Development and survival of the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Yponomeutidae) as a function of temperature: effect on the number of generations in tropical and subtropical regions

The diamondback moth, Plutella xylostella (L.), is the most important pest of brassicaceous crops worldwide. Since temperature is the major abiotic factor influencing insect development and thermal requirements may vary among insect populations, it is important to know the effect of temperature on development and survival of a subtropical strain of P. xylostella. Development and survival of the diamondback moth was evaluated under seven constant temperatures ranging from 10°C to 35°C. Development was completed between 10°C and 32.5°C, but at 35°C all individuals died in the larval stage. Data were fitted to one linear and five nonlinear models. Considering as criteria the goodness of fit and the ability to estimate parameters of biological significance, the models Briere-1 and Briere-2 were the most adequate to describe the relationship between temperature and development of P. xylostella. The linear model demonstrated that P. xylostella required 312.5 degree-days above a lower threshold of 6.3°C to complete development. The degree-day model showed that the number of diamondback moth generations in the tropical region of Brazil is nearly twice the number in the subtropical region of the country. This result explains, at least in part, the higher population levels of this species in the tropical region of Brazil, and also demonstrates that P. xylostella is tolerant to a wide range of temperatures (6.1-32.5°C). Therefore, temperature cannot be considered a limiting factor for the occurrence of diamondback moth throughout the year in most regions of Brazil.     

家用调料植物粗提物防治小菜蛾的研究

以安全性和简易性为前提的庭园害虫防治成为休闲生活不可或缺的技术。本文以花椒Zanthoxylum bungeamtm、八角茴香lUicium veturn、辣椒Capsicum frutescens等日用品为材料,研究了不同种类、浓度和作用时间对小菜蛾Plutella xylostella这一重要庭园害虫的拒食、触杀及产卵忌避效果。拒食试验表明,供试3种植物对小菜蛾有显著拒食作用,拒食率从大到小依次为八角茴香（0．94）、花椒（0．86）、辣椒（0．76）;拒食率随处理时问长短变化的差异不显著,但随提取液浓度的升高而升高。触杀试验表明,供试3种植物对小菜蛾有显著触杀作用,24h内死亡率从大到小依次为花椒（45．6％）、八角茴香（35．6％）、辣椒（23．3％）,均显著高于对照处理的1．1％;触杀死亡率随虫龄的增加而降低。产卵忌避试验表明,3种植物对小菜蛾成虫产卵忌避作用由大到小依次为辣椒、八角茴香、花椒。综合本研究所得结果来看,八角茴香对小菜蛾的控制效果最好。所选材料容易获得,对小菜蛾忌避作用明显,在庭园小菜蛾防治中具有潜在的应用价值。     

Host plant and population determine the fitness costs of resistance to Bacillus thuringiensis

Novel adaptations often cause pleiotropic reductions in fitness. Under optimal conditions individual organisms may be able to compensate for, or reduce, these fitness costs. Declining environmental quality may therefore lead to larger costs. We investigated whether reduced plant quality would increase the fitness costs associated with resistance to Bacillus thuringiensis in two populations of the diamondback moth Plutella xylostella. We also measured the rate of decline in resistance on two host-plant (Brassica) species for one insect population (Karak). Population X plant species interactions determined the fitness costs in this study. Poor plant quality increased the fitness costs in terms of development time for both populations. However, fitness costs seen in larval survival did not always increase as plant quality declined. Both the fitness and the stability experiment indicated that fitness costs were higher on the most suitable plant for one population. Theoretically, if the fitness cost of a mutation interacts additively with environmental factors, the relative fitness of resistant insects will decrease with environmental quality. However, multiplicative costs do not necessarily increase with declining quality and may be harder to detect when fitness parameters are more subject to variation in poorer environments.     

