The regulation of development during diapause in Ephestia elutella (Hübner) by temperature and photoperiod

Diapausing larvae of Ephestia elutella reared at 20°C in short photoperiods (LD 11:13), and then maintained 12 weeks or longer at 5–15°C before transfer to 20 or 25°C, pupated sooner than unchilled controls. At 25°C, all samples kept in long photoperiods (LD 15:9) survived better and pupated faster than similarly treated samples held in short photoperiods (LD 9:15). Samples kept at 20°C after chilling pupated much slower than those at 25°C, and, except after exposure at 5°C, pupated at similar rates at LD 11:13 or 15:9, although mortality was higher at the shorter photoperiod. After exposure at 5°C, larvae required increased day-length as well as increased temperature to hasten pupation whereas after exposure at 10°C most responded to increased temperature only.For samples maintained in slightly heated or unheated outbuildings, the summer emergence was poorly synchronized and males on average emerged ahead of females. Samples moved from the unheated outbuilding to 25°C and long days in the laboratory in early spring, however, pupated quickly and males and females emerged together. A late phase of diapause development thus exists requiring both high temperature and long photoperiods to ensure a prompt resumption of morphogenesis. Spring temperatures in the United Kingdom are seldom high enough to synchronize the completion of diapause.     

Effect of photoperiod and temperature on diapause in a Florida strain of the tropical warehouse moth Ephestia cautella

A photoperiodically-controlled diapause of the long-day, short-day type was identified in a brown-winged, yellow-eyed strain of Ephestia cautella (Walker). The proportion of larvae diapausing in very long photoperiods was less than in short photoperiods. The mean critical photoperiod, here defined as that photoperiod giving half the maximum percentage of insects that diapause in response to photoperiod at a given temperature, was between 12 and 13 hr for the long-day reaction at both 20 and 25°C. The principal sensitive phase occurred near the time of the last larval moult. The mean duration of diapause was 2–3 months at 20°C and slightly longer at 25°C. The optimum temperature for diapause development was near 15°C, all larvae pupating within 24 days after a 45-day exposure at this temperature. Diapause could be terminated whenever larvae diapausing at 20°C were exposed to as few as five long (15 hr) photoperiods at 25°C. Long photoperiods at 20°C, or short photoperiods (9 hr) at 25°C were less effective in terminating diapause.     

Effects of thermoperiod on photoperiodic determination of larval diapause in Ostrinia nubilalis

Some effects of thermoperiods on the photoperiodic determination of larval diapause in the European corn borer, Ostrinia nubilalis are reported. Thermoperiods and photoperiods were shown to interact to a highly significant degree in the induction of diapause. Diapause determination was found to be sensitive to the duration and temperature of the cryoscotophase of the thermophotoperiod; the thermophotophase characteristics exerted only minor influence. Cryoscotophase temperatures of 10°C or lower caused an unexpected increase in the apparent critical nightlength for diapause determination. A 0°C cryoscotophase effectively prevented the photoperiodic determination of diapause. Experimental investigation of the effects of 0°C cryoscotophases produced evidence that such low-temperature cryoscotophases may suppress the biological clock functions regulating the determination of diapause. Photoperiodic regimes in which 0°C pulses were inserted at successive points in 24-h cycle evoked responses suggesting that the early scotophase (saturation phase) was more strongly influenced by the low temperature pulse than was the later scotophase (scotonon); low temperature had little effect on the light-dependent processes (photonon) associated with the photophase.     

Geographic variation in critical photoperiod for diapause induction and its temperature dependence in Hyphantria cunea Drury (Lepidoptera: Arctiidae)

The fall webworm, Hyphantria cunea Drury (Lepidoptera: Arctiidae), was introduced from North America to Japan half a century ago. The critical photoperiod for diapause induction and its temperature dependence, as defined by the difference in the critical photoperiod between 20 and 25°C, were investigated in order to understand the mechanisms behind a shift from bi- to trivoltine life cycles. The critical photoperiod for diapause induction was shorter in the southern trivoltine populations than in the northern bivoltine populations, and this was more marked at 25°C than at 20°C. Although the critical photoperiod showed a positive correlation with the original latitude, the correlation was relatively low at both temperatures. Conversely, temperature dependence of the critical photoperiod for diapause induction correlated negatively with the original latitude. The trivoltine populations showed greater temperature sensitivity than the bivoltine populations. These results suggest that an increase in temperature sensitivity of the diapause response to photoperiods was involved in the shift to a trivoltine life cycle. The crossing experiments suggested that the photoperiodic control of diapause induction and its temperature dependence are under polygenic control without sex-linkage. Received: 29 October 1996 / Accepted: 26 February 1997     

