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.     

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 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 μg on day 14, as opposed to 15 and 80 μg 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.     

Developmental and diel changes of adipokinetic hormone in CNS and haemolymph of the flightless wing-polymorphic bug,Pyrrhocoris apterus (L.)

We have used an enzyme-linked immunoassay (ELISA) for determination of the AKH content in CNS (brain + corpora cardiaca + corpora allata) and haemolymph of adult macropterous and brachypterous females of the bug Pyrrhocoris apterus. The tests revealed that the AKH content fluctuates between 1 and almost 4 pmol/CNS during the first 14 days of adult life and significantly increased in order diapausing brachypters相似文献   

Differences in haemolymph proteins in relation to diapause and wing dimorphism inPyrrhocoris apterus (L.) (Heteroptera: Pyrrhocoridae)

Three different cultures of dimorphic bug,Pyrrhocoris apterus, were analyzed concerning diapause and its relation to wing morph pattern. The proportion of macropterous bugs was considerably higher (36%) in the Mediterranean culture from Israel than that (1.3%) in the temperate culture from the Czech Republic. The macropterous morph- and brachypterous morphrelated types of reproduction arrest, differing by the length of pre-oviposition period, were distinguished in cultures analyzed. The reproduction arrest with an average pre-oviposition period of 38.2 days in Mediterranean macropters and 18.5 days in macropters from selected macropterous strain, was found to be typical for macropterous morph. Two different photoperiodic conditions induced macropterous morph-related reproduction arrest, the long-day (18 h light-6 h dark) photoperiod in macropterous strain macropters and the short-day (12 h light-12 h dark) photoperiod in Mediterranean macropters. The brachypterous morph-related reproduction arrest, characterized by pre-oviposition period longer than 90 days, occurred predominantly in diapausing brachypterous bugs. While the hibernal diapause of brachypterous bugs was characterized by a very high level of the 78- and 82-kDa proteins in haemolymph, their content in haemolymph of macropters during macropterous morph-related reproduction arrest was almost as low as in the reproductive adults. The variation of reproduction arrest in relation to wing dimorphism represents an important feature in the life strategy ofP. apterus.     

Differential allocation of protein resources to flight muscles and reproductive organs in the flightless wing-polymorphic bug, <Emphasis Type="Italic">Pyrrhocoris apterus</Emphasis> (L.) (Heteroptera)

Differences in the growth of dorsolongitudinal flight muscles and gonads in 1–28 days old long-winged (macropterous) and short-winged (brachypterous) adults of the firebug (Pyrrhocoris apterus L.) and the resource allocation to these organs were studied by means of total protein analysis. We found predominant allocation of food resources to flight muscles compared to reproductive organs in both macropterous males and females during the first 5 days of adult life. Subsequent histolysis of developed flight muscles coincided with increased total protein content in some reproductive organs. Initiation of intensive food intake after starvation or application of higher dose of methoprene on macropterous adults changed the resource allocation in favour of growth of reproductive organs and induced precocious histolysis of flight muscles. It indicates an involvement of juvenile hormone in wing morph-related differential allocation of resources in the bug. Increased total protein contents in the ovaries and accessory glands of starved macropterous females and males treated with methoprene, respectively, indicate that proteins derived from the methoprene-induced histolysis of the flight muscles are re-utilized for the growth of the reproductive organs. It is the first report of persistence of differential resource allocation to flight muscles and reproductive organs in the wing-polymorphic insects with non-functional macropterism.     