Effects of host plant and genetic background on the fitness costs of resistance to Bacillus thuringiensis

Novel resistance to pathogens and pesticides is commonly associated with a fitness cost. However, measurements of the fitness costs of insecticide resistance have used diverse methods to control for genetic background and rarely assess the effects of environmental variation. Here, we explored how genetic background interacts with resource quality to affect the expression of the fitness costs associated with resistance. We used a serially backcrossed line of the diamondback moth, Plutella xylostella, resistant to the biopesticide Bacillus thuringiensis, to estimate the costs of resistance for insects feeding on two Brassica species. We found that fitness costs increased on the better-defended Brassica oleracea cultivars. These data were included in two meta-analyses of fitness cost experiments that used standardized protocols (and a common resistant insect stock) but which varied in the methodology used to control for the effects of genetic background. The meta-analysis confirmed that fitness costs were higher on the low-quality host (B. oleracea); and experimental methodology did not influence estimates of fitness costs on that plant species. In contrast, fitness costs were heterogeneous in the Brassica pekinensis studies: fitness costs in genetically homogenized lines were significantly higher than in studies using revertant insects. We hypothesize that fitness modifiers can moderate fitness costs on high-quality plants but may not affect fitness when resource quality is low.     

cDNA sequence analysis of ribosomal protein S13 gene in Plutella xylostella （Lepidoptera： Plutellidae）

Ribosomal protein S 13 gene has been cloned and analyzed in many organisms,but there are few documents relating to insects. In this communication, the full-length cDNA sequence of ribosomal protein S 13 gene in the diamondback moth, Plutella xylostella(Lepidoptera: Plutellidae), was determined by using PCR amplification technique. The features of the ribosomal protein S 13 gene sequence were analyzed and the deduced amino acids sequence was compared with those from other insects. The results of multi-alignment of the amino acid sequences between the diamondback moth and other insect species revealed that this gene sequence is highly conserved in insects. Based on maximum likelihood method, a phylogenetic tree was constructed from 10 different species using PHYLIP software. It showed that nematode is one separate lineage and the five insect speciesbe long to another lineage, whereas those species higher than insects form the third one. The pattern of this phylogenetic tree evidently represented the evolution of different species.     

Production and regulation of extracellular proteases from the entomopathogenic fungus Isaria fumosoroseus (Cordycipitaceae; Hypocreales) in the presence of diamondback moth,Plutella xylostella cuticle

Extracellular proteases of entomopathogenic fungi have been implicated as components of the insect infection process. To elucidate their role in the infection process, it is essential to characterize these enzymes. During the current studies synthesis and regulation of extracellular proteases (Pr1 and Pr2) by Isaria fumosoroseus (Cordycipitaceae; Hypocreales) isolate IF28.2 were investigated as a function of carbon source (with special reference to diamondback moth cuticle), temperature and pH. The highest level of Pr1 and Pr2 activity were found in the supernatants from 1% glucose plus 1% diamondback moth cuticle (18.83±1.25 and 12.44±1.36 U/mg per h for Pr1 and Pr2, respectively). Maximum Pr1 production by the depressed mycelia was observed from the supernatants having diamondback moth cuticle as a nutrient source, whereas depressed mycelia showed maximum Pr2 activity from the cultures having chitin as the basic nutrient. The optimum pH for Pr1 and Pr2 activity was 8 while 35°C was the best temperature for protease production.     

Could the intrinsic rate of increase represent the fitness in terrestrial ectotherms?

The intrinsic rate of increase (r_m) has been considered as an important indicator of fitness in terrestrial ectotherms since long. It is actually an equivalent to the instantaneous growth rate of the exponential equation for describing the density-independent population growth. In terrestrial ectotherms, r_m has been demonstrated to be temperature-dependent. The temperature at which r_m was maximal, was considered to be the “optimal” temperature for fitness in Amarasekare and Savage (2012), but this definition needs further analysis. Only r_m cannot provide thorough representation of fitness. Because body size can affect the competitive abilities in many terrestrial ectotherms, both population size and body size should be considered in measuring the fitness of ectotherms. The rule of “bigger is better” requires relatively low temperature to increase in body size, whereas relatively high temperature is required for a rapid increase in population size. Thus, there is presumably a trade-off in temperature for adjusting individual body size and population size to achieve maximum fitness. We hypothesized that this temperature could be reflected by the intrinsic optimum temperature for developmental rate in the Sharpe–Schoolfield–Ikemoto model, and it led to a temperature estimate around 20 °C. However, the traditional viewpoint based on the temperature corresponding to the maximal intrinsic rate of increase provides a temperature estimate around 30 °C. This study suggests that a low temperature around 20 °C might authentically represent the optimal ambient temperature for fitness in terrestrial ectotherms. It implies that thermal biologists who are interested in the effect of temperature on the fitness in terrestrial ectotherms should pay more attention to their performance at low temperature rather than high temperature.     

