Differences in adipokinetic response of Pyrrhocoris apterus (Heteroptera) in relation to wing dimorphism and diapause

Three experimental groups of adult females (reproductive and diapausing brachypters, and macropters with reproductive arrest) of Pyrrhocoris apterus (Linnaeus) (Heteroptera: Pyrrhocoridae) from a temperate population were analysed for their adipokinetic responses. The adipokinetic response, expressed as an increase of haemolymph lipids after injection of adipokinetic hormone from Locusta migratoria (Lom-AKH-I), was assessed in relation to age, wing dimorphism and type of reproductive arrest. Two pmols of Lom-AKH-I were used for determination of adipokinetic responses. The increase of haemolymph lipids in all experimental groups of females induced by this dose of the hormone was comparable with that induced by crude extract of the bug’s own corpora cardiaca. The level of adipokinetic response after injection of 2 pmol of Lom-AKH-I was significantly higher in macropterous and diapausing brachypterous females than in reproductive brachypterous females. However, significantly higher contents of haemolymph lipids in control macropterous females than those found in the control reproductive and diapausing brachypterous females of the corresponding age revealed wing-morph-related differences in lipid metabolism. The observed wing morph- and diapause-related differences in the content of haemolymph lipids and adipokinetic response, respectively, are discussed in relation to the different roles of two wing morphs in the life history of this heteropteran.     

Feeding, drinking and digestive enzyme activities in long- and short-day females of Pyrrhocoris apterus (Heteroptera)

Abstract. .Reproducing females of the firebug, Pyrrhocoris apterus (Linnaeus) (Heteroptera: Pyrrhocoridae), reared under long-day (LD 18:6h) photocycle (long-day females), and diapausing females reared under short-day (LD 12:12h) conditions (short-day females) were analysed for temporal patterns of feeding, drinking and gut digestive enzyme activities. In long-day females the highest activities of feeding and drinking were found at the beginning of each reproductive cycle. Significant correlation between feeding and drinking activities in both reproducing long-day females and short-day females during pre-diapause state was found. In pre-diapausing short-day females the cyclical pattern of feeding and drinking was maintained despite the inhibition of reproduction. After entering the diapause their feeding activity became very low. Analysis of five gut enzymes revealed that short-day females differ significantly from long-day females in higher protease, amylase and esterase activities, in case of lipase only at certain times of their adult life; no significant difference was found in the aminopeptidase activity. Positive correlations between intensity of feeding and gut enzyme activities were observed in short-day females. Using several protease inhibitors most protease activity in the gut of P.apterus was identified as cysteine and aspartyl proteases, and only a small part of the activity could be caused by some serine proteases.     

Locomotor activity in adult Pyrrhocoris apterus (Heteroptera) in relation to sex, physiological status and wing dimorphism

The pattern of locomotor (walking) activity was studied in adult males and females of short‐winged (brachypterous) and long‐winged (macropterous) morph of the flightless bug Pyrrhocoris apterus (Linnaeus) (Heteroptera: Pyrrhocoridae) under constant laboratory conditions. Walking activity was measured with a computerized video system and analysed with respect to sex, physiological status (reproduction, diapause and reproductive arrest of non‐diapause type) and wing dimorphism of the bugs. The largest duration was observed in the macropterous females with reproductive arrest of non‐diapause type (average 6 h per day) and the shortest duration in diapausing brachypterous females and males (average less than 2 h per day). This was reflected also in the overall time spent by walking during the first 14 days after imaginal ecdysis. The time spent by walking significantly increased in the macropterous morph as the bugs aged, whereas in diapausing brachypters the time spent by walking decreased with age. No linear relationship between walking activity and age was found in reproductive brachypterous morph. The bugs of all experimental groups moved mostly during the photophase and were almost inactive during the scotophase. Thus, walking activity in P. apterus is diurnal, irrespective of the wing morph, physiological status, sex and age. Contrary to the macropterous morph, where the locomotor activity of females during photophase was significantly higher than in males, no significant differences were found between the locomotor activities of brachypterous males and females. The observed differences in locomotor activity are discussed in relation to different roles of two wing morphs in the life history of this heteropteran.     

