Glycerol accumulation in developmentally arrested pupae of Papilio machaon obtained by brain removal

Brains were removed from non-diapausing pupae of Papilio machaon within 24 hr after pupation. The developmentally arrested pupae were examined for glycerol metabolism by comparison with naturally diapausing pupae. The decerebrate non-diapausing pupae accumulated appreciable amounts of glycerol (13.0 mg/ml haemolymph) during exposure to 5°C for 2 months. However, this amount was considerably lower than that accumulated in the naturally diapausing pupae (24.5 mg/ml) under the same conditions. A difference was also observed in the initial level of the glycogen in the fat body. The naturally diapausing pupae contained about 105 mg/g glycogen, while the decerebrate non-diapausing pupae contained only about 70 mg/g, one day after pupation.     

Water and carbohydrate levels at different developmental stages and dynamics in hibernating pupae of Pieris melete (Lepidoptera: Pieridae)

For insight into the physiological indicators of diapause in Pieris melete, water and carbohydrate (glycogen and trehalose) levels were measured under both natural and laboratory conditions. The highest water content (3.71–3.79 mg/mg dry weight) was found in larvae and developing pupae, which was substantially higher than in diapausing pupae (2.59 mg/mg dry weight). Water content was almost stable during diapause, except for individuals approaching diapause termination (3.43–3.58 mg/mg dry weight). The total carbohydrate level was significantly higher in pre‐pupae (47.41 μg/mg) compared to larvae (22.80 μg/mg) and developing pupae (21.48 μg/mg). The highest level of trehalose was detected in winter diapausing pupae, and no trehalose was found in larvae or developing pupae. Levels of glycogen were highest in pre‐pupae and lowest in diapausing pupae. Levels of total carbohydrate decreased as diapause proceeded, and no significant changes were found in trehalose levels for diapausing pupae under natural conditions or treated for 60–90 days at 5°C. Pupae treated at 20°C for 60–90 days had significantly lower levels of trehalose than those treated for 30 days. Glycogen content was relatively stable at 5°C, but increased after treatment under natural conditions and 20°C for more than 60 days. These results suggest that the dynamics of water and carbohydrate levels are potential physiological diapause indicators, which show metabolic differences between trehalose and glycogen during diapause development.     

大斑芫菁滞育幼虫在滞育不同阶段体内糖类和醇类含量的变化

在以卵滞育的昆虫中昆虫滞育时的生理代谢特点已经得到了大量研究。本文对以末龄幼虫（5龄）滞育的大斑芫菁Mylabris phalerate（Pallas）在不同滞育阶段体内糖类和醇类代谢的特征进行了研究。结果表明： 滞育个体血淋巴中的海藻糖含量高于非滞育个体,且随滞育时间的加大逐渐升高,滞育5个月时达到最大值,为5.61 μmol/mL。糖原的含量随滞育的进程逐渐减少,滞育初期（0.5个月）为0.72 mg/mL,到滞育末期（5个月）时仅为0.1 mg/mL。滞育个体脂肪体中的海藻糖含量都高于非滞育个体,滞育1个月时为非滞育个体的3倍,至滞育末期时达非滞育个体的5倍,为2.5 μmol/g脂肪体。糖原含量总体变化趋势是随滞育时间的加大逐渐减少,滞育早期和中期都高于非滞育个体。在滞育过程中血淋巴积累的小分子多元醇主要为甘油,其次是山梨醇;而在脂肪体中主要为甘油,其次是甘露醇,少量积累山梨醇：表明大斑芫菁滞育幼虫主要积累的是海藻糖和一些小分子多元醇。滞育幼虫在准备滞育时储备了大量糖原,这些糖原可能为滞育期间海藻糖、山梨醇和甘油的代谢提供了原料。     