Effects of photoperiod and temperature on diapause induction and termination in the swallowtail,Sericinus montelus

Abstract Sericinus montelus overwinters as diapausing pupae. In the present study, the effects of photoperiod and temperature on diapause induction and termination of diapause are investigated. The results obtained demonstrate that high temperature can reverse the effect of short day‐lengths on diapause induction. Under an LD 12 : 12 h photoperiod, all pupae enter diapause at 15, 20 and 25 °C, whereas all pupae develop without diapause at 35 °C. No pupae enter diapause under an LD 14 : 10 h photoperiod when the temperature is above 20 °C. Photoperiodic response curves obtained at 25 and 30 °C indicate that S. montelus is a long‐day species and the critical day‐length is approximately 13 h at 25 °C. At 25 °C, the duration of diapause is shortest when the diapausing pupae are maintained under an LD 16 : 8 h photoperiod and increases under LD 14 : 10 h and LD 12 : 12 h photoperiods. Under an LD 16 : 8 h photoperiod, the duration of diapause is shortest when the diapausing pupae are maintained at 25 °C, followed by 20 and 30 °C, and then at 15 °C. These results suggest that a moderate temperature favours diapause development under a diapause‐averting photoperiod in this species. The duration of diapause induced by an LD 12 : 12 h photoperiod is significantly longer at 25 °C than those at 15, 20 and 30 °C, and is shortest at 15 °C. At 25 °C, the duration of diapause induced by LD 6 : 18, LD 12 : 12 and LD 13 : 11 h photoperiods is similar and longer than 90 days. Thus, the diapause‐inducing conditions may affect diapause intensity and a photoperiod close to the critical day‐length has significant influence on diapause intensity in S. montelus.     

Diapause induction in the oblique-banded leafroller Choristoneura rosaceana (Lepidoptera: Tortricidae): Role of photoperiod and temperature

Induction of diapause in the larval stage of the oblique-banded leafroller, Choristoneura rosaceana (Harris), was found to be dependent on both photoperiod and temperature. At constant temperatures of 24, 20 and 16°C, short photoperiods induced diapause. The critical photoperiod was between 14–15 h of light per day at 20 and 16°C. At 14 h light: 10 h dark, all larvae expressed diapause. Temperature had a modifying effect, and slightly shifted the larval response to diapause-inducing photoperiods. High constant temperatures of 28°C and above induced diapause in some individuals (< 20%), while fluctuating temperatures of 32 and 16°C in a 12-h cycle resulted in 67% diapause induction, suggesting that diapause could also be induced by fluctuating temperatures, particularly if the higher temperature exceeds 25°C.The first- and the second-instar larvae were the only two stages sensitive to diapause induction. Exposure of adult, egg and third, fourth, and fifth-larval instars to diapause-inducing conditions did not produce diapause. Although diapause was induced in the first or the second instars, it was always expressed in the third or fourth instar.     

Timing of diapause induction outside the natural distribution range of a species: an outdoor experiment with the bean bug Riptortus clavatus

The phytophagous bug Riptortus clavatus (Thunberg) (Heteroptera: Alydidae) produces two or three generations per year in Central Japan and overwinters in the adult stage. In bugs from the Kyoto population (35°00 N, 135°45 E), we studied (1) the effects of day-length on the nymphal and preoviposition periods under constant photoperiod at 20.5 °C, and (2) photoperiodic induction of adult diapause at 20.5 °C and under a combination of constant photoperiod and natural daily rhythm of temperature in the forest-steppe zone of Russia (50°38 N, 35°58 E). Then, we examined (3) the timing of diapause induction under quasi-natural conditions in the same region, far outside the species' natural geographical range. At 20.5 °C, the nymphal period in both males and females was significantly longer under regimes with shorter photophases than under those with longer photophases. The preoviposition period in females was significantly longer under the near-critical long-day regime L14:D10 than under typical long-day regimes (L15:D9, L16:D8, and L17:D7). The critical day-length for diapause induction was shorter under conditions of natural daily rhythm of temperature than those reported at constant 20, 25, and 30 °C. Under quasi-natural conditions in the forest-steppe zone, R. clavatus entered diapause in September, much later than the local populations of true bugs studied to date. This experiment showed that R. clavatus was maladapted to new environmental conditions: diapause was induced too late with the result that all or most nymphs hatched in late August or early September will die.     