始红蝽越冬聚集行为对其能量代谢的影响

聚集行为是动物适应不良环境的一种生态对策。始红蝽Pyrrhocoris apterus在成虫滞育越冬期间具有显著的聚集行为。采用LI-6400型光合仪测定了5种聚集度(分别用1头、5头、10头、20头和50头的群体大小表示)下始红蝽的呼吸量,通过呼吸量计算得到各聚集度始红蝽的呼吸速率(rate of respiration,Rr)及能量代谢速率(rate of metabolism,Rm),旨在明确聚集行为是否能够对始红蝽越冬时的能量代谢产生影响。结果表明： 不同聚集度间始红蝽的Rm值存在显著性差异; 并且随聚集度的增加,Rm值与聚集度呈显著负相关,Rm值依次为0.052,0.044,0.041,0.037和0.033 W/g。结果说明聚集行为可以有效降低始红蝽新陈代谢速率,有利于种群成功越冬。     

How climate change affects the occurrence of a second generation in the univoltine Pyrrhocoris apterus (Heteroptera: Pyrrhocoridae)

1. Understanding the conditions that allow for the occurrence of an additional generation in populations that are usually univoltine is important under the present climate warming. In temperate areas, a second generation is enabled through the emergence of a time window that opens when first-generation individuals are ready to reproduce and closes when second-generation individuals cannot complete development before the onset of winter. 2. The conditions that limit the width of this window were studied in Pyrrhocoris apterus (Heteroptera: Pyrrhocoridae), a ground-inhabiting heteropteran overwintering in facultative adult diapause, whose populations in Central Europe have typically been univoltine until the 1980s. 3. The frequency of females of the first generation that started to lay eggs decreased from 70% in June to zero in early August, but oviposition of these females continued until the end of August. Using thermal constants for egg–adult development and temperature data, this study found that the development of most second-generation individuals could only be completed before the start of winter if hastened through behavioural thermoregulation. 4. Consequences of temperature increase on the width of the thermal window were calculated. Increasing temperature causes the time window to open earlier and close later by accelerating maturation of first-generation females and improving conditions for maturing of the second-generation individuals in late summer and autumn. 5. Climate warming will create conditions that facilitate the occurrence of a second generation in a year in typically univoltine populations of this species.     

The effect of allatectomy and photoperiod on the supercooling point in Pyrrhocoris apterus adults

Abstract Adults of Pyrrhocoris apterus (L.) (Heteroptera: Pyrrhocoridae) reared at 26°C had a considerably lower supercooling point (SCP) in diapause-inducing (LD 12:12 h) than in diapause-inhibiting (LD 18:6 h) photoperiodic regimes. Exposure of the insects to diapause-inducing acclimation conditions (LD 8:16 h and a temperature of 20°C during photophase and 5°C during scotophase) for 2–3 weeks had very little effect on the SCP, irrespective of the rearing photoperiod. Allatectomized adults kept continuously at LD 18:6 h and 26°C had high SCP similar to intact or sham-operated insects. In contrast to non-allatectomized insects, the SCP of allatectomized insects decreased after an exposure of from 2 to 3 weeks to diapause-inducing acclimation conditions (see above) almost to the level found in the intact diapausing insects. The relationship between the decrease of SCP and ‘diapause syndrome’ (de Wilde, 1970) is discussed.     

Feeding behaviour, digestive physiology and lipid content in macropterous females of Pyrrhocoris apterus (L.) (Heteroptera: Pyrrhocoridae)

Abstract. Macropterous females of Pyrrhocoris apterus (L.) reared under short-day conditions (LD 12:12 h) were analysed for temporal patterns of feeding and drinking behaviour, activities of digestive enzymes in the gut, and lipid and glycogen content in the haemolymph and fat body. Peaks of drinking activity were recorded at days 3, 7 and 10 during the first 14 days after imaginal ecdysis. Feeding activity peaked on the third day, ceasing completely after the fourth day of adult life. Esterase, protease, amylase and aminopeptidase activities exhibited the highest overall activity in the first days after imaginal emergence; then enzyme activities decreased. In the fat body, the content of lipids was highest on day 5, then a decrease of about 40% was observed at day 14; the amount of glycogen was highest on day 1 at 11 μg of glucose equivalents/mg of fat body, then decreased to 2 μg at day 14 after the imaginal moult. In the haemolymph, the lipid content rose until day 8 when it reached almost 0.3 μmol/μl; at day 14 the value was slightly lower. The association of fasting with reproductive arrest in macropterous females of P. apterus, accompanied by a decrease in digestive enzyme activities and a mobilization of lipid reserves from the fat body, was demonstrated.     