稻飞虱翅型纯系后代个体的翅型分化对光周期变化不敏感

【目的】为了明确光周期和遗传因子在稻飞虱翅型分化中的作用, 研究了3种稻飞虱（褐飞虱Nilaparvata lugens、白背飞虱Sogatella furcifera和灰飞虱Laodelphax striatellus）翅型纯系或近纯系在不同光照时数下的翅型分化比率。【方法】以经过5~45代连续翅型筛选后的褐飞虱、白背飞虱和灰飞虱的长翅型和短翅型纯系或近纯系为材料, 在室内分别测定了其在长光照（16和20 h）、短光照(4~12 h)和正常光照(14 h) 3类光周期条件下饲养后, 雌、雄成虫中长翅和短翅个体出现的比率及存活率。【结果】白背飞虱和灰飞虱的长翅型纯系M♂×M♀或短翅型纯系B♂×B♀在不同光周期下的翅型比率均无显著差异（P>0.05）。褐飞虱短翅型近纯系B♂×B♀的雌虫短翅率和成虫总短翅率在不同光周期下也无显著差异（P>0.05）, 但雄虫短翅率在正常光照14 h和短光照4 h下显著高于长光照20 h下的（P<0.05）。当褐飞虱短翅型达到纯系后, 其后代翅型在6~16 h光照条件下无显著差异。褐飞虱长翅型近纯系M♂×M♀的后代虽有短翅个体出现, 但是雌虫和雄虫的各自短翅率在不同光周期下无显著差异（P>0.05), 仅总体短翅率在12 h光照条件下的显著高于16 h下的（P<0.05）。褐飞虱长、短翅型杂交筛选品系M♂×B♀的雌虫短翅率随光照时数的延长而升高; 灰飞虱杂交筛选品系M♂×B♀的短翅雄虫随光照时数的缩短而增多（P<0.05）, 但当筛选代次达到45代时, 这种趋势不再显著。3种稻飞虱长翅型和短翅型纯系或近纯系若虫的存活率会稍低于长、短翅型杂交后代的存活率, 但长、短翅型品系的存活率在6~16 h光照条件下差异不显著(P>0.05)。【结论】稻飞虱翅型分化对光周期的反应受飞虱本身遗传背景的影响, 翅型纯系后代个体的翅型分化对光周期变化不敏感。     

A male fitness advantage to wing reduction in the oriental chinch bug,Cavelerius saccharivorus okajima (heteroptera: Lygaeidae)

A male fitness advantage to wing reduction was investigated for the oriental chinch bug, Cavelerius saccharivorus, which is wing dimorphic. Field surveys for the frequency of matings between wing morphs showed that the mating probability was much higher in brachypterous males than in macropterous males in the early breeding season. Brachypterous males copulated with not only brachypterous females but also macropterous females in this season. This led to a considerable rate (30%) of insemination of macropterous females just before emigration even in the early breeding season. A rearing experiment for the pre-reproductive period of females revealed that females mated with brachypterous males copulated and oviposited earlier than females mated with macropterous males. These results indicate that there is a fitness advantage to wing reduction in the males of C. saccharivorus in terms of the earlier sexual maturation.     

Wing morph-related differences in developmental pattern of accessory gland proteins in adult males of Pyrrhocoris apterus (L.) and their endocrine control

The study showed that the amounts of the total proteins and 53 kDa protein in male accessory glands (AGs) of the firebug Pyrrhocoris apterus (L.) increased with age of the adult life. The 53 kDa protein, the most abundant polypeptide detected in the secretion of the AGs, and some other smaller peptides were identified as glycoproteins. Changes in the amounts of the total proteins and 53 kDa protein in AGs were found to be wing morph-dependent and their levels were significantly higher in 1-10 days old brachypterous males than in macropterous males of the same age. Macropterous males were characterized by delayed growth of the AGs. Treatment of adult macropterous males with methoprene significantly increased the amounts of total proteins and 53 kDa protein in their AGs when compared to acetone-treated macropterous controls of the same age. Allatectomy of brachypterous males decreased the levels of the total proteins and 53 kDa protein in their AGs, while application of methoprene enhanced the quantity of allatectomy-suppressed proteins in these tissues. This is the first report of juvenile hormone-dependent wing morph-related differences in the synthesis of AG proteins and their endocrine control in wing-polymorphic insects.     

