大猿叶虫生殖力、交配、发育速率等生物学特性的研究
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摘要
大猿叶虫Colaphellus bowringi Baly属鞘翅目Coleptera,叶甲科Chrysomelidae,无缘叶甲属Colaphellus,是我国十字花科蔬菜上的一种重要食叶害虫。本论文以大猿叶虫为材料,系统研究了不同滞育持续时间对该虫产卵的影响,该虫的交配行为,温度对该虫交配行为的影响。最后利用该虫作为材料测试了发育速率等比例假说。
1.春季不同出土时间大猿叶虫寿命和生殖力的比较：在田间,大猿叶虫C.bowringi的繁殖期出现在春季和秋季,以成虫在土中越夏和越冬。本实验在试验室20℃,LD12：12条件下详细比较了大猿叶虫龙南种群同一时间(10月份)自然条件下入土滞育,第二年春季2,3月份出土繁殖的成虫的寿命和产卵量。结果显示：2月出土雌雄成虫寿命明显短于3月出土成虫,日平均产卵量差异不显著,总产卵量由于寿命的影响也显著低于3月出土成虫。这一结果显示同一个季节中大猿叶虫早期出土成虫的比晚期出土的成虫适应性差。
2.大猿叶虫交配行为的观察：大猿叶虫C. bowringi一生能多次交配。本实验在25℃,光周期LD12：12条件下系统的观察了大猿叶虫成虫连续7天的交配行为。结果表明：该虫每日平均交配5.67+0.26次,最高可达11次,不同日龄间的交配次数存在显著差异。平均每日用于交配的时间为238+10min,占总时间的33.5%,最长可达493min,占总时间的68.5%,不同日龄间的交配时间有极显著差异。交配持续时间最短8min,最长达289min,平均为48+2min；同一日内,随着交配次数增加,交配持续时间逐渐缩短；不同日龄间的交配持续时间存在显著差异。相邻两次交配之间的间隔时间最短5min,最长300min,平均交配间隔75+3min；交配间隔时间随日龄的增加而明显延长。
3.温度对大猿叶虫交配行为的影响：在光周期LD12：12,22℃,25℃和28℃温度下,我们详细观察了该虫的交配节律,日交配频率,交配持续时间和每日总交配时间。结果表明,该虫的交配节律受温度的影响,22℃下的交配高峰(上午9：00)较25℃和28℃下的交配高峰(上午8：00)推迟了1小时。温度对日交配频率和交配持续时间也有明显影响,22℃下日交配频率显著低于25℃和28℃,而交配持续时间则显著长于25℃和28℃。然而,该虫每日总交配时间在这三种温度下相似,提示了在低温下日交配频率的下降是由于交配持续时间延长所致。
4.五种统计方法对发育速率等比例假说的测定：生物各发育阶段所持续的时间的比率不受温度影响被称为发育速率等比例。在16-30℃(以2℃递增)8个温度下,我们详细记载了大猿叶虫C. bowringi各发育阶段的数据并以五种分析方法测试了发育速率等比例的假说,这五种分析方法包括：1)直接利用基于传统线性方程描述发育速率的线性函数比较有标准误的发育起点温度(Campbell et al.,1974),2)基于Ikemoto和Takai (2000)改进的线性方程分析协方差比较不同阶段的发育起点温度,3)Jarosik等(2002)提出的方法,4)各发育时间比率的方差分析,5)时培健等(2010)提出的方法。我们的结果表明发育起点温度或上述比率不存在显著差异,并且很有力的证实了发育速率等比例假说。
The cabbage beetle, Colaphellus bowringi Baly (Coleptera:Chrysomelidae) is a serious pest of crucifers in China. In this thesis, the effect of diapause duration on oviposition, mating behavior and the effect of temperature on mating behavior of this beetle were investigated, at last, rate Isomorphy Hypothesis was examined using this beetle.
A comparison of longevity and fertility between the post-diapause adults with different emerge time in spring in the cabbage beetle, C. bowringi
There are two distinct infestations in the field of the cabbage beetle, C. bowringi Baly, one occurs in spring, another in autumn. The beetle aestivates and hibernates as an adult in the soil. In the present study, the longevity and egg production between the adults enter diapause at the same time (October) and emerged in February and March in.the next spring were compared at 20℃, LD12:12. The results showed that the longevity of both male and female adult emerged in February were significantly shorter than the adult emerged in March. There was no significant difference in the mean daily egg production between the two treatments. However, mean egg production of females emerged in March was significantly higher than that of females emerged in February because the impact of longevity, The result suggests that early emerged adult of Colaphellus bowringi has a weaker adaptation than the lately emerged adult in the same season.