Structural proteins of Amsacta moorei,Euxoa auxiliaris, and Melanoplus sanguinipes entomopoxviruses

The structural proteins of Amsacta moorei, Euxoa auxiliaris, and Melanoplus sanguinipes entomopoxviruses (EPVs) were separated by electrophoresis on sodium dodecyl sulfate (SDS)-polyacrylamide gels. More than 35 structural proteins were detected in each virus. Based on the distribution and the variation in the molecular weights of the virus structural proteins little homology was detected between the EPVs and vaccinia virus. The molecular weight of Amsacta EPV occlusion body matrix protein (110,000) was determined by SDS-acrylamide gel electrophoresis. The occlusion body matrix protein of Amsacta EPV occluded virus isolated from infected E. acrea larvae was rapidly degraded at pH 10.6 to peptides of approximately 94,000 and 60,000 daltons. After 2 hr incubation at alkaline pH, Amsacta EPV occlusion body protein was degraded to approximately 56,000 daltons. Proteolysis of occlusion body protein was inhibited by SDS. No proteolytic degradation was detected in occlusion body matrix protein isolated from Amsacta EPV infected BTI-EAA cells. Amino acid analysis indicates that entomopoxvirus occlusion body matrix protein consists of approximately 20% acidic amino acids and 9% of the sulfur-containing amino acids cysteine and methionine.     

Thermoperiodic induction of larval diapause in the European corn borer, Ostrinia nubilalis

Thermoperiodic induction of larval diapause was shown to occur in the European corn borer, Ostrinia nubilalis (Hübner). Using continuous darkness and thermoperiods of the format XC:24-XT(15:30°C), incidence of diapause was shown to be dependent on phase durations, with a critical cryophase of about 9.5 hr. A thermoperiodic response threshold was demonstrated; it was found to be very close to 17.5°C. Thermoperiodic amplitude (temperature difference between cryophase and thermophase) was shown to have no influence on the induction of diapause, within a relatively broad range of physiological temperatures. Thermoperiodic induction of diapause was shown to be at least partially independent from the effects of temperature on larval developmental rates.     

Effect of photoperiod on the maternal induction of an egg diapause in the grasshopper Chorthippus bornhalmi

The effect of the photoperiod experienced maternally on the induction of an egg diapause was studied in the grashopper Chorthippus bornhalmi Harz (Acridoidea: Gomphocerinae), which lives on the southern Balkan peninsula. Photoperiods used ranged from L12:D12 to L20:D4 and were kept constant, or were increased or decreased after the imaginal ecdysis in other experiments. Under most conditions in the study, the females laid almost only diapause eggs (not less than 93%). Using constant day-lengths, only at L14:D10 the percentage of non-diapause eggs was somewhat higher (20%). An increase in day-length from moderately long (L14:D10) to long day (L16:D8) was necessary for non-diapause eggs to predominate. However, the percentage of non-diapause eggs decreased with the age of the females, except when the day-length was raised a second time. An increase from short day (L12:D12) to moderately long day (L14:D10) or from long day (L16:D8) to extremely long day (L18:D6) as well as a decrease of day-length, all resulted in the production of diapause eggs. The consequences for the life cycle are discussed.