Fat body stimulation of ecdysone synthesis in Heliothis zea

Prothoracic glands of Heliothis zea pupae require both a humoral factor and prothoracicotropic hormone (PTTH) to synthesize ecdysone. The humoral factor is absent when pupae are maintained at diapause-sustaining temperatures. Thus, pupae remain in diapause despite the release of PTTH at or before larval-pupal ecdysis.Tissue implantation experiments revealed that a diapause-terminating factor is present in the fat body of non-diapausing pupae. Other tissue implantation experiments showed that, when diapausing pupae were transferred from 19 to 27°C, diapause-terminating activity appeared first in the fat body and then the fat body into the haemolymph. HPLC separation of the haemolymph and fat body fractions followed by bioassay demonstrated that fractions containing diapause-terminating activity eluted from both tissues within 28–30 min. These results suggest that the factors found in the fat body and haemolymph may be the same compound.Evidence from ecdysone radioimmunoassay experiments ruled out the possibility that the diapauseterminating activity was due to either free or conjugated ecdysteroids. Corresponding in vitro experiments in which the prothoracic glands were cultured with brain extracts versus fat body and haemolymph fractions also indicated that the haemolymph/fat body factor was not PTTH.     

Comparison of cold hardiness and sugar content between diapausing and nondiapausing pupae of the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae)

Abstract. To understand overwintering of the cotton boll worm Helicoverpa armigera, cold hardiness and sugar content are compared between diapausing and nondiapausing pupae. Diapausing and nondiapausing pupae reared at 20 °C under short and long photoperiods are acclimatized with a reduction of 5 °C per 5 days to 0 °C. When the acclimation temperature reaches 0 °C, the survival of diapausing pupae is assessed. The survival gradually decreases as the period of treatment progresses and approximately half survive for 112 days. However, nondiapausing pupae survive only 14 days after exposure to 0 °C. The surpercooling points of nondiapausing, diapausing and acclimatized pupae are approximately −17 °C. The major sugars contained in pupae are trehalose and glucose. Even though trehalose contents in diapausing pupae (initial level: 0.6 mg 100 mg⁻¹ fresh weight) increase significantly during cold acclimation and continue increasing until 58 days after exposure to 0 °C (maximum level: 1.8 mg 100 mg⁻¹), glucose is maintained at low levels (0.02 mg 100 mg⁻¹) for 56 days at 0 °C. However, glucose contents increase (maximum level: 0.8 mg 100 mg⁻¹) with decreasing contents of trehalose 84 days after exposure to 0 °C. Glycogen content gradually decreases during cold acclimation. When nondiapausing pupae are acclimatized with a reduction of 5 °C per 5 days to 5 °C from the beginning of pupation until the eyespots move, trehalose content increases (maximum level: 1.0 mg 100 mg⁻¹). Glucose contents in nondiapausing pupae increase before eclosion (0.09 mg 100 mg⁻¹). From these results, diapausing pupae of H. armigera can overwinter in regions where average winter temperatures are higher than 0 °C, but nondiapausing pupae cannot.     

The biochemical effects of potassium chloride on the silkworm, （ Bombyx mori L.）

The supplementation with 50, 100 and 150μg/mL potassium chloride to the fifth instar larvae of the silkworm Bombyx mori on fat body glycogen, protein, total lipids and haemolymph protein and trehalose were analyzed. The fat body glycogen and protein and haemolymph protein were increased significantly in all the treated groups; whereas fat body total lipids increased only in 100 and 150μg/mL and haemolymph trehalose increased only in 150μg/mL potassium chloride-treated groups when compared with those of the corresponding parameters of the carrier controls.     

Cold hardiness and sugar content in photo‐insensitive individuals of the cotton bollworm Helicoverpa armigera

Abstract A proportion of Helicoverpa armigera collected from fields in Okayama Prefecture (Western Japan; 34.6°N, 134.1°E) does not enter diapause when reared under a short days at 20 °C during the larval stages. However, diapause in such photo‐insensitive individuals can be induced when they are reared at moderately low temperatures, such as 15 °C, regardless of photoperiod. To determine whether such photo‐insensitive individuals can survive overwintering in fields, the present study compares the cold hardiness and sugar content between nondiapausing and diapausing pupae of photo‐insensitive individuals selected over several generations at 20 °C under a short day photoperiod (LD 10 : 14 h). Diapausing and nondiapausing pupae are obtained under the short days by rearing at 15 and 20 °C, respectively, during larval and pupal stages. These pupae are stepwise acclimated at a reduction of 5 °C every 5 days to 0 °C. Maximum survival periods of nondiapausing and diapausing pupae at 0 °C are approximately 30 and 90 days, respectively. Trehalose content in diapausing pupae increases, reaches a maximum level (1.95 mg 100 mg^?1 in males and 2.1 mg 100 mg^?1 in females) 28 days after exposure to 0 °C and then decreases. On the other hand, glucose content in diapausing pupae increases (maximum level: 0.32 mg 100 mg^?1 in males and 0.21 mg 100 mg^?1 in females) with decreasing trehalose content 42 days after exposure to 0°C. The decrease in trehalose content and the increase in glucose content may be linked to termination of diapause in H. armigera. These results suggest that, in Japan, the photo‐insensitive individuals can only survive in the mild winters of southern regions, and not in the severe winters of northern regions.     