Estimating the economic cost of one of the world's major insect pests, Plutella xylostella (Lepidoptera: Plutellidae): just how long is a piece of string?

Since 1993, the annual worldwide cost of diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), control has been routinely quoted to be US$1 billion. This estimate requires updating and incorporation of yield losses to reflect current total costs of the pest to the world economy. We present an analysis that estimates what the present costs are likely to be based on a set of necessary, but reasoned, assumptions. We use an existing climate driven model for diamondback moth distribution and abundance, the Food and Agriculture Organization country Brassica crop production data and various management scenarios to bracket the cost estimates. The "length of the string" is somewhere between US$1.3 billion and US$2.3 billion based on management costs. However, if residual pest damage is included then the cost estimates will be even higher; a conservative estimate of 5% diamondback moth-induced yield loss to all crops adds another US$2.7 billion to the total costs associated with the pest. A conservative estimate of total costs associated with diamondback moth management is thus US$4 billion-US$5 billion. The lower bound represents rational decision making by pest managers based on diamondback moth abundance driven by climate only. The upper estimate is due to the more normal practice of weekly insecticide application to vegetable crops and the assumption that canola (Brassica napus L.) is treated with insecticide at least once during the crop cycle. Readers can decide for themselves what the real cost is likely to be because we provide country data for further interpretation. Our analysis suggests that greater efforts at implementation of even basic integrated pest management would reduce insecticide inputs considerably, reducing negative environmental impacts and saving many hundreds of millions of dollars annually.     

Insect resistance to Bacillus thuringiensis: uniform or diverse

Resistance to the insecticidal proteins produced by the soil bacterium Bacillus thuringiensis (Bt) has been documented in more than a dozen species of insect. Nearly all of these cases have been produced primarily by selection in the laboratory, but one pest, the diamondback moth (Plutella xylostella), has evolved resistance in open-field populations. Insect resistance to Bt has immediate and widespread significance because of increasing reliance on Bt toxins in genetically engineered crops and conventional sprays. Furthermore, intense interest in Bt provides an opportunity to examine the extent to which evolutionary pathways to resistance vary among and within species of insect. One mode of resistance to Bt is characterized by more than 500-fold resistance to at least one Cry1A toxin, recessive inheritance, little or no cross-resistance to Cry1C, and reduced binding of at least one Cry1A toxin. Analysis of resistance to Bt in the diamondback moth and two other species of moth suggests that although this particular mode of resistance may be the most common, it is not the only means by which insects can attain resistance to Bt.     

用甘蓝苗连续饲养小菜蛾的技术

室内实验结果表明 :在 ( 2 5± 1 )℃、相对湿度 5 0 %～ 70 %、光周期 1 6L∶8D条件下 ,用甘蓝苗(BrassicanapusL .)饲养小菜蛾PlutellaxylostellaL .可取得较好的饲养结果 ,连续饲养多代种群未出现退化现象 ,能够得到大量发育整齐的供试虫源。在低温 ( 4℃ )条件下 ,卵冷藏 7d ,孵化率 77 8% ,蛹冷藏 1 5d化蛹率 86 9% ,冷藏后卵和蛹的发育等与各自的对照基本一致 ,可以较好地调控小菜蛾发育进度 ,便于取得虫龄基本一致的虫源。     

Developmental parameters and seasonal phenology of Calepitrimerus vitis (Acari: Eriophyidae) in wine grapes of western Oregon