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Zusammenfassung Der einfluß der Photoperiode auf die maternelle Induktion einer Eidiapause beim Grashüpfer Chorthippus bornhalmi wurde untersucht. Die Art lebt auf der südöstlichen Balkanhalbinsel. Die getesteten Photoperioden reichten von L12:D12 bis L20:D4. In einem Teil der Versuche wurde die Photoperiode während des gesamten Lebens der Grashüpfer konstant gehalten, in anderen nach der Imaginalhäutung erhöht oder erniedrigt. Unter den meisten Bedingungen legten die Grashüpfer-Weibchen fast ausschließlich Diapause-Eier (mehr als 93%). Bei konstanten Tageslängen stieg der Anteil der Subitaneier lediglich nach einer Eiablage bei L14:D10 auf 20%. Eine Erhöhung der Photophase von 14 auf 16 Stunden war notwendig, damit der Anteil an Subitaneier überwog. Aber auch unter diesen Bedingungen nahm ihr Anteil mit dem Alter der Weibchen ab, außer wenn die Photophase noch ein zweites Mal (auf 18 Stunden) verlängert worden war. Eine Zunahme der Tageslänge von Kurztag (L12:D12) auf mäßigen Langtag (L14:D10) oder auch von Langtag (L16:D8) auf extremen Langtag (L18:D6) sowie eine Abnahme der Tageslänge hatten zur Folge, daß die Weibchen fast ausschließlich Diapause-Eier legten. Auf grund der spezialisierten Form der Diapause-Induktion können die Grashüpfer im Freiland nur im Frühjahr Subitaneier legen.

     

Effect of photoperiod experienced by parents on diapause induction in Trichogramma cacoeciae

In the genus Trichogramma, the prepupal stage can survive the cold season in diapause. However, optimal conditions for the induction of this cessation of development during the process of mass production of the parasitoid in a biological control program depend on the species. In Trichogramma cacoeciae Marchal (Hymenoptera: Trichogrammatidae), diapause is induced more easily if the parents are reared under short‐day conditions (L10:D14), and if the temperature is 10 °C rather than 13 °C. However, the effect of parental photoperiod on diapause induction is weaker at lower temperatures (10 °C). Following diapause induction, individuals can be stored at 3 °C for several months, up to 1 year. Non‐optimal conditions led to the establishment of a quiescent state in some or all individuals. In such cases, it was necessary to reduce the storage period to 1 or 2 months only, to prevent high mortality rates and low fecundity.     

Rapid seasonal adaptation of an alien bruchid after introduction: geographic variation in life cycle synchronization and critical photoperiod for diapause induction

Whether alien insects that are introduced into temperate regions adapt to seasonally changing environmental conditions is an important question in evolutionary biology. If rapid evolution has occurred in a non‐native environment, a latitudinal cline in critical photoperiod for diapause induction (i.e., the photoperiod at which half of the individuals enter diapause) and in life cycle synchronization with host plant phenology should be evident among locations. The alien bruchid Acanthoscelides pallidipennis (Motschulsky) (Coleoptera: Bruchidae) is native to North America and introduced into Japan with the host plant Amorpha fruticosa L. (Fabaceae) in the late 1940s. To examine whether seasonal adaptation has occurred in A. pallidipennis, we conducted a laboratory experiment and phenological observations using three latitudinally different populations. We bred F1 eggs at 22 °C and five photoperiodic regimens – L:D = 10:14, 13:11, 14:10, 15:9, or 16:8 hours – and examined whether diapause was induced. The estimated critical photoperiod for diapause induction was longest in the most northern population and shortest in the most southern population. Life cycle was found to be synchronized with host phenology in each location. Also voltinism varied geographically, from univoltine in the northern population to bivoltine in the southern populations. These results showed that A. pallidipennis rapidly adapted to seasonal environmental conditions in Japan after its introduction.     

Contribution of male-produced proteins to vitellogenesis in Melanoplus sanguinipes

The transfer during copulation of radioactively labelled male accessory reproductive gland (ARG) protein and its accumulation by the ovary of Melanoplus sanguinipes have been studied. Most of the transferred material leaves the spermatheca within 24 hr and enters the haemolymph from which it can be accumulated by the ovary. Injection of labelled male ARG protein into vitellogenic females demonstrates that during the first 24 hr after injection, accumulation by the ovary is rapid. Immunoprecipitation and immunoelectrophoretic studies indicate that some of the ARG protein is accumulated unchanged. It is proposed that when the male transfers several spermatophores during copulation, he may make a significant contribution of protein to the developing oöcytes.     