The in vitro biosynthesis and secretion of glycerol by larval fat bodies of chilled Ostrinia nubilalis

Fat bodies from diapausing fifth-instar larvae of Ostrinia nubilalis were incubated in vitro at 5 or 23°C in Grace's medium and the glycerol contents of the organ and incubation medium determined. Fat bodies from diapausing larvae chilled 3 weeks at 5°C secreted glycerol into the medium at 5°C at a net rate of approx. 0.75 nmol/mg fat body dry wt/h for at least 96 h while the tissue levels remained essentially constant. Depending upon the experiment, from 6 to 15 times more glycerol was produced in 24 h at 5°C by these fat bodies than by those taken from diapausing unchilled larvae and incubated at either 5 or 23°C. A minimal chilling period of 10–12 days was recognized as necessary for chilled larval fat bodies to demonstrate rates of glycerol synthesis greater than those of unchilled larvae and the lag showed a temporal correlation with changes in haemolymph glycerol concentrations. These results suggest that this response to chilling by O. nubilalis is relatively slow. While incubation, at 23°C, of fat bodies from previously chilled larvae did not result in cessation of glycerol secretion, the rate of its appearance in the culture medium decreased during the 24-h incubation period. Although the ability of chilled fifth-instar larvae to accumulate glycerol is not dependent upon the diapause state results show that clearance of glycerol from the haemolymph by rewarmed O. nubilalis is related to diapause intensity.     

Study on the physiology of diapause, cold hardiness and supercooling point of overwintering pupae of the pistachio fruit hull borer, Arimania comaroffi

The pistachio fruit hull borer, Arimania comaroffi (Ragonot) (Lepidoptera: Pyralidae), is a key pest of pistachio orchards in Iran. This pest passes the winter as diapausing pupae. In this study, some physiological changes in relation to environmental temperature were investigated in field collected pupae. The relationship between supercooling point, cold hardiness and physiological changes of a wild population of this pest was also investigated. The glycogen content decreased with decrease in environmental temperature. Decrease in glycogen content was proportional to increase in total body sugar, trehalose, myo-inositol and sorbitol contents. In January with mean ambient temperature of 5.4°C, glycogen (5 mg/g fresh body weight) content was at the lowest level whereas total body sugar (10.3 mg/g fresh body weight), trehalose (8.6 mg/g fresh body weight), myo-inositol (5.3 mg/g fresh body weight) and sorbitol (2.6 mg/g fresh body weight) were at the highest levels. Total body sugar, trehalose, myo-inositol and sorbitol contents increased as mean temperature decreased from 22.7°C in October to 5.4°C in January. Total body lipid decreased during overwintering and reached to the lowest level at the end of March. Supercooling points were decreased from October to January and reached to the lowest level (-16°C) in January with minimum ambient temperature of -10°C. Survival at low temperature after 24 h was also greatest in January with 72% survival at -10°C, 39% survival at -15°C and 0% survival at -20°C. Increase in temperature from February onward, was proportional with increase in supercooling points and decrease in survival rate. Regardless of sampling date, all pupae died after 24 h at -20°C, whereas none pupae died after 24 h at -5°C. This indicates that this insect is freeze-intolerant.     

EFFECT OF A HIGH TEMPERATURE ON VITELLOGE-NESIS IN THE JAPANESE OAK SILKWORM,ANTHERAEA YAMAMAI (LEPIDOPTERA: SATURNIIDAE)*