Topical application of Pya-AKH stimulates lipid mobilization and locomotion in the flightless bug, Pyrrhocoris apterus (L.) (Heteroptera)

Abstract Two different methods of applying Pya -AKH to long-winged (macropterous) females of the firebug, Pyrrhocoris apterus (Linnaeus) (Heteroptera) were compared: both injection and topical application increased the levels of lipids in the haemolymph and stimulated locomotor activity. Lipid mobilization was maximal when 10 pmol was applied by injection or 40–100 pmol by topical application, with the first significant responses occurring 1.5 h after injection and 2 h after topical application. The highest elevations of lipid concentration in the haemolymph were comparable between the treatments, i.e. 14.36 ± 3.59 mg/mL for injection and 14.43 ± 4.07 mg/mL for topical application. However, these maximal elevations were achieved at different times: 3 h after the injection and 7 h after the topical application.
Injection of 10 and 40 pmol of Pya -AKH stimulated locomotor activity with maximal activity 3 h later but, surprisingly, injection of 80 pmol showed no effect initially and than a slight inhibitory effect after 6–8 h.
Increased locomotor activity was found after topical application of Pya -AKH, but the response was lower than after injection and appeared later, 5–9 h after the hormone application. In addition, the greatest increase in walking activity required topical application of 300 pmol and was still less dramatic than the response to injection. The stimulatory effect of Pya -AKH on locomotion was positively correlated with its effect on lipid mobilization only for injection of the hormone. It is argued that a stress caused by injection could play a role in the appearance of the complex response to adipokinetic hormone.     

Quantitative studies on the release of locust adipokinetic hormone

Fractionation of methanolic extracts of haemolymph on Sephadex LH-20 made possible the measurement of the titre of adipokinetic hormone in the haemolymph of locusts. Experimentally produced high concentrations of haemolymph carbohydrate caused a delay in the mobilization of lipid during flight, and very low titres of the hormone were present in the haemolymph of locusts injected with trehalose immediately before a 25 min flight. In these locusts flight speed was higher than saline-injected controls. Although delayed lipid mobilization during flight was also seen in locusts injected with sucrose, sucrose is not utilized for flight metabolism and flight speed was not increased by the injection. Tentative estimates of the release rate (c. 1000pg/20min flight) and half life (c. 20 min) of adipokinetic hormone during flight are made. The results described suggest that during flight the rate at which trehalose disappears from the haemolymph does not play a major role in the initiation of the release of adipokinetic hormone.     

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.     

Temporal pattern of feeding activity in the firebug Pyrrhocoris apterus and its relation to sex, wing dimorphism and physiological state of adults

Abstract.  The present study tested whether the pattern of feeding activity in the firebug Pyrrhocoris apterus (L.) is sex- and wing morph-related, diurnal or nocturnal, as well as whether the feeding rhythm persists in constant darkness. Temporal patterns of feeding activity are analysed in macropterous and brachypterous adults reared under long-day (LD 18 : 6 h) and short-day (LD 12 : 12 h) photoperiods, and in adults transferred to constant darkness. In females, the total feeding activity is highest in long-day reproductively active brachypters, intermediate in short-day diapausing brachypters, and lowest in macropters; the differences among males are substantially smaller. Although the total feeding activity of macropterous males is higher than in macropterous females, no sex-related differences are found in feeding activity of diapausing and reproductively active brachypters. The frequency of feeding exhibits sex-related differences, with obviously higher values in males. Mean feeding periods of macropterous and reproductively active brachypterous males are shorter than in females of the same wing morph. Mean interfeeding periods are longest in macropters, intermediate in diapausing brachypters, and shortest in reproductively active brachypters, and always lower in males than in females. The study shows that the feeding activity of P. apterus adults is age-, sex- and wing morph-related, and exhibits a diurnal pattern, except in reproductively active brachypterous females. The latter do not express a clear diurnal rhythm of feeding, presumably because of interactions with cycles of egg development and oviposition. The persistence of diurnal rhythm of feeding activity in short-day brachypterous females transferred to constant darkness indicates an endogeneity of this rhythm in P. apterus .     