Mating behavior of C. bowringi Baly
The cabbage beetle, C. bowringi mates repeatedly within their lifespan. The mating behavior of the C. bowringi was systematically observed at a photophase of LD 12:12 at 25℃for 7 consecutive days. The results showed that pairs mated on average (5.67±0.26) times per day with the highest incidence of mating of 11 times. There were significant differences in mating frequency among different age pairs. Beetle spent an average of (238±10) min mating per day, i.e.33.5% of the total time. The longest recorded amount of time engaged in mating was 493 min per day; i.e.68.5% of the total time. The total time spent mating per day was significantly different among different aged pairs. Mating duration ranged from 8 to 289 min with a mean of (48±2) min. In the course of one day, mating duration gradually shortened with increasing mating frequency. Mating duration was significantly different among different aged pairs. The interval between two consecutive mating bouts ranged from 5 to 300 min with a mean of (75±3) min. The interval between consecutive mating bouts significantly increased with age.
The effect of temperature on mating behavior of the C. bowringi Baly
The mating rhythm, daily mating frequency, mating duration and total mating time per day in the cabbage beetle were investigated at 22℃,25℃and 28℃under LD 12:12. The results showed that mating rhythm was influenced by temperature. The daily mating peak at 22℃delayed one hour compared with 25℃and 28℃. Temperature had also a significant influence on daily mating frequency and mating duration. The daily mating frequency at 22℃was significantly lower than 25℃and 28℃; whereas the mating duration at 22℃was significantly longer than at 25℃and 28℃. However, the total mating time per day was similar among the three temperatures, suggesting that the decline of mating frequency at lower temperature was due to the prolongation of mating duration.
Testing the Rate Isomorphy Hypothesis Using Five Statistical Methods
Organisms are said to be in developmental rate isomorphy when the proportions of developmental stage durations are unaffected by temperature. Comprehensive stage-specific developmental data were generated on the cabbage beetle, C. bowringi, at eight temperatures ranging from 16℃to 30℃(in 2℃increments) and five analytical methods were used to test the rate isomorphy hypothesis, including i) direct comparison of lower developmental thresholds with standard errors based on the traditional linear equation describing developmental rate as the linear function of temperature (Campbell et al.,1974), ii) analysis of covariance to compare the lower developmental thresholds of different stages based on the modified linear equation of Ikemoto and Takai (2000), iii) the method proposed by Jarosik et al. (2002), iv) analysis of variance to test for significant differences between the ratios of developmental stage durations to that of preimaginal development, and v) the approach proposed by Shi et al. (2010). The results revealed no significant difference among the lower developmental thresholds or among the aforementioned ratios, and thus convincingly confirmed the rate isomorphy hypothesis.
1.春季不同出土时间大猿叶虫寿命和生殖力的比较：在田间,大猿叶虫C.bowringi的繁殖期出现在春季和秋季,以成虫在土中越夏和越冬。本实验在试验室20℃,LD12：12条件下详细比较了大猿叶虫龙南种群同一时间(10月份)自然条件下入土滞育,第二年春季2,3月份出土繁殖的成虫的寿命和产卵量。结果显示：2月出土雌雄成虫寿命明显短于3月出土成虫,日平均产卵量差异不显著,总产卵量由于寿命的影响也显著低于3月出土成虫。这一结果显示同一个季节中大猿叶虫早期出土成虫的比晚期出土的成虫适应性差。
2.大猿叶虫交配行为的观察：大猿叶虫C. bowringi一生能多次交配。本实验在25℃,光周期LD12：12条件下系统的观察了大猿叶虫成虫连续7天的交配行为。结果表明：该虫每日平均交配5.67+0.26次,最高可达11次,不同日龄间的交配次数存在显著差异。平均每日用于交配的时间为238+10min,占总时间的33.5%,最长可达493min,占总时间的68.5%,不同日龄间的交配时间有极显著差异。交配持续时间最短8min,最长达289min,平均为48+2min；同一日内,随着交配次数增加,交配持续时间逐渐缩短；不同日龄间的交配持续时间存在显著差异。相邻两次交配之间的间隔时间最短5min,最长300min,平均交配间隔75+3min；交配间隔时间随日龄的增加而明显延长。
3.温度对大猿叶虫交配行为的影响：在光周期LD12：12,22℃,25℃和28℃温度下,我们详细观察了该虫的交配节律,日交配频率,交配持续时间和每日总交配时间。结果表明,该虫的交配节律受温度的影响,22℃下的交配高峰(上午9：00)较25℃和28℃下的交配高峰(上午8：00)推迟了1小时。温度对日交配频率和交配持续时间也有明显影响,22℃下日交配频率显著低于25℃和28℃,而交配持续时间则显著长于25℃和28℃。然而,该虫每日总交配时间在这三种温度下相似,提示了在低温下日交配频率的下降是由于交配持续时间延长所致。
4.五种统计方法对发育速率等比例假说的测定：生物各发育阶段所持续的时间的比率不受温度影响被称为发育速率等比例。在16-30℃(以2℃递增)8个温度下,我们详细记载了大猿叶虫C. bowringi各发育阶段的数据并以五种分析方法测试了发育速率等比例的假说,这五种分析方法包括：1)直接利用基于传统线性方程描述发育速率的线性函数比较有标准误的发育起点温度(Campbell et al.,1974),2)基于Ikemoto和Takai (2000)改进的线性方程分析协方差比较不同阶段的发育起点温度,3)Jarosik等(2002)提出的方法,4)各发育时间比率的方差分析,5)时培健等(2010)提出的方法。我们的结果表明发育起点温度或上述比率不存在显著差异,并且很有力的证实了发育速率等比例假说。
The cabbage beetle, Colaphellus bowringi Baly (Coleptera:Chrysomelidae) is a serious pest of crucifers in China. In this thesis, the effect of diapause duration on oviposition, mating behavior and the effect of temperature on mating behavior of this beetle were investigated, at last, rate Isomorphy Hypothesis was examined using this beetle.