The influence of diet on the duration and thermal sensitivity of development in the linden bug Pyrrhocoris apterus L. (Heteroptera: Pyrrhocoridae)

Temperature and nutrition are crucial environmental variables that determine rates of growth and development in insects. However, the simultaneous effect of these factors on life‐history traits is rarely addressed. In the present study, the influence of two diets (linden fruit and sunflower seeds) on the duration of immature stages and thermal reaction norms for development is tested in the bug Pyrrhocoris apterus L. (Heteroptera: Pyrrhocoridae). Eggs and larvae are reared at five constant temperatures (20, 22, 24, 26 and 28 °C) under an LD 20 : 4 h photocycle. Development rates deviate from linearity in the studied thermal range, especially in larvae; therefore, a nonlinear (power‐law) approximation is also attempted. Parental diet causes no change in thermal reaction norms for egg development. However, the progeny of sunflower‐fed bugs are more variable in terms of their development time, suggesting a transgenerational effect. Larval mortality rates increase in cooler conditions and are always higher on sunflower seeds. This is accompanied by more variable, less temperature‐dependent and generally slower larval development. A review of previously published case studies on temperature–diet interactions in the control of insect development leads to two general conclusions. First, there are two approaches for assessing the temperature‐dependent development in insects: one based on the concept of the sum of degree‐days and the other based on the concept of reaction norm. Despite an obvious non‐exclusiveness, the two approaches appear to have developed in isolation from each other. Second, three principal patterns of temperature–diet interactions can be recognized. The pattern found in P. apterus (the direct effects of diet are stronger at higher temperatures and much weaker or absent at lower temperatures) appears to be the most widespread.     

Analysis of pteridines in Pyrrhocoris apterus (L.) (heteroptera,pyrrhocoridae) during development and in body-color mutants

By using thin-layer chromatography (TLC) and high-performance liquid chromatography (HPLC), five different pteridines have been quantified in extracts from Pyrrhocoris apterus: neopterin, isoxanthopterin, isoxantholumazine (violapterin), 7-methylxanthopterin, and erythropterin. Biopterin was also detected using HPLC. Pteridines have been analyzed separately in bodies and eyes of the wild type regarding developmental stage and sex. The pteridine content in both bodies and eyes increased from nymphs to 2-day-old adults. After this period, the concentration of pteridines in the eyes of adults remained approximately constant, while in the bodies isoxantholumazine, 7-methylxanthopterin, and isoxanthopterin kept increasing until 20 days. Considering the total amount of pteridines per insect, no qualitative or quantitative differences between males and females have been observed except a lower concentration of erythropterin in female bodies. But when pteridines are analyzed per unit body weight, a lower amount of pteridines is always detected in females due to its bigger size. A new body-color mutant, the yolk body mutant, that goes through developmental colour changes, was also analyzed during development, showing a general deficient accumulation of all pteridines, especially of 7-methylxanthopterin and isoxantholumazine in adult bodies. The effect on pteridine content of three other recessive (white, yellow, mosaic) and two dominant (Pale and Apricot) body-color mutations, has also been studied. A specific pteridine pattern different from the wild type was obtained for each mutant. © 1997 Wiley-Liss, Inc.     

Study of intrapopulational variability of duration and temperature norms of development of the linden bug Pyrrhocoris apterus (Heteroptera, Pyrrhocoridae)

The existence of significant variability in duration and temperature norms of development between families within insect populations has been shown for the first time. This variability is inferfamily and therefore has genetic ground. Revealed for the first time is the statistically significant positive correlation between the regression coefficient of the development rate for temperature and the temperature threshold for development of eggs and larvae from different families. The greater the slope of the regression line of the development rate for temperature, the higher the temperature threshold value in this particular family. These results demonstrate for the first time the existence of genetic co-variation between the regression coefficient and the temperature threshold within the insect populations. It is suggested that the source of the interpopulational and interspecies changes in the temperature reaction norms of the insect development might be the intrapopulational hereditary variability of the development duration, regression coefficient, and the development threshold, this variability being an object of natural selection. It was shown that in all studied families and populations the values of the linear regression coefficient of development rates for temperature in eggs of the linden bug Pyrrhocoris apterus were markedly and statistically significantly higher, while the temperature threshold values—lower as compared with the corresponding parameters in larvae. These results obviously are in contradiction with the concept of the “isomorphism of development rates” (Jarosik et al., 2002), according to which the development threshold for all life cycle stages of a species should be the same, whereas only slopes of the regression lines of the development rate for temperature can differ. For the first time the absence of genetic covariation has been shown between the temperature norms of development of different life cycle stages of the species—eggs and larvae. This means that the regression coefficient as well as the sum of the degree-days and the development threshold in eggs and larvae are inherited independently and therefore they can be independently changed in evolution in correspondence with specific environmental conditions, under which these life cycle stages take place.     