The subgenus Persicargas (Ixodoidea: Argasidae: Argas), 26. Argas (P.) arboreus: Effect of photoperiod on diapause induction and termination

Prefeeding exposure of female Argas (Persicargas) arboreus to short photoperiod (SP) induces diapause, expressed as oviposition delay for more than 40 days postfeeding. Diapause incidence increases as prefeeding SP exposure periods increase. Postfeeding SP exposure is not essential to maintain diapause. Postfeeding exposure to SP without prefeeding SP exposure does not induce diapause. A 2 week prefeeding exposure to long photoperiod (LP) terminates diapause in two-thirds of the females exposed to LP after feeding. The remaining one-third, which may differ genetically, require a 6 week LP prefeeding exposure to terminate diapause. An 8 week prefeeding exposure to LP is required to terminate diapause in females exposed to SP after feeding.     

Seasonal occurrence and diapause induction of a predatory bug Andrallus spinidens (F.) (Heteroptera: Pentatomidae)

The bug Andrallus spinidens (F.) (Heteroptera: Pentatomidae) is a polyphagous predator of insect larvae, which is distributed in tropical and warm temperate zones worldwide. Seasonal occurrence and diapause induction of this bug were studied in a population in Miyazaki, southern Kyushu, Japan. The field research showed that A. spinidens produces three or four generations a year and is most abundant from mid‐summer to autumn. Reproductive activity of field‐collected adults decreased from late summer to autumn, indicating that this bug enters adult diapause in autumn. Only adults were found in early spring and these were reproductive. Laboratory experiments showed that, irrespective of photoperiod, adult diapause is induced at lower temperatures (≤22.5°C), whereas it is avoided at higher temperatures (≥25°C). Ambient temperature falls across the critical range from late summer to autumn. Thus, the bug clearly overwinters in adult diapause induced by low temperatures and this diapause is terminated during the course of winter.     

Photoperiodic and temperature control of diapause induction and colour change in the southern green stink bug Nezara viridula

Abstract. The effect of photoperiod and temperature on the duration of the nymphal period, diapause induction and colour change in adults of Nezara viridula (L.) (Heteroptera: Pentatomidae) from Japan was studied in the laboratory. At 20 °C, the developmental period for nymphs was significantly shorter under LD 10 : 14 h (short day) and LD 16 : 8 h (long day) than under intermediate photoperiods, whereas at 25 °C it was slightly shorter under intermediate than short- and long-day conditions. It is assumed that photoperiod-mediated acceleration of nymphal growth takes place in autumn when day-length is short and it is unlikely that nymphal development is affected by day-length under summer long-day and hot conditions. Nezara viridula has an adult diapause controlled by a long-day photoperiodic response. At 20 °C and 25 °C in both sexes, photoperiodic responses were similar and had thresholds close to 12.5 h, thus suggesting that the response is thermostable within this range of temperatures and day-length plays a leading role in diapause induction. Precopulation and preoviposition periods were significantly longer under near-critical regimes than under long-day ones. Short-day and near-critical photoperiods induced a gradual change of adult colour from green to brown/russet. The rate of colour change was significantly higher under LD 10 : 14 h than under LD 13 : 11 h, suggesting that the colour change is strongly associated with diapause induction. The incidences of diapause or dark colour did not vary among genetically determined colour morphs, indicating that these morphs have a similar tendency to enter diapause and change colour in response to short-day conditions.     

Diapause induction and coloration in the Senegalese grasshopper, Oedaleus senegalensis

Abstract. Embryonic diapause induction in the Senegalese grasshopper, Oedaleus senegalensis Krauss (Orthoptera: Acrididae), is influenced both by the photoperiod and the temperature experienced by females. High temperatures (40°C) and long photoperiods (LD 14:10h), which characterize the beginning of the rainy season in the Sahel, cause non-diapausing eggs to be laid. Lower temperatures (25°C) and shorter photoperiods (LD 12:12h), which occur at the end of the rains, result in the production of diapausing eggs. At 30°C and constant photoperiods, O. senegalensis exhibited a long-day-short-day response with critical photoperiods of c. 13 h and c. 20 h, only the former value being of ecological significance. The photoperiodically sensitive stages to diapause induction in females occurred from the fifth stadium onwards. Temperature also affected the coloration of both nymphs and adults. Dark-black and pale-white individuals were produced by low (25°C) and high (40°C) temperatures respectively, whereas an intermediate temperature (30°C) produced individuals which were greyish brown. These results are discussed in relation to the ecology of O. senegalensis.     

