发育历期与其栖息地纬度的关系。结果表明:不同温度下的不同地理种群的卵期不存在显著差异,卵期与其栖息地纬度呈正相关。在22、25和28℃下,不同地理种群的幼虫期存在显著差异,幼虫期与其栖息地纬度呈正相关;在20和31℃下,不同地理种群的幼虫期不存在显著差异,幼虫期与其栖息地纬度呈负相关。除31。C外,不同地理种群的蛹期存在显著差异,蛹期与其栖息地纬度呈正相关。这些结果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。
     4、亚洲玉米螟发育起点温度和有效积温的地理变异
     详细比较了来自5个不同地理种群亚洲玉米螟(广西阳朔24.8°N,110.5°E,江西南昌28.8°N,115.9°E,山东泰安36.2°N,117.1°E,河北廊坊39.5°N,116.7°E和黑龙江哈尔滨44.9°N,127.2°E)的卵、幼虫、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:(1)卵的发育起点温度与栖息地纬度呈正相关关系,从南到北依次为8.50、9.24、9.18、10.02和9.96℃;有效积温则与栖息地纬度呈负相关关系,从南到北依次为79.27、76.19、77.96、73.37和75.32日度。(2)雌幼虫的发育起点温度与栖息地纬度呈正相关关系,从南到北,雌幼虫依次为6.71、9.85、9.81、10.82和7.83℃,雄幼虫的发育起点温度与栖息地纬度呈负相关关系,从南到北,雄幼虫依次为11.80、11.90、10.81、12.43和9.59℃;雌雄幼虫的有效积温则均与栖息地纬度呈正相关关系,从南到北,雌幼虫依次为480.49、440.16、450.82、397.32和591.54日度,雄幼虫依次为370.86、378.73、418.20、346.10和532.93日度。(3)雌雄蛹的发育起点温度均与栖息地纬度呈正相关关系,从南到北,雌蛹依次为10.95、12.02、11.74、11.97和11.62℃,雄蛹依次为11.25、11.93、12.01、11.97和12.61℃;雌雄蛹的有效积温均与栖息地纬度呈正相关,从南到北,雌蛹依次为111.17、107.36、111.23、106.32和114.98日度,雄蛹依次为108.58、106.84、107.45、108.01和109.46日度。实验结果揭示了亚洲玉米螟的发育起点温度和有效积温的存在地理变异现象,但变异程度因虫态不同而异。
     5、亚洲玉米螟繁殖力的地理变异
     详细比较了来自5个不同地理种群亚洲玉米螟(海南乐东18.8°N,109.2°E,江西南昌28.8°N,115.9°E,山东泰安36.2°N,117.1°E,河北廊坊39.5°N,116.7°E和黑龙江哈尔滨44.9°N,127.2°E)成虫寿命、产卵历期、产卵量,并分析了成虫寿命、产卵历期、产卵量与栖息地纬度的关系。雌雄成虫寿命随纬度升高而延长(雌虫寿命从10.20 d延长到16.40 d;雄虫寿命从8.35延长到14.30 d)。产卵历期随纬度升高而延长,产卵量随纬度升高而增大,从低纬度到高纬度,产卵历期分别为7.45,10.45,11.90,10.62,13.15 d,产卵量分别为351.55,500.09,522.90,546.76,577.95粒。本结果首次揭示了亚洲玉米螟繁殖略策的地理变异。
     6、亚洲玉米螟体重和体型的地理变异
     详细比较了来自4个不同地理种群亚洲玉米螟(海南乐东18.8°N,109.2°E,广西阳朔24.8°N,110.5°E,江西南昌28.8°N,115.9°E和河北廊坊39.5°N,116.7°E)的体重、体型大小及其与栖息地纬度的关系。结果表明:不同地理种群的亚洲玉米螟卵重随纬度的升高而逐渐增大,符合贝格曼法则(Bergmann's law),而雌雄蛹重及成虫体长、后足腿节长和前翅长均随纬度的升高而逐渐减小,符合反贝格曼法则(Converse Bergmann's law)。雌虫的前翅显著长于雄虫,其性体型二型性符合任希法贝(?)(Rensch'srule),即在雌虫体型较大的种群中,雄虫前翅比雌虫前翅增长幅度相对较大。本文结果进一步揭示了即使在同一种类昆虫中,其各个虫态体重和体型的地理变异也可能不同。
The Asian corn borer, Ostrinia furnacalis (Guenee) (Lepidoptera:Crambidae), is widely distributed from south to north in China and is a major pest of corn, Zea mays L. The Asian corn borers damage corn by boring into stem and fringe. The borers are full-grown by fall, but remain in their host plants over the winter. In this thesis, we investigated geographic variation in diapause and biological characteristics of the Asian corn borer. The results are as follows.