Flight fuel related differences between solitary and gregarious locusts (Locusta migratoria migratorioides)

Abstract. Flight fuel relations of crowded and isolated Locusta migratoria migratorioides were investigated in younger (12–16 days after fledging) and older (27–30 or 27–32 days after fledging) adult males.No phase polymorphism dependent differences were found in resting haemolymph carbohydrate levels of the younger locusts.In the older age group, resting haemolymph carbohydrate levels were slightly though significantly higher in the isolated than in the crowded locusts.Injection of various doses of synthetic adipokinetic hormones (AKHs) did not induce marked changes in haemolymph carbohydrate levels and no differences were found between crowded and isolated locusts.A 30 min flight led to the same decrease in haemolymph carbohydrate levels of isolated and crowded locusts, 43.3% and 44.6% of the resting levels, respectively.We concluded, therefore, that the results do not seem to indicate that isolated locusts rely more heavily on carbohydrates as flight fuel than crowded locusts.Hyperlipaemic responses to flight were less intense in isolated than in crowded locusts, but phase polymorphism dependent differences in flight-induced increase of haemolymph lipid levels were not parallel in 12–16-day-old and 27–32-day-old males.In the younger age group the difference was mainly in the duration of flight needed to induce full response which appeared already after 20 min of flight in the crowded locusts, but only after 45 or 60 min of flight in the isolated ones.In contrast, the older isolated locusts showed markedly lower haemolymph lipid elevations than the crowded locusts even after 30, 45 or 60 min of flight.The hypothesis is forwarded that isolated locusts have a rather coarse adipokinetic strategy focused on a single long-distance migratory flight, whereas gregarious locusts possess a fine adipokinetic balance for reiterative migratory flights and saving fuel reserves for unpredictable long-distance migrations.     

The effect of constant darkness on the content of adipokinetic hormone, adipokinetic response and walking activity in macropterous females of Pyrrhocoris apterus (L.)

Abstract.  Changes in the content of adipokinetic hormone (AKH), the adipokinetic response and the walking activity of 10-day-old adult macropterous females of the firebug, Pyrrhocoris apterus (L.), reared under long-day (LD) photoperiod (LD 18 : 6 h) are compared with those exposed for 3 days to constant darkness (DD). Diel changes of all the parameters studied in LD females persist in females kept in constant dark. A positive correlation exists between diel changes of AKH content in the central nervous system (CNS) in the LD and DD females, and a negative correlation in the AKH level in haemolymph and walking activity. In addition, there is a positive correlation between diel changes of AKH level in haemolymph and walking activity in macropterous females reared under LD conditions, as well as in those transferred to constant darkness. The data suggest that there is some feedback between the release of AKH from CNS into the haemolymph and walking activity in macropterous females. Preliminary studies on the simultaneous expression of mRNA for the period gene and a positive reaction to an antibody against AKH in the same corpus cardiacum cells suggest that the period gene may be involved in regulating the AKH content in this gland.     

褐飞虱长翅型和短翅型雌雄成虫体内磁性物质定位和定量检测

【目的】地磁定向是昆虫远距离迁飞定向的重要机制之一。本研究以迁飞性害虫褐飞虱Nilaparvata lugens长翅型和短翅型雌雄成虫为研究对象,系统开展虫体内磁性物质定量研究。【方法】本研究利用MPMS-7型超导量子磁强计检测褐飞虱长翅型和短翅型雌雄成虫体内的磁性物质,并将经普鲁士蓝染色后的虫体超薄石蜡切片于JEM-2100型透射电子显微镜下观察磁性物质的分布状况,最后利用原子发射光谱法对虫体内的磁性物质进行定量分析与比较。【结果】超导量子磁强计检测发现,仅在褐飞虱长翅型和短翅型雌雄成虫腹部的磁滞曲线有明显闭合现象,证明该部位存在磁性物质;经透射电镜观察发现,普鲁士蓝沉淀主要呈点簇状存在于虫体腹部,证明该部位存在铁磁性物质;通过等离子发射光谱检测发现,同一性别长翅型成虫体内铁离子含量显著高于短翅型成虫,且相同翅型雄成虫体内铁离子含量显著高于雌成虫。【结论】褐飞虱长翅型和短翅型雌雄成虫腹部普遍存在铁磁性物质,且不同翅型和性别间磁性物质含量差异显著。     