Abstract The effect of a high temperature, i. e. 32°C. on vitellogenesis of the Japanese oak silkworm, Antheraea yamamai (Lepidoptera: Saturniidae) was markedly significant. Its extent was dependent on the development stage of the silkworm exposed to 32°C. When exposed to 32 °C since the 1st day after cocooning, titres of both vitellogenin (Vg) and soluble proteins in the fat body and hemolymph of mature larvae were evidently lower than those at 26°C. When pupae were maintained at 32°C since the 1st day after pupation. the titres of Vg in the fat body showed no significant difference from those at 26°C, but those in the hemolymph and the titres of vitellin (Vt) in the ovary mostly were obviously lower in contrast to those at 26°C. While exposed to 32°C since the 6th day after pupation, at most instance the tires of Vg both in the fat body and hemolymph were not markedly different from those at 26°C, and those of Vt in the ovary were significantly higher than those at 26°C, In addition, the changes in the titres of soluble proteins in the fat body and hemolymph as well as the ovary were monitored when pupae were maintained at 32°C since the 1st or 6th day after pupation. It is recommended that both mature larvae and pupae at cocooning stage and earlier pupal stage should not be exposed to 32°C when the silkworm is reared for egg raising.     

Comparative studies of oxidative enzyme systems in epidermis and fat body of diapausing and non-diapausing silkmoths

The activities of four oxidative enzyme systems, including NADH oxidase, succinate-cytochrome c reductase, NADH-cytochrome c reductase, and cytochrome c oxidase, were compared in mitochondrial-microsomal preparations from wing epidermis and fat body of diapausing Samia cynthia pupae, presumptively non-diapausing S. cynthia ricini pupae which were caused to diapause by removal of the brain, and non-diapausing S. cynthia ricini during the pupal and pharate adult period. In diapausing pupae the activities of all enzyme systems were low and presented a profile similar to that previously reported for the Cecropia silkmoth. By contrast, in non-diapausing individuals the activities showed substantially higher levels, and an essentially unchanging pattern from just after the larval-pupal ecdysis through most of adult development. These events are functionally correlated with the patterns of biosynthetic activity in diapausing and non-diapausing silkmoths and are discussed in relation to the endocrine control of diapause and development.     

Characteristics of larvae of the southwestern corn borer, Diatraea grandiosella, obtained from populations present in tropical and temperate regions

A comparative study was undertaken of southwestern corn borers (Diatraea grandiosella) collected from south central Mexico (19°N latitude) and southeast Missouri (37°N latitude). All the life stages of the Mexican insects were found to be larger or heavier, or both, than were those of the Missouri insects. Mexican larvae grew at a higher rate and attained a significantly heavier body weight than did Missouri larvae. Although both Mexican and Missouri larvae underwent ecdyses during diapause, Mexican larvae ecdysed more frequently than did the Missouri larvae at 23°C light:dark 12 h:12 h. Larvae that ecdysed most frequently did not necessarily spend longer in diapause. Electrophoresis of the fat body and haemolymph proteins of Mexican and Missouri larvae revealed similar patterns. The fat body of diapausing Mexican larvae contained substantial amounts of a diapause-associated protein which has been characterized previously from the fat body of Missouri Larvae. Double immunodiffusion confirmed that the diapause-associated protein of the Mexican larvae was identical to that present in the Missouri larvae. Smaller amounts of this protein appear to be present in the fat body of diapausing Mexican larvae than are present in that of diapausing Missouri larvae.     

Overexpression and functional characterization of an Aspergillus niger phytase in the fat body of transgenic silkworm,Bombyx mori

In a previous study, we isolated 1,119 bp of upstream promoter sequence from Bmlp3, a gene encoding a member of the silkworm 30 K storage protein family, and demonstrated that it was sufficient to direct fat body-specific expression of a reporter gene in a transgenic silkworm, thus highlighting the potential use of this promoter for both functional genomics research and biotechnology applications. To test whether the Bmlp3 promoter can be used to produce recombinant proteins in the fat body of silkworm pupae, we generated a transgenic line of Bombyx mori which harbors a codon-optimized Aspergillus niger phytase gene (phyA) under the control of the Bmlp3 promoter. Here we show that the Bmlp3 promoter drives high levels of phyA expression in the fat body, and that the recombinant phyA protein is highly active (99.05 and 54.80 U/g in fat body extracts and fresh pupa, respectively). We also show that the recombinant phyA has two optimum pH ranges (1.5–2.0 and 5.5–6.0), and two optimum temperatures (55 and 37 °C). The activity of recombinant phyA was lost after high-temperature drying, but treating with boiling water was less harmful, its residual activity was approximately 84 % of the level observed in untreated samples. These results offer an opportunity not only for better utilization of large amounts of silkworm pupae generated during silk production, but also provide a novel method for mass production of low-cost recombinant phytase using transgenic silkworms.     