Infection of Locusta migratoria with entomopoxviruses from Arphia conspersa and Melanoplus sanguinipes grasshoppers

Entomopoxvirus (EPV) occlusion bodies isolated from Arphia conspersa and Melanoplus sanguinipes grasshoppers were fed to 3rd and 4th instar Locusta migratoria nymphs. Locus mortality induced by A. conspersa EPV was first detected 18 days after addition of virus to the diet, and reached a level of approximately 68% of the colony population by 60 days after virus inoculation. In a similar population of L. migratoria nymphs, mortality induced by M. sanguinipes virus reached 90% 60 days after virus inoculation. Entomopoxvirus was isolated from M. sanguinipes EPV infected locust nymphs and the viral DNA was cleaved with several restriction endonucleases. The DNA fragment patterns obtained after agarose gel electrophoresis were compared with the fragment patterns from the original sample of M. sanguinipes EPV DNA cleaved with the same restriction endonucleases. No differences in the cleavage patterns were detected between the two virus DNA samples. Virus structural proteins of M. sanguinipes EPV purified from infected locust nymphs were compared by polyacrylamide gel electrophoresis with virus proteins isolated from the original sample of M. sanguinipes EPV. A total of six different virus protein bands were detected between the two poxvirus preparations.     

光周期和温度对亚洲玉米螟滞育诱导的影响

亚洲玉米螟Ostrinia furnacalis (Guenée)属兼性滞育昆虫, 其滞育特性与种群利用适生境的最大化和季节性活动密切相关。解析季节性因素如光周期和温度对其滞育的诱导作用可为预测种群发生动态提供科学依据。本实验在HPS 500型和HPG-320H型自动控制环境气候箱中, 测定了RH 80%下温度20℃, 27℃和30℃与11个光周期组合对吉林农安（NA）、 河北衡水（HS）、 广东惠州（HZ）和海南海口（HK）等4个地理种群的滞育诱导作用。结果表明： 光周期、 温度及其交互作用均对亚洲玉米螟滞育诱导具有重要影响, 其中光周期在滞育诱导中起主导作用。在20℃条件下, 其光周期反应曲线属典型的长日照反应型, 临界光周期随种群分布的地理纬度降低而缩短, 4个地理种群NA, HS, HZ和HK的临界日昼长依次为14 h 3 min, 13 h 59 min, 13 h 32 min和13 h 7 min, 最有效的滞育诱导光周期是日昼长12 h。另一方面, 随种群分布纬度升高, 其对光周期诱导滞育的敏感性降低。温度为27℃时, 其光周期反应曲线为短日照 长日照反应型, 各种群有两个临界光周期, 临界日昼长依次为12 h 50 min和13 h 32 min, 11 h 35 min和13 h 8 min, 12 h 58 min和13 h 1 min, 以及11 h 50 min和12 h 26 min, 且最有效的滞育诱导光周期范围的滞育率明显低于20℃。一定温度范围内（20℃～27℃）, 随温度升高, 临界光周期缩短; 温度达到30℃时则滞育显著被抑制, 滞育率仅为4.3%或更低。这些结果说明温度对亚洲玉米螟滞育的光周期诱导具有很强的补偿作用。因此, 在自然界亚洲玉米螟的滞育属具有温度补偿作用的长日照反应型昆虫, 临界光周期随地理纬度北移而增长; 种群滞育的短日照 长日照反应型特性可能是产生局部世代种群的重要内因。     

Extraglandular synthesis of accessory reproductive gland components in male Melanoplus sanguinipes

The accessory reproductive glands (ARG) of the male migratory grasshopper, Melanoplus sanguinipes, are able to accumulate injected labelled ARG protein from the haemolymph. Accumulation is slight in the ARG of 2-day-old virgins, or 7-day-old allatectomized (CA^?) insects. The ARG of 7-day-old virgins, or 7-day-old CA^? insects treated with synthetic juvenile hormone, accumulate about 1.5 times more label than those of 2-day-old insects in a 24-hr period. The ARG of recently mated males accumulate almost four times more label than those of 2-day-old controls. Immunoprecipitation studies indicate that about one fifth of the labelled protein is accumulated unchanged.The fat body and haemolymph contain proteins which are precipitable by antiserum to whole ARG homogenate. The concentration of these proteins in the fat body increases after removal of the ARG, or after copulation. It is concluded that the fat body synthesizes certain proteins which are accumulated by the ARG. Both the synthesis and the accumulation of these proteins are regulated by the corpora allata.