     1. Geographic variation in the induction and termination of diapause in the Asian corn borer
     The photoperiodic response, the critical photoperiod, sensitive stage to photoperiod and diapause termination of the Asian corn borer from six different latitudes:Ledong (LD), Hainan, (18.8°N,109.2°E); Yangshuo (YS), Guangxi, (24.8°N,110.5°E); Nanchang (NC), Jiangxi (28.8°N,115.9°E); Taian (TA), Shandong (36.2°N,117.1°E); Langfang (LF), Hebei (39.5°N,116.7°E); Haerbin (HEB), Heilongjiang (44.9°N,127.2°E) were investigated. The results indicated that the photoperiodic response curves showed a typical long day response in YS, NC, TA, LF and HEB populations at all temperatures tested. The incidence of diapause increased with latitude. The critical photoperiod prolonged with the increase in latitude and was positively correlated with latitude. Nearly all individuals in LD population develop without diapause at 25,28 and 31℃. However, LD population showed a weak photoperiodic response at 22℃. When the diapause-averting photoperiod of L16:D 8 was interrupted by 4 short days at 25℃or when the diapause-inducing photoperiod of L12: D12 was interrupted by 4 long days, there were significant differences in the incidence of diapause among different populations. However, all populations showed that the L3-L4 larvae were the stages most sensitive to the photoperiod. The duration of diapause was significantly different among different populations. The duration of diapause in HEB was longest, followed by TA, LF and NC populations. The duration of diapause was positively correlated with latitude.
     2. Diapause response of night-interruption in two different geographic populations of the Asian corn borer
     The photoperiodic response and effects of night-interruption experiments, in which the scotophases of L9:D15 and L12:D12 were interrupted by 1 h light pulse, on diapause-averting, were investigated at 25 and 28℃in Nanchang (28°41′N,115°53′E) and Haerbin (44°56′N,127°10′E) populations of O. furnacalis. The results showed that the critical daylength in Haerbin population at 25 and 28℃was 1h 40 min and 2 h longer than that in Nanchang population, respectively. At all night-interruption experiments, the incidences of diapause in almost all positions of light pulse in Nanchang population were significantly lower than that in Haerbin population. When the light pulse was used to interrupt L9:D15 and L12:D12, the incidence of diapause was lower in L9:D15 than that in L12:12. Furthermore, the most highly photo-sensitive position occurred in the middle of scotophase at L9:D15, whereas the most highly photo-sensitive position occurred in the early scotophase at 25℃. The effect of the light pulse on diapause-averting at 28℃was greater than that at 25℃. Our results further suggest that effect of night-interruption on diapause-averting may be different even in the same species depending on different geographic populations and the experimental conditions.
     3. Geographic variation in developmental duration of the Asian corn borer
     The developmental duration of egg, larva and pupa of the Asian corn borer from five different latitude (Yangshuo, Guangxi,24.8°N,110.5°E; Nanchang, Jiangxi,28.8°N, 115.9°E; Taian, Shandong,36.2°N,117.1°E; Langfang, Hebei,39.5°N,116.7°E; Haerbin, Heilongjiang,44.9°N,127.2°E) were compared at 20-25℃. And then the relationship between egg duration, larval duration, pupal duration and latitude was analyzed. The results indicated that there were not significant differences in egg durations among the different latitudes at different temperatures and the egg duration was positively correlated with latitude. There were significant differences in larval durations among different latitudes at 22,25 and 28℃, and the larval duration was positively correlated with latitude. However, the larval duration was not significantly different at 20 and 31℃, and there were negative correlations between the larval duration and latitude. There were significant differences in pupal durations among different latitudes except 31℃and there were positive correlations between the pupal duration and latitude. These results further suggest that the geographic variation in developmental duration may be different in different stage and different experimental conditions in the same specie of insects.
