Comparison of sexual compatibility in crosses between the southern and northern populations of the cabbage beetle Colaphellus bowringi

It is widely accepted that the genetic divergence and reproductive incompat- ibility between closely related species and/or populations is often viewed as an important step toward speciation. In this study, sexual compatibility in crosses between the southern XS population and the northern TA population of the polyandrous cabbage beetle Co- laphellus bowringi was investigated by testing their mating preferences, mating latency, copulation duration, and reproductive performances of post-mating. In choice mating ex- periments, the percentages ofmatings were significantly higher in intra-population crosses than in inter-population crosses. Both isolation index （/） and index of pair sexual isolation （/PSi） indicated partial mating incompatibility or assortative mating in crosses between the two different geographical populations. In single pair mating experiments, XS females in inter-population crosses mated significantly later and copulated significantly shorter than those in intra-population crosses. However, TA females in inter-population crosses mated significantly earlier and copulated longer than those in intra-population crosses, suggesting that larger XS males may enhance heterotypic mating. The lifetime fecundity was highest in XS homotypic matings, lowest in TA homotypic matings, and intermedi- ate in heterotypic rnatings between their parents. The inter-population crosses resulted in significantly lower egg hatching rate and shorter female longevity than intra-population crosses. These results demonstrated that there exist some incompatibilities in premating, postmating-prezygotic, and postzygotic stages between the southern XS population and northern TA population of the cabbage beetle Colaphellus bowringi.     

大猿叶虫滞育进程中成虫内生殖系统的形态特征

大猿叶虫Colaphellus bowringi Baly是十字花科蔬菜的一种重要害虫,以成虫在土壤中滞育越冬和越夏。本研究通过解剖大猿叶虫非滞育成虫,观察描述了雌雄成虫内生殖系统的结构特点,并绘制了雌雄成虫内生殖系统模式图。对滞育初期、滞育期间、滞育解除后未取食和滞育解除后取食成虫解剖显示,滞育初期,雌雄成虫内生殖系统几乎与羽化初期成虫一致。滞育期间雄成虫的附腺和射精管亦不发达,滞育解除后未取食雄成虫的附腺膨大且粗于输精管,射精管略膨大,取食雄成虫的射精管呈不透明淡桔黄色。滞育期间雌成虫的卵巢小,略大于滞育初期卵巢,少数卵巢小管的基部可见具卵黄原沉淀的卵粒。滞育解除后未供食雌成虫卵巢明显膨大可见大量成形卵粒,部分卵粒可见卵黄原沉淀;取食雌成虫的卵巢膨大,可见大量成熟卵粒,侧输卵管和总输卵管中可见待产的卵。作者认为,大猿叶虫成虫在滞育期间能够缓慢发育,部分滞育前积累的代谢物质被用于滞育后发育,但只有经过取食,成虫才能正常交配和产卵。     

The effect of mating frequency and mating pattern on female reproductive fitness in cabbage beetle,Colaphellus bowringi

It is generally thought that females can receive more of the material benefits from males by increasing mating frequency and polyandry can lead to greater reproductive success. The cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), is a highly promiscuous species, in which females or males can readily mate repeatedly with a given partner or multiple partners at a very high frequency. In the present study, the effect of mating frequency (number of matings) and mating pattern (polyandry vs. monogamy) on female reproductive fitness was investigated by measuring fecundity, fertility, and female longevity. The results indicated that increased female mating frequency with the same male did not result in variation in lifetime fecundity, but significantly increased fertility and decreased female longevity. Moreover, five copulations were sufficient to acquire maximal reproductive potential. Female lifetime fecundity also did not differ between polyandrous and monogamous treatments. However, monogamous females exhibited a significant increase in fertility and significant prolongation of longevity compared with polyandrous females, further demonstrating that monogamy is superior to polyandry in this beetle.     

影响大猿叶虫交配持续时间的因素

为了探明影响大猿叶虫Colaphellus bowringi Baly交配持续时间的因素,在室内系统观察了一天中的交配起始时间、日龄、交配经历、体重、雄虫密度和温度对交配持续时间的影响.结果表明：一天中不同时间交配的成虫,其交配持续时间有显著差异,在上午8:00的交配持续时间最长,随着时间的推移,交配持续时间逐渐缩短,两者之间呈负相关.不同日龄成虫的交配持续时间有显著差异,且与成虫日龄呈明显的正相关.当雄虫与有交配经历的雌虫交配,其交配持续时间明显长于与未交配的雌虫交配,但雄虫的交配经历对交配持续时间没有影响.交配持续时间随着雌、雄体重的增加而延长,但没有显著差异.在雄虫高密度条件下的交配持续时间明显短于低密度下的交配持续时间.在22℃时成虫的交配持续时间最长,其次是25℃,28℃下最短, 温度间的交配持续时间有明显差异.这些结果表明在自然条件下大猿叶虫的交配持续时间受到许多因素的影响.     