Effect of thermoperiod on diapause intensity in Pyrrhocoris apterus (Heteroptera Pyrrhocoridae)

The intensity of adult diapause in Pyrrhocoris apterus was measured in two series of experiments as the duration of pre-oviposition period at a constant temperature of 25 degrees C after transfer from short (12L:12D) to long day conditions (18L:6D). Higher diapause intensity was induced with a thermoperiod than at constant temperatures. After the induction throughout larval instars 3-5 and during 4 weeks of adult life at short days and a thermoperiod of 25/15 degrees C the pre-oviposition period was 30+/-4 and 26+/-3 days. After induction at constant 25 degrees C the pre-oviposition period was 22+/-3 and 23+/-4 days, while after induction at constant 20 degrees C it was 17+/-4 and 19+/-4 days. Induction at a lower constant temperature of 20 degrees C was thus followed by a less intense diapause than the induction at a higher constant temperature of 25 degrees C. These counterintuitive results are discussed. The oxygen consumption rate measured at experimental temperatures prior to transfer from short to long days was higher at thermoperiodic conditions than at constant temperatures and it was similar at constant 20 and 25 degrees C. Thus, the oxygen consumption rate measured prior to the transfer was highest (indication of the least intense diapause) in the insects that showed later, after the transfer to long days, the longest pre-oviposition period (indication of the most intense diapause). Within the first two days after transfer to constant 25 degrees C, oxygen consumption rate measured at 25 degrees C decreased in the thermoperiodic insects, while it transiently increased in insects from constant 20 degrees C. Two days and later after the transfer, oxygen consumption rate was similar in all groups. Cold hardiness was not correlated with diapause intensity. The low lethal temperature in diapausing insects was correlated with the night temperature during diapause induction.     

二化螟水稻种群与茭白种群光周期反应的比较

本文报道了用水稻和茭白分别饲养二化螟水稻种群和茭白种群的光周期反应。光周期反应曲线显示,用水稻饲养茭白种群或用茭白饲养水稻种群,无论是在短光照还是在长光照条件下,绝大多数幼虫被诱导进入滞育,丧失了各自原有的光周期反应特性,表明这两个种群已分化到仅适应其本身寄主的程度。用水稻饲养的茭白种群仅有5%～6%的个体化蛹,且其幼虫期较用茭白饲养的延长了15～18天;而用茭白饲养的水稻种群有30%～40%的个体化蛹,其幼虫期与水稻饲养的仅相差3～8天。茭白种群用水稻饲养时32日龄幼虫体重仅为茭白饲养的53.1%,而水稻种群用水稻饲养时32日龄幼虫体重为茭白饲养的79.5%。这些结果表明,茭白种群不适应取食水稻,而水稻种群对取食茭白则有一定的适应能力。根据这些结果,我们认为：（1）这两个种群已出现种下分化的迹象;（2）茭白种植不会对水稻田二化螟的发生产生大的影响。     

Some effects of low temperature on the photoperiodic clock in Drosophila triauraria

Abstract. Prior studies revealed that the photoperiodic clock in Drosophila triauraria operates during the scotophase and the processes measuring scotophase are composed of two phases: the A phase which lasts 6–7 h from the onset of the scotophase and the B phase which follows the A phase. In this study the photoperiodic clock in this species was analysed by low-temperature pulses. Low temperature (5C) or a low-temperature (2C) pulse of 3 or 4 h decreased the induction of diapause when applied during the scotophase, but not when applied within the photophase. The low-temperature pulse showed two maxima of the effect, 0–4 h and 8–10 h after the onset of the scotophase, approximately corresponding to the A and B phases, respectively. It is considered that low temperature retards the time measurement processes operating during these phases. In the period which was rather insensitive to the diapause-preventing action of low temperature, i.e. 5-7h after the onset of the scotophase, substances or stimuli accumulated during the A phase may be converted to information required for operation of the machinery in the B phase. Experiments using a combination of low-temperature and light pulses suggested that low-temperature decreases the sensitivity to light, especially in the B phase.     