Haemolymph ecdysone concentrations in Hyalophora cecropia pupae,dauer pupae and adults

Haemolymph ecdysone concentrations were determined by radioimmunoassay in diapausing pupae, pharate adults, adults, and chilled dauer pupae. The concentration in diapausing pupae after 6 months chilling (5.35 pg/μl) increased dramatically after 3 days at 27°C (>200 pg/μl) and then decreased to low levels in adult females (1.63 pg/μl). In adult males ecdysone was undetectable in all except one animal. Dauer pupae showed a decrease from 6.1 to 1.7 pg/μl 1 day after being transferred from 6 to 27°C. Over a 3-day period the value increased to 3.19 pg/μl and remained constant for more than a year. These results suggest that diapausing pupae with and without brain neurosecretory cells maintain a low concentration of ecdysone in the haemolymph.     

Diapause phenomena in non-diapausing last instar larvae,pupae, and pharate adults of the Colorado beetle

From the first day of the last (fourth) larval instar no trace of juvenile hormone (JH) can be detected in the haemolymph by Galleria bioassay. Three specific diapause proteins, which are also found in diapausing adults, appear in the haemolymph. These proteins disappear towards the end of the pupal stage. Study of the ultrastructure of the fat body revealed the formation from lysosomes of proteinaceous bodies which are also characteristic for adult diapause. The behaviour of last instar larvae and pupae resembles that of prediapausing and diapausing adults respectively. Injection of synthetic JH delays the appearance of the diapause proteins in the haemolymph and of proteinaceous bodies in the fat body for 2 to 3 days. The absence of JH seems to trigger off these diapause phenomena.     

Carbohydrate metabolism during the pupal molt of the tobacco hornworm,Manduca sexta

During the pupal molt of the tobacco hornworm, Manduca sexta, the percentage of active fat body glycogen phosphorylase increased from 5–10 to 20%, but only for a period of 5 h prior to the molt. From the time of the appearance of two sclerotized dorsal bars to the time of the molt, the concentration of total hemolymph carbohydrates doubled to 100 mM trehalose. Initially, the glucose level was high (16 mM) when compared with feeding larvae (approximately 1 mM) but decreased to zero just prior to the molt. The amount of cuticular chitosan decreased from approximately 100 mg to 10 mg at pupation; the exuvia contained approximately 7 mg. While the levels of total lipids in hemolymph were not affected, the lipid content of the fat body decreased significantly prior to the molt but increased sharply thereafter. Fat body glycogen phosphorylase in pharate pupae and pupae of M. sexta was substantially activated by the Manduca adipokinetic peptide hormone, which in pharate pupae, produced the same response at 2 and 20 pmol per insect as in ligated larval abdomens. In pupae the response was clearly reduced. Using chilling to stimulate glycogen phosphorylase, it was found that the enzyme in pharate pupae and pupae responded both in vivo and in vitro as in ligated abdomens of larvae. Thus, a transition to the adult response seems to occur during the pupal and pharate adult development. © 1995 Wiley-Liss, Inc.     

Physiology of diapause and cold hardiness in overwintering pupae of the apple leaf miner Phyllonorycter ringoniella in Japan

Abstract The apple leaf miner Phyllonorycter ringoniella (Matsumura) (Lepidoptera: Gracillariidae) overwinters as a diapausing pupa. The diapause rate reaches 100% in early October. Diapause intensity decreases gradually from early October and diapause terminates in early February. The fresh body weight of diapausing pupae is 1.6 times that of non-diapausing pupae. The main cryoprotectant in P. ringoniella pupae is trehalose. Three stages are distinguishable as indicated by the correlations between diapause intensity, levels of cold hardiness and the trehalose content: diapause induction occurred in October, diapause development from November to December, and post-diapause quiescence from January to April. During diapause induction, the pupae accumulate low levels of trehalose and do not survive exposure to −15 °C. During diapause development, the pupae gradually accumulate more trehalose and show some ability to survive exposure to −15 °C, but not to −20 °C. During post-diapause quiescence, the pupae accumulate relatively more trehalose and cold hardiness fully develops, but decreases quickly in April. The trehalose content in pupae sampled in December is unaffected by acclimation temperatures in the range 0–30 °C, but decreases in pupae sampled in March after acclimation at temperatures from 5 to 15 °C. These results suggest that overwintering pupae of P. ringoniella have the ability to accumulate trehalose and develop a high level of cold hardiness during diapause development.     