Effects of thermoperiods on diapause induction in the cabbage beetle, Colaphellus bowringi (Coleoptera: Chrysomelidae)

Abstract. The effects of thermoperiods on diapause induction in continuous darkness or under a 12 : 12 h LD photoperiod were investigated in the cabbage beetle, Colaphellus bowringi Baly, a typical short‐day species. The diapause response curves both at different constant temperatures and at the thermocycle of format CT x: (24 ? x) h (16 : 28 °C) under continuously dark rearing conditions showed that the incidence of diapause depended mainly on whether or not the mean temperature was ≤20 °C or >20 °C. If the mean temperature was ≤20 °C, all individuals entered diapause; if >20 °C, the incidence of diapause declined gradually with increasing mean temperatures. The thermocycle (CT 12 : 12 h) with a series of different cryophases (8–22 °C) and thermophases (24–32 °C) under continuous darkness demonstrated a cryophase response threshold temperature of approximately 19 °C and a thermophase response threshold temperature of approximately 31 °C. Thermoperiodic amplitude (temperature difference between cryophase and thermophase) was shown to have a significant influence on diapause induction at the mean temperatures of 22, 23 and 24 °C, but not at ≥25 °C. Thermoperiodic responses under LD 12 : 12 h clearly showed that the incidence of diapause was influenced strongly by the photophase temperature. The thermoperiod under LD 12 : 12 h induced a much lower incidence of diapause than the thermoperiod with the same temperature in continuous darkness. The ecological significance of thermoperiodic induction of diapause in this species is discussed.     

温度对大猿叶虫成虫繁殖和繁殖后滞育的影响

大猿叶虫Colaphellus bowringi Baly是十字花科蔬菜上的重要害虫。在田间,该虫的繁殖期出现在春季和秋季,以成虫在土中越夏和越冬。本试验在光周期L∶D=16∶8条件下,系统调查了不同温度(20、25和28℃)对大猿叶虫非滞育成虫繁殖的影响。结果表明:每雌平均日产卵量在28℃最大(51.24±2.90)粒,显著高于25℃和20℃的(39.57±2.86)粒和(31.53±2.02)粒,但由于28℃下的产卵持续时间明显短,导致了总产卵量在这3种温度间没有显著差异;28℃下的卵孵化率低于20℃和25℃,但没有显著差异;随着温度的升高,成虫寿命逐渐缩短,20℃与28℃间的雌虫寿命存在显著差异,20℃与25℃,28℃的雄虫寿命存在显著差异;在20、25、28℃下,均有30%的个体经历一段时间繁殖后进入滞育,其滞育前的产卵持续时间随着温度的升高而明显缩短,总产卵量也随着温度的升高而下降。     

Mating behavior of the cabbage beetle, Colaphellus bowringi (Coleoptera: Chrysomelidae)

Abstract  Circadian mating rhythms, mating frequency, mating duration, and the effect of mating duration on fecundity and fertility in the cabbage beetle, Colaphellus bowringi were investigated in the laboratory. Mating occurred throughout the 24-h cycle but the majority of copulations occurred in the photophase with two apparent peaks, one at 8:00 and another at 16:00. Mating frequency observations for 10 consecutive days indicated that pre-mating period of C. bowringi was about 4 days, and pairs mated an average of 5 times per day and an average of 40 times during the first 10 days. There was a negative correlation between mating frequency and mating duration during the consecutive mating. The mean duration of the first copulation (136.24 ± 4.62 min) was significantly longer than those of the second (57.87 ± 2.03 min), third (53.05 ± 2.05 min) and fourth copulation (30.86 ± 2.98 min). Fecundity showed a slight increase with increasing mating duration but no significant difference among treatments. However, fertility was significantly influenced by the mating duration in this species. Mating of 20-min duration did not produce viable eggs. The mean percentage of fertile eggs with completed mating duration (204.43 ± 18.96 min, 56.75% fertile eggs) was significantly higher than those with 60 min (39.55%) and 30 min (17.91%) mating duration, suggesting that the longer mating duration might be associated with transfer of more sperm that are used to increase the fertility of eggs.     

Effects of diapause duration on future reproduction in the cabbage beetle,Colaphellus bowringi: positive or negative?