Maternal temperature has different effects on the photoperiodic response and duration of larval diapause in blow fly ( Calliphora vicina ) strains collected at two latitudes

Abstract .The blow fly Calliphora vicina Robineau-Desvoidy (Diptera: Calliphoridae) has a wide distribution across northern and temperate Europe. It has a facultative, maternally-induced larval diapause in response to short days. The photoperiodic response, measured at 15 and 20°C, of two populations was compared. A southern population (originating at 51° N) was sensitive to temperature at all daylengths; the incidence of diapause was greatly reduced at 20°C compared with 15°C. The photoperiodic response of a northern population (from 65° N) was sensitive to temperature only in long days; in short days (< 14 h of light) the response of this strain was identical at each temperature.
Variation in parental photoperiod and temperature were found to affect the duration of larval diapause, indicating a role for maternal effects in diapause intensity as well as incidence. However, the between-strain variation was greater than that within strains, indicating qualitative differences in diapause response. These differences may arise from the ecological conditions at the points of origin of the two strains. The northern strain from the harsher climate has a more intense diapause that follows a relatively temperature-insensitive photoperiodic response. In contrast, the southern strain has a shallow diapause and its photoperiodic response may be overridden by the experience of concurrent high temperature.     

The role of circadian clock genes in the photoperiodic timer of the linden bug Pyrrhocoris apterus during the nymphal stage

Many insects survive seasonal adversities during diapause, a form of programmed developmental and metabolic arrest. Photoperiodically regulated entry into diapause allows multivoltine insect species to optimize the number of generations. The molecular mechanism of the photoperiodic timer is unknown in insects. In the present study, we take advantage of the robust reproductive diapause response in the linden bug Pyrrhocoris apterus and explore the fifth‐instar nymphal stage, which is the most photoperiod‐sensitive stage. The nymphs display daily changes in locomotor activity during short days; this differs from the activity observed during long days. We find evidence of cyclical expression of the circadian clock genes, per and cyc, in nymphal heads; in addition, per expression is also photoperiod‐dependent. The RNA interference‐mediated knockdown of the two circadian clock genes, Clk and cyc, during the nymphal stage results in reproductive arrest in adult females. Furthermore, Clk and cyc knockdown induces the expression of the storage protein hexamerin in the fat body, whereas the expression of vitellogenin diminishes. Taken together, these data support the involvement of circadian clock genes in photoperiodic timer and/or diapause induction.     

Wing morph-related physiological differences in adults of temperate population of Pyrrhocoris apterus (L.) (Heteroptera: Pyrrhocoridae)

The reproductive and diapausing adult females of brachypterous morph and macropterous females with reproductive arrest of non-diapause type, originating from the laboratory cultures of Pyrrhocoris apterus, were studied for their feeding and drinking behaviour, digestive enzyme activities, and carbohydrate and lipid contents. The highest feeding and drinking activities were observed in reproductive brachypters, the lowest in macropters. Macropters also differed from brachypters by lower activities of gut lipase, peptidase and protease, lower concentration of haemolymph sugars, and lower weight of fat body, which probably reflects their low feeding activity. The total content of fat body lipids was also lower in macropters (0.6 mg) than in reproductive and diapausing brachypters (4.6 and 7.5 mg, respectively) on day 14. A very high amount of glycogen was found in the fat body of diapausing brachypters, 363 micrograms on day 14, as opposed to 15 and 80 micrograms in macropterous and reproductive brachypterous females, respectively. The obtained data indicate that the most important difference between macropterous and brachypterous females with different types of reproductive arrest consists of an enhanced mobilization of lipids for dispersal in macropters and accumulation of energetic reserves for hibernation in brachypters.     

Wing Movement Behavior in Long- and Short-Winged Morphs of the Flightless Bug Pyrrhocoris apterus L. (Heteroptera: Pyrrhocoridae)

The aim of the present paper was to study, with the use of video-recordering and computerized methods, the flight and wing movement behavior in two wing morphs of the flightless bug Pyrrhocoris apterus during the first 14 days of their adult life. We demonstrated the capability of macropterous males and females of this heteropteran to flap their fore- and hindwings for the first time. Flight was not observed in any of the tested males and females. We revealed that wing flapping does not occur earlier than on the fourth and fifth day after adult emergence in males and females, respectively. Time-series analysis of the obtained data showed that the maximum activity of wing flaps is limited to the light phase of the 12-h light/12-h dark photocycle, with the peak occurring in the morning, i.e., about 3 h after turning the light on. No substantial differences in the peaks of wing flapping activities in males versus females were found. The present paper is the first experimental demonstration of wing flap activity in a representative species with a nonfunctional alary polymorphism.     