13C and 31P NMR studies on sugar metabolism in Bombyx mori and Philosamia cynthia ricini larvae

Studies were made on ¹³C and ³¹P NMR in larvae of two species of silkworm, Bombyx mori and Philosamia cynthia ricini, in vivo as well as in vitro to determine the pathways of glucose utilization, especially those to amino acids as components of silk fibroin. Results showed that the ¹³C of [1-¹³C]glucose administered orally into 5th instar larvae of both species was incorporated into glucose-1-phosphate, glucose-6-phosphate and trehalose. Serine, glutamate, glutamine, citrate, malate, trehalose and sorbitol-6-phosphate were detected in the hemolymphs of these larvae as metabolites of [1-¹³C]glucose. Two days after [1-¹³C]glucose administration, labeled alanine, glycine, serine, urea, glycogen, trehalose and glycerol were clearly detected in Bombyx larvae. Starvation caused rapid consumption of administered [1-¹³C]glucose with very little accumulation of ¹³C in glycogen or trehalose. In the in vivo³¹P NMR spectra of Bombyx larvae, ATP, arginine phosphate, sorbitol-6-phosphate, uridine diphosphoglucose, phosphoenolpyruvate and inorganic phosphate were detected with some sugar phosphates, such as glucose-1-phosphate and glucose-6-phosphate. During starvation, the intensity of the signal of inorganic phosphate increased and those of sugar phosphate other than sorbitol-6-phosphate decreased, but these changes were reversed by oral administration of glucose.     

Temperature-sensitive mechanism regulating diapause in Heliothis zea

Pupal diapause in Heliothis zea is regulated by a temperature-sensitive mechanism which prevents ecdysone production despite the release of prothoracicotropic hormone. To determine how this mechanism functioned, donor prothoracic glands were implanted into prothoracic gland-ablated hosts to test their ability to produce ecdysone in a diapause-sustaining temperature of 19°C. Results of these experiments ruled out the possibility that ecdysis production was regulated by the nervous system or by a mechanism intrinsic to the prothoracic glands, and suggested that a humoral factor was required for diapause termination.Haemolymph injection experiments supported this humoral factor hypothesis, i.e. haemolymph from non-diapausing donor pupae terminated diapause in hosts maintained at 19°C, whereas haemolymph from diapausing donor pupae had no such effect. These findings indicate that the temperature-sensitive mechanism regulating H. zea diapause functions by controlling the availability of a humoral factor necessary for ecdysone production by the prothoracic glands.     

取食Bt棉与玉米的越冬代棉铃虫抗寒性特征比较

【目的】针对由Bt棉花和玉米构成的华北农田景观, 探讨不同寄主作物对棉铃虫 Helicoverpa armigera 越冬抗寒的影响。【方法】2013年在河北廊坊科研中试基地的作物田中将棉铃虫幼虫接到Bt棉和玉米上, 比较取食不同作物后棉铃虫的化蛹率、存活率和越冬蛹羽化率;在室内控制条件下分别用棉蕾和鲜玉米粒饲喂棉铃虫幼虫。 测定滞育蛹和非滞育蛹的鲜重、干重、以及脂肪、糖原和低分子物质含量等指标, 比较取食不同作物后棉铃虫的抗寒能力。【结果】取食玉米的棉铃虫滞育蛹干重（117.5 mg）、脂肪含量（457.2 μg/mg DW）以及海藻糖浓度（86.45 μg/g）均显著高于取食Bt棉的棉铃虫滞育蛹干重（56.6 mg）、脂肪含量（239.6 μg/mg DW）以及海藻糖浓度（13.87 μg/g）;取食玉米的棉铃虫冰点(-10.2℃)显著低于取食Bt棉的棉铃虫冰点(-6.5℃)。【结论】结果表明取食玉米更加有利于棉铃虫越冬。 据历史数据, 近年来玉米种植面积不断增加, 这将提高棉铃虫成功越冬比率, 对棉铃虫种群扩张起到促进作用, 因此注重玉米上棉铃虫的防治尤为重要。