     4. Geographic variation in lower development threshold and sum of effective temperatures of the Asian corn borer
     The LDT and SET of egg, larva and pupa of the ACB from five different latitudes: Yangshuo, Guangxi, (24.8°N,110.5°E); Nanchang, Jiangxi (28.8°N,115.9°E); Taian, Shandong (36.2°N,117.1°E); Langfang, Hebei (39.5°N,116.7°E); Haerbin, Heilongjiang (44.9°N,127.2°E) were compared. And then the relationship between LDT/SET and latitude was analyzed. The results indicated:(1) The LDT increased with increasing latitude. form south to north, the LDT is 8.50、9.24、9.18、10.02 and 9.96℃, respectively; while the SET decreased with increasing latitude, form south to north, the SET is 79.27、76.19、77.96、73.37 and 75.32 day-degrees (dd), respectively. (2) The LDT of female larvae increased with increasing latitude and The LDT of male larvae decreased with increasing latitude. Form south to north, the LDT of female larvae is 6.71、9.85、9.81、10.82 and 7.83℃, respectively; and the LDT of male larvae is 11.80、11.90、10.81、12.43 and 9.59℃, respectively, while the SET of female and male larvae increased with increasing latitude. Form south to north, the SET of female larvae is 480.49、440.16、450.82、397.32 and 591.54 dd, respectively, and the SET of male larvae is 370.86、378.73、418.20、346.10 and 532.93 dd, respectively. (3) the LDT of female and male pupae increased with increasing latitude. Form south to north, the LDT of female pupae is 10.95、12.02、11.74、11.97 and 11.62℃, respectively, and the LDT of male larvae is 11.25、11.93、12.01、11.97 and 12.61℃, respectively, while the SET of female and male larvae also increased with increasing latitude. Form south to north, the SET of female larvae is 111.17、107.36、111.23、106.32 and 114.98 dd, respectively, and the SET of male larvae is 108.58、106.84、107.45.108.01 and 109.46 dd, respectively. These results indicated the LDT and SET of the ACB varied from populations, while the variation extent differed by stages.
     5. Geographic variation in fecundity of the Asian corn borer
     The longevity of adult, oviposition period and egg-production of the Asian corn borer from five different latitudes:Ledong, Hainan (18.8°N,109.2°E); Nanchang, Jiangxi (28.8°N,115.9°E); Taian, Shandong (36.2°N,117.1°E); Langfang, Hebei (39.5°N,116.7°E) and Haerbin, Heilongjiang (44.9°N,127.2°E) were compared. The relationship between longevity, oviposition period, egg-production and latitude was analyzed. The results indicated that the longevity of female and male gradually increased with latitude (from 10.20 to 16.40 d for female; from 8.35 to 13.30 d for male). Both oviposition period and egg-production also increased with latitude. From low latitude to high latitude, the oviposition period was 7.45,10.45,11.90,10.62,13.15 d, and the egg-production was 351.55,500.09,522.90,546.76,577.95 eggs, respectively. This result first reveals the geographic variation in reproductive tactics of O. furnacalis.
     6. Geographic variation in body weight and size of the Asian corn borer
     The body weight and body size of the Asian corn borer from four different latitudes: Ledong, Hainan (18.8°N,109.2°E); Yangshuo, Guangxi, (24.8°N,110.5°E); Nanchang, Jiangxi (28.8°N,115.9°E); Langfang, Hebei (39.5°N,116.7°E) were compared. And then the relationship between body weight, body size and latitude was analyzed. The results indicated that egg weight of the Asian corn borer gradually increased with an increase in latitude that followed Bergmann's law, whereas the pupal weight, body length, hind femur length and fore wing length for both female and male gradually increased with an increase in latitude that followed Converse Bergmann's law. Fore wing length of the female is significantly longer than that of the male in the four geographic populations and the sexual size dimorphism followed the Rensch's rule, with the fore wing length of males relatively larger compared to that of females in populations with large body size. These results further suggest that the geographic variation in body weight and size may be different in different stage in the same specie of insects.
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