A comparison of longevity and fertility between the post-diapause adults with different emerge time in spring in the cabbage beetle, C. bowringi
There are two distinct infestations in the field of the cabbage beetle, C. bowringi Baly, one occurs in spring, another in autumn. The beetle aestivates and hibernates as an adult in the soil. In the present study, the longevity and egg production between the adults enter diapause at the same time (October) and emerged in February and March in.the next spring were compared at 20℃, LD12:12. The results showed that the longevity of both male and female adult emerged in February were significantly shorter than the adult emerged in March. There was no significant difference in the mean daily egg production between the two treatments. However, mean egg production of females emerged in March was significantly higher than that of females emerged in February because the impact of longevity, The result suggests that early emerged adult of Colaphellus bowringi has a weaker adaptation than the lately emerged adult in the same season.
Mating behavior of C. bowringi Baly
The cabbage beetle, C. bowringi mates repeatedly within their lifespan. The mating behavior of the C. bowringi was systematically observed at a photophase of LD 12:12 at 25℃for 7 consecutive days. The results showed that pairs mated on average (5.67±0.26) times per day with the highest incidence of mating of 11 times. There were significant differences in mating frequency among different age pairs. Beetle spent an average of (238±10) min mating per day, i.e.33.5% of the total time. The longest recorded amount of time engaged in mating was 493 min per day; i.e.68.5% of the total time. The total time spent mating per day was significantly different among different aged pairs. Mating duration ranged from 8 to 289 min with a mean of (48±2) min. In the course of one day, mating duration gradually shortened with increasing mating frequency. Mating duration was significantly different among different aged pairs. The interval between two consecutive mating bouts ranged from 5 to 300 min with a mean of (75±3) min. The interval between consecutive mating bouts significantly increased with age.
The effect of temperature on mating behavior of the C. bowringi Baly
The mating rhythm, daily mating frequency, mating duration and total mating time per day in the cabbage beetle were investigated at 22℃,25℃and 28℃under LD 12:12. The results showed that mating rhythm was influenced by temperature. The daily mating peak at 22℃delayed one hour compared with 25℃and 28℃. Temperature had also a significant influence on daily mating frequency and mating duration. The daily mating frequency at 22℃was significantly lower than 25℃and 28℃; whereas the mating duration at 22℃was significantly longer than at 25℃and 28℃. However, the total mating time per day was similar among the three temperatures, suggesting that the decline of mating frequency at lower temperature was due to the prolongation of mating duration.
Testing the Rate Isomorphy Hypothesis Using Five Statistical Methods
Organisms are said to be in developmental rate isomorphy when the proportions of developmental stage durations are unaffected by temperature. Comprehensive stage-specific developmental data were generated on the cabbage beetle, C. bowringi, at eight temperatures ranging from 16℃to 30℃(in 2℃increments) and five analytical methods were used to test the rate isomorphy hypothesis, including i) direct comparison of lower developmental thresholds with standard errors based on the traditional linear equation describing developmental rate as the linear function of temperature (Campbell et al.,1974), ii) analysis of covariance to compare the lower developmental thresholds of different stages based on the modified linear equation of Ikemoto and Takai (2000), iii) the method proposed by Jarosik et al. (2002), iv) analysis of variance to test for significant differences between the ratios of developmental stage durations to that of preimaginal development, and v) the approach proposed by Shi et al. (2010). The results revealed no significant difference among the lower developmental thresholds or among the aforementioned ratios, and thus convincingly confirmed the rate isomorphy hypothesis.
引文
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Wang X P, Xue F S, Hua A, et al.2006. Effects of diapause duration on future reproduction in the cabbage beetle, Colaphellus bowringi:positive or negative? Physiol. Entomol.,.31:190-196.
Wei X T, Zhou Y C, Xiao H J, et al.2010. Relationship between the natural duration of diapause and post-diapause reproduction in the cabbage beetle, Colaphellus bowringi (Coleoptera: Chrysomelidae). Eur. J. Entomol.,107:337-340
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Willmer P.1991. Thermal biology and mate acquisition in ectotherms. Trends Ecol. Evol.,6:396-399.
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Xue F S, Yang A Q, Li A Q, et al.2001. Observation of hibernation and aestivation of Colaphellus bowingi (Col, chrysomelidae). Jiangxi plant protection,24:1-2[薛芳森,杨爱卿,李爱青等.2001.大猿叶虫夏眼与冬蛰的观察.江西植保,24(1)：1-2.]
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Xue Y, Chen L P.2007. Statistical Models and R Software. Tsinghua University Press, Beijing. pp. 255-256.
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