Developmental parameters of protogyne Calepitrimerus vitis (Nalepa) (Acari: Eriophyidae) were determined at 12, 15, 17, 22, 25, 28, 31, and 34 °C to better understand seasonal activity, population growth, and ultimately more effectively manage pest mites in wine grapes. Net reproductive rate (R(o)) was greater than zero at all temperatures with the maximum R(o) (9.72) at 25 °C. The lowest estimated R(o) (0.001) occurred at 34 °C. There was a gradual decrease in mean generation time (T) as temperatures increased from 17 to 31 °C. The shortest and longest generation time was recorded at 31 °C (T = 5.5 d) and 17 °C (T = 17.5 d). Rates of natural increase were lowest at 17°C (0.035) and increased with increasing temperatures, respectively. The peak rate of natural increase value (0.141) was at 25 °C. Estimations for minimum and maximum developmental thresholds were 10.51 and 39.19 °C, respectively, while the optimum developmental temperature was 26.9 °C. The thermal constant for egg to adult development was estimated at 87.7DD. The highest fecundity was observed at 25 °C. These parameters indicated that mites begin feeding at the onset of shoot growth when tissue is most susceptible in spring. Historical weather data showed that vines are in this susceptible growth stage for longer periods in the cool Willamette Valley compared with warmer Umpqua and Applegate/Rogue Valley regions. Estimation of degree-days indicated when deutogyne mites move to overwintering refuge sites. Degree-day accumulations indicated up to 14 generations per growing season.     

Site-Specific Cassette Exchange Systems in the Aedes aegypti Mosquito and the Plutella xylostella Moth

Genetically engineered insects are being evaluated as potential tools to decrease the economic and public health burden of mosquitoes and agricultural pest insects. Here we describe a new tool for the reliable and targeted genome manipulation of pest insects for research and field release using recombinase mediated cassette exchange (RMCE) mechanisms. We successfully demonstrated the established ΦC31-RMCE method in the yellow fever mosquito, Aedes aegypti, which is the first report of RMCE in mosquitoes. A new variant of this RMCE system, called iRMCE, combines the ΦC31-att integration system and Cre or FLP-mediated excision to remove extraneous sequences introduced as part of the site-specific integration process. Complete iRMCE was achieved in two important insect pests, Aedes aegypti and the diamondback moth, Plutella xylostella, demonstrating the transferability of the system across a wide phylogenetic range of insect pests.     

Effect of temperature on post-wintering development and total lipid content of alfalfa leafcutting bees

Temperature plays an important role in effective management of the alfalfa leafcutting bee [Megachile rotundata (F.); Megachilidae], the major commercial pollinator of seed alfalfa [Medicago sativa (L.); Fabaceae] in North America. To improve our understanding of threshold and optimum rearing temperatures of M. rotundata, we examined the effect of temperature on postwintering development by using a greater number of temperature treatments than applied in previous studies (19 versus eight or fewer) and analytical tools formulated to model nonlinear relationships between temperature and insect development rates. We also tested the hypothesis that rearing temperature influences adult body lipid content at emergence, which could affect adult survival, establishment and performance as a pollinator, and reproductive success. We found that the Lactin-2 and Briere-2 models provided the best fits to data and gave reasonable estimates of lower (16-18°C) and upper (36-39°C) developmental thresholds and optimum (33-34°C) rearing temperatures for maximizing development rate. Bees successfully emerged over a broad range of temperatures (22-35°C), but variation in development rate among individuals reared at the same temperature was lowest at 31-33°C. The optimum rearing temperature to maximize the proportion of body lipids in adults was 27-29°C. Our results are discussed in relation to previous findings and speak to the difficulties in designing practical rearing guidelines that simultaneously maximize development rate, survival, and adult condition, while synchronizing adult emergence with alfalfa bloom.     

TWO HERBIVORES AND CONSTRAINTS ON SELECTION FOR RESISTANCE IN BRASSICA RAPA

Although most plants experience herbivory by several insect species, there has been little empirical work directed toward understanding plant responses to these simultaneous selection pressures. In an experiment in which herbivory by flea beetles (Phyllotreta cruciferae) and diamondback moths (Plutella xylostella) was manipulated in a factorial design, I found that selection for resistance to these herbivores is not independent in Brassica rapa. Specifically, the effect of flea beetle damage on B. rapa fitness depends on the amount of diamondback moth damage a plant experiences: damage by these herbivores has a nonadditive effect on plant fitness. When diamondbacks are abundant, plants that sustain high levels of damage by flea beetles are favored by natural selection, but when diamondbacks are rare, a low level of damage by flea beetles is favored. However, resistance to the later-feeding diamondback moth is not affected by the presence or absence of damage by early-feeding flea beetles. Thus, there are no plant-mediated ecological interactions between these herbivores that affect the outcome of selection for resistance. Because these herbivores do not independently affect plant fitness, neither is likely to develop a pairwise coevolutionary relationship with its host. Instead, coevolution is diffuse.     