Flight muscles polymorphism in a flightless bug, Pyrrhocoris apterus (L.): developmental pattern, biochemical profile and endocrine control

The flightless bug Pyrrhocoris apterus (L.) is polymorphic for both wing length and flight muscle development. The developed flight muscles of macropterous adults of both sexes first enlarge their volume during the first 5 days after adult emergence, but are then histolyzed in all males and females older than 10 and 14 days, respectively. The flight muscles of brachypterous adult males and females are underdeveloped due to their arrested growth. The total protein content of histolyzed dorsolongitudinal flight muscles from 21-day-old macropterous adults of both sexes is lower than that of developed dorsolongitudinal flight muscles in 5-10-days-old macropterous bugs, but substantially higher than the protein content of underdeveloped dorsolongitudinal flight muscles from adult brachypters. Histolyzed dorsolongitudinal flight muscles differ from the developed ones by decreased quantities of 18 electrophoretically separated proteins. Histolysis of developed dorsolongitudinal flight muscles is accompanied by significant decreases in citrate synthase, glyceraldehyde-3-phosphate dehydrogenase and β-hydroxyacyl-CoA dehydrogenase enzyme activities and an increase in alanine aminotransferase activity, and can be precociously induced by application of a juvenile hormone analogue. This is the first report of flight muscle polymorphism, histolysis of developed flight muscles and its endocrine control in insects displaying non-functional wing polymorphism.     

褐飞虱的交配行为及与保幼激素滴度的关系

受5龄末期和羽化初期较高保幼激素滴度的影响,褐飞虱Nilaparvata lugens(Stal)短翅型成虫性成熟比长翅型成虫早40～50h。5日龄前短翅型雌成虫的交配成功率较高,雄成虫的交配竞争能力较强;1、2日龄的长翅型雌成虫不交配,5日龄后长翅型成虫的交配能力渐渐超过短翅型成虫。实验种群的交配持续时间为(70.38±41.77)s;交配成功率为51.63%;雌、雄虫均可重复交配。一日中0～4h和10～14h交配成功率较高。     

褐飞虱的迁人和翅型分化规律的研究

褐飞虱Nilaparvata lugens(Stal)属迁飞性水稻害虫,其成虫有短翅和长翅两种翅型。 褐飞虱迁入雌虫均未经过交配,其短翅型由一对显性等位基因控制。翅型分化同时受到遗传和内分泌系统的协调控制,外界条件如密度和寄主等因子通过内分泌系统来影响翅型的分化。褐飞虱翅型分化的敏感龄期雌虫为1～3龄,而雄虫为1-5龄;不同若虫蜜度处理对褐飞虱成虫的前翅形成有一定的影响。分蘖期水稻饲养的褐飞虱短翅型比例明显高于孕穗期水稻饲养的褐飞虱。不同密度下各生物型间的翅型分化差异不显著。迁入地的浙江各种群属温带型,其雌虫短翅率低且与密度呈显著负相关;菲律宾热带种群雌虫在不同密度下均为短翅型,而雄虫的短翅率随密度增加而上升。广西种群接近热带型,其雌虫短翅率高但不随密度而变化。试验各种群的雄虫在中等密度甚至高密度时其短翅率出现最高。     

Adaptive significance of wing dimorphism in males ofNilaparvata lugens

The trade-offs associated with macroptery in males were investigated in a wing-dimorphic planthopper, Nilaparvata lugens (Stål) (Delphacidae, Hemiptera). Developmental time of brachypters was significantly shorter than that of macropters across a wide range of population densities, while no within-population differences between wing morphs in body weight were found. In mating experiments, brachypterous males 12 to 24 h old were almost three times more successful in competition for females than macropterous males of the same age, while in five days old males, macropters were twice as successful as brachypters. In competition for females involving direct body contact between males, brachypterous and macropterous males were equally successful. It is suggested that macroptery and delayed peak in sexual activity in males may both be influenced by the titre of juvenile hormone, as is known from females. The combination of prolonged development and delayed maximum competitive ability for mates, associated with macroptery, is likely to be disad vantageous, representing probably a penalty associated with macroptery in males in undisturbed habitats.