Regulation of body size in a heteropteran bug, Pyrrhocoris apterus

The determination of body size (length or weight) of teneral adults of Pyrrhocoris apterus L. was investigated. About 50% of size variation was determined already in the egg stage (average egg weight adult length correlations were 0.67 in males and 0.71 in females). During the larval development, starvation, low temperature, and pollution by excreta combined with crowding reduced adult size and usually increased the development length. The effects were directly proportionate to the length of the stress period, and increased with larval age. Photoperiod or intrinsic variation of development time did not affect adult size. The effects of stress were largely compensated when it was followed by a period of compensation growth at optimum conditions. A dramatic reduction of size appeared when last inster larvae starved and moulted to adults without access to food. Average adult weight was then reduced by up to 48%. Threshold weight for adult moulting was 18 mg, being attained after 1–2 day feeding of the last instar.

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Zusammenfassung Die Körpergrösse bei den Imagines eines Wildstammes der Feuerwanze Pyrrhocoris apterus wurde teilweise durch die Veränderlichkeit der Eigrösse determiniert, teilweise auch durch Hungern der Larven, niedrige Temperatur und metabolischen Produkten beeinflusst. Der Effekt letzterer Faktoren stieg mit deren Dauer und war je grösser desto näher sich die Wirkungsperiode zur Imaginalhäutung befand. Unterschiedliche Photoperioden wirkten auf die Körperlänge nicht. Wurden die Larven zwischen der ungünstigen Periode und der Imaginalhäutung noch den optimalen Bedingungen ausgesetzt, kompensierten sich die negativen Einflüsse weitgehend. Zu einer drastischen Verkleinerung der Imagines kam es nur, wenn das letzte Larven-stadium mit Hunger beeinflusst wurde und bis zur Imaginalhäutung keinen Zutritt zum Futter hatte. Minimales Gewicht, wo die Metamorphose der Larven noch möglich war, wurde zu 18 mg festgestellt.

     

Voltinism and seasonal changes in haemolymph protein pattern of Pyrrhocoris apterus (Heteroptera: Pyrrhocoridae) in relation to diapause

Abstract. The red firebug, Pyrrhocoris apterus (L.), was shown not to be strictly a monovoltine species in Czechoslovakia. A second generation arises from eggs laid by females emerging between June and the beginning of August. Oviposition takes place before diapause is induced in the females by shortening of photophase. Adults that moult later, enter diapause without having laid eggs.
Electrophoretic patterns of haemolymph proteins of adults collected in the field throughout a year showed a characteristic temporal pattern of changes correlated not only with the stages of ovarian development, but also with the progression of diapause and post-diapause quiescence. The changes mainly concerned vitellogenin and polypeptides of molecular weight 78 and 80 kDa.
Vitellogenin was present only in the haemolymph of reproductive females. The polypeptides of 78 and 80 kDa, belonging to the group of storage proteins, accumulated conspicuously in the haemolymph of adults undergoing diapause and post-diapause quiescence. However, the occurrence of these polypeptides seems not to be connected exclusively with cold, because their titres increased with the length of diapause development, even in diapausing adults reared in the laboratory at a constant 26.C     

Intermittent character of adult diapause in Aelia acuminata (Heteroptera)

The response to photoperiod of field-collected Aelia acuminata was studied by exposing insects to three photoperiodic regimes. After the completion of diapause the insects do not respond to photoperiod: in mid-February the length of the pre-oviposition period is the same under both long and short photoperiods. After the onset of oviposition, insects again become responsive to photoperiod regardless of the time of year: under long day, oviposition lasts until death, whereas under short day the females cease oviposition after about 3 weeks. After a further 2 or 3 months without ovipositing, these females which are still experiencing short-day conditions resume reproduction for a further 4–7 weeks and long-lived individuals have three oviposition periods. A regulatory mechanism is suggested for the alternation between oviposition and diapause.