Thermodynamics constrains the evolution of insect population growth rates: "warmer is better"

Diverse biochemical and physiological adaptations enable different species of ectotherms to survive and reproduce in very different temperature regimes, but whether these adaptations fully compensate for the thermodynamically depressing effects of low temperature on rates of biological processes is debated. If such adaptations are fully compensatory, then temperature-dependent processes (e.g., digestion rate, population growth rate) of cold-adapted species will match those of warm-adapted species when each is measured at its own optimal temperature. Here we show that cold-adapted insect species have much lower maximum rates of population growth than do warm-adapted species, even when we control for phylogenetic relatedness. This pattern also holds when we use a structural-equation model to analyze alternative hypotheses that might otherwise explain this correlation. Thus, although physiological adaptations enable some insects to survive and reproduce at low temperatures, these adaptations do not overcome the "tyranny" of thermodynamics, at least for rates of population increase. Indeed, the sensitivity of population growth rates of insects to temperature is even greater than predicted by a recent thermodynamic model. Our findings suggest that adaptation to temperature inevitably alters the population dynamics of insects. This result has broad evolutionary and ecological consequences.     

Temperature-related fitness costs of resistance to spinosad in the diamondback moth, Plutella xylostella (Lepidoptera: Plutelidae)

Fitness costs associated with resistance genes expressed in the absence of insecticides affect the evolution of insecticide resistance and the outcome of resistance management programmes. However, measurements of fitness costs may not be straightforward as they vary with environmental conditions. The diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae), has developed resistance to spinosad, the first insecticide of the Naturalyte class, after only a few years of field application of this product. In this study, we compared the performance of two homogenous strains of P. xylostella, one susceptible (SS) and the other resistant (RR) to spinosad at an unfavourable, low natural temperature regime, a favourable median-fluctuating temperature regime and an unfavourable high-fluctuating temperature regime. The RR strain showed only marginal fitness cost at the median temperature regime. At the low temperature regime, the RR strain failed to produce any viable offspring, while the SS strain achieved positive population growth. At the high temperature regime, the RR strain showed a 33% decrease in intrinsic rate of increase compared to the SS strain. The results demonstrate that fitness costs of resistance to spinosad are temperature-dependent, increasing in scale at unfavourably low and high temperatures; costs were particularly high at low temperatures. Suggestions for designing effective management programmes are made to delay or avoid development of resistance to spinosad by P. xylostella under different temperature conditions.     

Genetic Structure and Demographic History Reveal Migration of the Diamondback Moth Plutella xylostella (Lepidoptera: Plutellidae) from the Southern to Northern Regions of China

The diamondback moth Plutella xylostella (Linnaeus) (Lepidoptera: Plutellidae) is one of the most destructive insect pests of cruciferous plants worldwide. Biological, ecological and genetic studies have indicated that this moth is migratory in many regions around the world. Although outbreaks of this pest occur annually in China and cause heavy damage, little is known concerning its migration. To better understand its migration pattern, we investigated the population genetic structure and demographic history of the diamondback moth by analyzing 27 geographical populations across China using four mitochondrial genes and nine microsatellite loci. The results showed that high haplotype diversity and low nucleotide diversity occurred in the diamondback moth populations, a finding that is typical for migratory species. No genetic differentiation among all populations and no correlation between genetic and geographical distance were found. However, pairwise analysis of the mitochondrial genes has indicated that populations from the southern region were more differentiated than those from the northern region. Gene flow analysis revealed that the effective number of migrants per generation into populations of the northern region is very high, whereas that into populations of the southern region is quite low. Neutrality testing, mismatch distribution and Bayesian Skyline Plot analyses based on mitochondrial genes all revealed that deviation from Hardy-Weinberg equilibrium and sudden expansion of the effective population size were present in populations from the northern region but not in those from the southern region. In conclusion, all our analyses strongly demonstrated that the diamondback moth migrates within China from the southern to northern regions with rare effective migration in the reverse direction. Our research provides a successful example of using population genetic approaches to resolve the seasonal migration of insects.     