Abstract.  Cabbage beetles, Colaphellus bowringi , undergoing an imaginal summer and winter diapause in the soil, show a great difference in diapause duration (from several months to more than 3 years) under natural conditions. The effects of diapause duration on future reproduction in the beetle are investigated at 25 °C with an LD 14 : 10 h photoperiod and under natural conditions. The fecundity of postdiapause adults with a short diapause of 5 months and nondiapause adults is similar, showing that a short diapause has no affect on reproduction, whereas the longevity of postdiapause adults with a short diapause of 5 months is significantly shorter than nondiapause adults, showing that a short diapause has a negative affect on longevity. The mean total egg production per female and longevities of postdiapause adults with long diapause periods of 16, 22, 29 and 34 months are similar to nondiapause adults, but the mean daily egg production per female is significantly higher than nondiapause adults, showing that extended diapause has a positive effect on postdiapause reproduction. The offspring of postdiapause parents require a relatively shorter time for egg development compared with the offspring of nondiapause parents, showing that diapause has a positive effect on their offspring's performance. However, there are no significant differences among offspring performance in terms of survival, adult longevity, mean egg production per female and mean daily egg production per female.     

Life history responses of the cabbage beetle Colaphellus bowringi to temperature change

Temperature is considered one of the most important mediators of phenotypic plasticity in ectotherms. Here, we investigated life history traits of the cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), at a wide range of temperatures (16, 19, 22, 24, 26 and 28°C). The larval and pupal times were significantly decreased with increasing rearing temperature and growth rate was positively correlated with temperature. However, the relationship between body size and rearing temperature in C. bowringi did not follow the temperature–size rule; both males and females reached the highest body weight at 19°C. Females were significantly larger than males at all temperatures. Male pupae lost significantly more weight at metamorphosis compared to females. However, diapausing males gained significantly higher weight after feeding compared to diapausing females at higher temperatures of 22, 24, 26 and 28°C. Body weight tended to decrease with increasing rearing temperature, whereas sexual size dimorphism (SSD) tended to increase with increasing rearing temperature; thus, Rensch's rule is upheld. The degree to which SSD changed with temperature varied with different development stages. SSD was lowest in pupae, highest in newly emerged adults and intermediate in diapausing adults.     

大猿叶虫卵的颜色与其后代滞育率和繁殖力的关系

大猿叶虫Colaphellus bowringi Baly卵的颜色有浅黄色和深红色2种类型。调查卵的颜色与其后代滞育的发生和繁殖力的关系,结果表明,在25℃、28℃,不同光周期L∶D(11∶13,12∶12,13∶11和15∶9)下,浅黄色卵和深红色卵的后代滞育率没有显著差异;但在L14∶D10下,浅黄色卵的后代滞育率(25℃为58.6%;28℃为33.9%)显著高于深色卵的后代(25℃为45.4%;28℃为11.2%)。卵的颜色与其后代成虫的产卵前期和成虫寿命没有显著相关,但浅黄色卵后代的平均产卵量和平均日产卵量(1663粒和58.6粒)均显著高于深红色卵的后代(1176粒和45.1粒)。     

光周期和温度对大猿叶虫滞育后成虫生物学特性的影响

大猿叶虫Colaphellus bowringiBaly是我国十字花科蔬菜上的一种重要害虫,以成虫在土壤中越冬和越夏,滞育发生主要受温度和光周期调节。本文就光周期和温度对滞育后成虫生物学特性的影响进行了研究。结果表明,在25℃下,光周期（L14∶D10和L12∶D12）对成虫滞育后的存活率、寿命、日平均产卵量、总产卵量均无显著影响。在长光照（L14∶D10）下,温度18℃、22℃和25℃）对滞育后成虫的生物学特性具有明显的影响;随温度的降低,总产卵量呈下降趋势,平均产卵量显著降低,雌虫滞育后寿命显著延长,18℃下少数个体能被诱导再次滞育。     

Host plant mediation of diapause induction in the cabbage beetle, Colaphellus bowringi Baly （Coleoptera： Chrysomelidae）

The cabbage beetle, Colaphellus bowringiBaly （Coleoptera： Chrysomelidae）, is a serious pest of crucifers in China, undergoing an imaginal summer and winter diapause in the soil. The effects of host plants on diapause incidence were tested in the beetle. The ratio of adults entering diapause was significantly low when they fed on the mature leaves of Chinese cabbage Shanghaiqin （Brassica chinesis var communis） than those feeding on Chinese cabbage Suzhouqin （Brassica chinesis var communis）, radish （Raphanus sativus var longipinnatus） and stem mustard （Brassicajuncea vat tumida） at 25℃ combined with 13：11 （L： D） h. Fewer adults entered diapause on young leaves compared to physiologically aged and mature radish leaves at 25℃ combined with 13：11 （L： D） h. The effect of host plant species on diapause induction was also evident under continuously dark rearing conditions or at different photoperiods. These experimental results demonstrate that host plant mediation of diapause induction exists in the cabbage beetle. However, at temperatures ≤20℃ or photoperiods of 16：8 （L： D） h combined with 25℃, all individuals entered diapause regardless of the host plants, indicating that the effects of host plants on diapause induction could be expressed only within a limited range of temperatures and photoperiods.     

大猿叶虫生活史的研究

大猿叶虫Colaphellus bowringi是江西山区十字花科蔬菜上的重要食叶害虫,以成虫在土中越夏和越冬。由于成虫滞育期的差异,该虫的化性显示了明显的种内变异。有些个体隔年繁殖;有些个体是一化性的,仅在春季或秋季繁殖1代;有些个体是二化性的,春季和秋季各繁殖1代;有些个体是多化性的,春季1代,秋季2～3代。因此,大猿叶虫在田间一年可发生4代。在春季,滞育成虫于2月底至4月初陆续出土繁殖;在秋季,滞育成虫于8月中旬至10月初陆续出土繁殖。春季羽化的成虫于4月底至6月上旬陆续入土越夏,秋季羽化的成虫于9月中旬至12月底陆续入土越冬。大猿叶虫一生能交配多次,大多数雌虫的产卵期超过1个月,最长达67天。平均每雌产卵量：春季世代为644粒,秋季世代为963粒,最高达1.950粒。各虫态的发育历期：在15～30℃间,卵为13.78～3.14天,幼虫为22.83～6.95天,蛹为12.10～3.18天。发育阈值温度：卵为10.7℃,幼虫为8.8℃,蛹为9.6℃。非滞育成虫的寿命约为1～2个月,滞育成虫的寿命为5～38个月。滞育成虫均入土蛰伏,在土中的蛰伏深度为9～31 cm 。     

大猿叶虫滞育品系和非滞育品系交配行为的比较

大猿叶虫Colaphellus bowringi Blay是一种重要的十字花科蔬菜害虫,以成虫越冬和越夏。明确大猿叶虫非滞育品系生殖力下降的现象是否与交配行为的改变相关,将促进该非滞育品系在滞育遗传、滞育调控分子机制研究中的应用。本研究在25℃和L∶D=12∶12 h条件下,测定了大猿叶虫滞育品系和非滞育品系的交配节律、交配持续时间和交配频次。结果表明,大猿叶虫滞育品系和非滞育品系交配高峰均出现在光期开始后2 h,非滞育品系的交配比率显著低于滞育品系。滞育品系和非滞育品系第1次交配持续时间无显著差异,分别为122.7±7.4 min和100.1±10.2 min;但非滞育品系的总交配持续时间显著低于滞育品系。非滞育品系的交配频次为2.2±0.3次,显著低于滞育品系的3.8±0.5次。滞育品系和非滞育品系交配行为的比较发现,大猿叶虫非滞育品系生殖力降低与其交配比率下降、交配持续时间缩短和交配频次减少显著相关,本研究为进一步利用非滞育品系为材料解析滞育遗传、滞育调控机制奠定了基础。     

大猿叶虫夏滞育的诱导：基于定量的光周期反应

为了探明大猿叶虫Colaphellus bowringi Baly夏滞育诱导的光周期时间测量特性, 我们通过室内实验系统比较了该虫在25℃、 不同长光照条件下,夏滞育的发生以及诱导50%个体进入夏滞育所需求的光 暗循环数。结果表明：不同长光照诱导的夏滞育比率有显著差异, 其中15 h或16 h光照诱导的滞育比率最高, 短于或长于这两个光照其滞育率均明显下降。在不同光 暗循环实验中, 14 h诱导的滞育比率均低于50%, 诱导50%个体进入夏滞育所需求的光 暗循环数在L15∶D9, L16∶D8, L17∶ D7和L18∶D6分别为2.61, 3.72, 4.64和5.92 d, 处理间存在显著差异。这些结果提示该虫夏滞育的诱导是基于定量的光周期反应。     

温度对大猿叶虫发育、存活和繁殖的影响

在光周期14L:10D、75%RH的条件下研究不同温度对大猿叶虫Colaphellus bowringi Baly生长发育、存活率和繁殖的影响。结果表明,卵、幼虫和蛹的发育起点温度分别为11·86,11·17和10·15℃,有效积温分别为61·6,110·8和65·3日·度;22℃时卵的孵化率和蛹的羽化率最高,分别达96·04%和94·26%;滞育后成虫寿命雌虫普遍长于雄虫,25℃下的产卵期11～34d,每雌产卵量平均861粒,最高达1527粒。     

Female age: Male preference and reproductive success in primates

A survey of the literature on male sexual preferences according to the age of the female partner supported the hypothesis that adult males should prefer older females of higher parity. Reproductive success was also surveyed by age and parity, and younger females of lower parity were universally found to have lower success than older females of higher parity. This appears to be true for either age or parity independently, as well as in combination. These findings are discussed and applied to human societies, where similar trends would be expected.     