四种十字花科蔬菜上小菜蛾自然种群连续世代生命表

利用作用因子生命表技术,组建芥菜、芥蓝、小白菜和菜心4种十字花科蔬菜上小菜蛾自然种群连续世代生命表,分析寄主植物和生物因子对小菜蛾种群数量的控制作用。结果表明：在4种十字花科蔬菜的一造菜上,小菜蛾均能完成两个世代。虽然小菜蛾在芥菜上的初始卵量最高,但害虫种群总增长倍数在芥蓝上最高,其次为小白菜,菜心和芥菜,分别为17．64、11．90、11．43和3．76。这说明尽管芥菜对小菜蛾成虫的产卵有一定的吸引作用,但不适合小菜蛾生长发育。芥蓝是最适宜小菜蛾种群增长的寄主。生物因子在小菜蛾自然种群控制中起着重要的作用,但是在不同种类十字花科蔬菜上,天敌类群对小菜蛾控制作用存在一定差异。除芥菜之外,寄生性天敌对芥蓝、菜心和小白菜上的小菜蛾种群控制作用最大,其次为“捕食及其它”,病原微生物的控制作用最小。“捕食及其它”对芥菜上小菜蛾种群的作用非常明显,如果排除此因子作用,小菜蛾种群两代后将增长126．03倍。该因子是导致芥菜小菜蛾自然种群增长趋势指数低的主要原因。因此在制定小菜蛾防治策略时,应考虑蔬菜的种类和布局,加大对芥蓝小菜蛾种群的防治力度;芥菜可作为一种诱杀植物种植,以吸引小菜蛾产卵,并集中防治。这些防治策略在小菜蛾综合治理中具有重要的实际意义。     

亚麻荠对小菜蛾幼虫取食和成虫行为反应的影响

亚麻荠是一种很少有害虫危害的油料作物。用室内生测和Y 型嗅觉仪研究了亚麻荠对小菜蛾Plutella xylostella幼虫取食和成虫行 为反应的影响。以甘蓝作对照,用亚麻荠叶片喂养的小菜蛾初孵幼虫3天后校正死亡率为79 .2%,显示了较强的致死作用;喂养小菜蛾3龄幼虫至化蛹,其存活率、化蛹率、蛹重及成 虫寿命都显著降低,表明亚麻荠对小菜蛾幼虫的生长发育有不利影响。在幼虫的取食选择实 验中,有甘蓝叶供选择时,小菜蛾幼虫不取食亚麻荠;在无可选择的情况下,小菜蛾幼虫也 取 食亚麻荠叶片,但取食量很小,与取食甘蓝叶的量相比,差异极显著。行为反应测试表明, 小菜蛾成虫对甘蓝和亚麻荠植株的挥发物都有明显的趋性反应,与对照（净化空气）相比, 差异极显著,而在甘蓝和亚麻荠之间无选择性。说明小菜蛾成虫对亚麻荠植株的挥发物具有 较强的定向反应。     

Diurnal temperature amplitude affects diamondback moth adult and next-generation traits

Temperature fluctuation is one of the most important factors in climate change, and research into its effects on organisms is increasing. Nonetheless, few studies have addressed its effects on insects and particularly on the parental generation. We therefore simulated temperature amplitudes (± 0, 4, 6, 8, 10°C) for a mean environmental temperature of 25°C and studied their immediate and maternal effects on a global pest, the diamondback moth Plutella xylostella (Lepidoptera: Plutellidae), focusing on adult longevity and fecundity and egg development and survivorship in the next generation. Relative to constant temperature, moderate amplitudes (4 and 6°C) did not affect the adults, whereas 8 and 10°C amplitudes hindered adult longevity and fecundity, but significantly increased the proportional of early fecundity after each treatment. Temperature amplitude 10°C inhibited development of Day 2 and Day 3 laid eggs. These findings show that temperature amplitude affects both the maternal and offspring phenotype and indicate that the influence of temperature amplitude on diamondback moth population models requires further research.