Parental effects: physiological age, mating pattern, and diapause duration on diapause incidence of progeny in the cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae)

The cabbage beetle, Colaphellus bowringi Baly, is a short-day species that has both a facultative summer and winter adult diapause. In this study, we systematically investigated the effects of parental physiological age, mating pattern (including mating frequency and delayed mating), and diapause duration on the diapause incidence of progeny in the beetle. As parents aged, progeny diapause incidences showed considerable inter-parental variability and an abnormally varying pattern, but their predicting line was approximate to horizontal when reared under constant conditions. There was significant difference in progeny diapause incidence between multiple-mating and once-mating treatments. Higher diapause incidences in the progeny were observed in delayed-mating treatments. Progeny diapause incidences increased as diapause duration in the parental generation increased, and hibernating individuals produced more diapausing progeny than aestivating ones. Reciprocal cross tests between the post-diapause adults and non-diapause adults with no diapause history showed that the effect of parental diapause duration on the progeny diapause incidence was determined by both mothers and fathers but the mothers appeared to have a stronger effect than the fathers. Our results suggest that the parental age effect in Colaphellus bowringi might not be simply age-dependent, but may be controlled by the interaction of multiple factors, including physiological, environmental, behavioral and genetic factors. We discuss the ecological and physiological significance of these findings.     

Inheritance of the photoperiodic response controlling imaginal summer diapause in the cabbage beetle, Colaphellus bowringi

Adults of the cabbage beetle Colaphellus bowringi display a summer diapause in response to the exposure of their larvae to long photoperiods. In the present study, the inheritance of the photoperiodic response controlling summer diapause in C. bowringi by crossing a high diapause strain (D strain) with a laboratory selected nondiapause strain (N strain) was investigated under different photoperiods at 22, 25 and 28 °C. The beetles in both reciprocal crosses and backcrosses showed a clear short-day response for the induction of diapause at all temperatures, similar to that of the D strain, suggesting that photoperiodic response of this beetle is heritable. The diapause incidences in the progeny from all the crosses under LD 15:9 or LD 12:12 at 25 °C suggest that genetic and genetic-environmental interactions are involved in diapause induction. The incidence of diapauses in F₁ progeny was significantly lower than that in the D × D strain but significantly higher than that in the N × N strain, indicating that the diapause capability is inherited in an incomplete dominant manner. The incidence of diapause was greater among the offspring of hybrid females when those females had a D strain mother or grandmother rather than a N strain mother or grandmother, indicating that maternal effects on diapause induction are stronger than paternal effects. The laboratory selected nondiapause strain also showed a short-day photoperiodic response at a low temperature of 22 °C, indicating that the photoperiodic photoreceptor and photoperiodic clock still function in the nondiapause strain.     

Diapause induction and clock mechanism in the cabbage beetle, Colaphellus bowringi (Coleoptera: Chrysomelidae)

Photoperiodic control of diapause induction was investigated in the short-day species, Colaphellus bowringi, which enters summer and winter diapause as adult in the soil. Photoperiodic responses at 25 and 28 degrees C revealed a critical night length between 10 and 12 h; night lengths > or =12 h prevented diapause, whereas night lengths <12 h induced summer diapause in different degree. Experiments using non-24-h light-dark cycles showed that the duration of scotophase played an essential role in the determination of diapause. Night-interruption experiments with T=24 h showed that diapause was effectively induced by a 2-h light pulse in most scotophases; whereas day-interruption experiments by a 2-h dark break had a little effect on the incidence of diapause. The experiments of alternating short-night cycles (LD 16:8) and long-night cycles (LD 12:12) during the sensitive larval period showed that the information of short nights as well as long nights could be accumulated. Nanda-Hamner experiments showed three declining peaks of diapause at 24 h circadian intervals. Bünsow experiments showed two very weak peaks for diapause induction, one being 8 h after lights-off, and another 8 h before lights-on, but it did not show peaks of diapause at a 24 h interval. These results suggest that the circadian oscillatory system constitutes a part of the photoperiodic clock of this beetle but plays a limited role in its photoperiodic time measurement.