Lipid, carbohydrate and nitrogen content of long- and short-winged Gryllus firmus: Implications for the physiological cost of flight capability

Concentrations of total lipid, triglyceride, soluble carbohydrate, total nitrogen and water were measured in the long-winged (LW) and short-winged (SW) morphs of the cricket, Gryllus firmus. In addition, the weights and composition of wings and oviposited eggs were compared between morphs. This was done to obtain information on the energetic cost of flight capability in the LW morph. Whole-cricket content (% dry mass) of triglyceride was significantly higher in LW vs SW individuals of both sexes. Since triglyceride is a likely flight fuel in G. firmus, the biosynthesis of elevated levels of this high energy substance in the LW morph may represent an important energetic cost of flight capability. The existence of such a cost is consistent with the elevated respiratory metabolism previously observed in LW vs SW G. firmus. A highly significant negative correlation was observed between triglycerides and non-triglycerides in LW but not SW crickets. This suggests that lipid biosynthesis may be operating under some constraint in the LW morph. Increased triglyceride biosynthesis may require a concomitant decreased biosynthesis of non-triglycerides. In contrast to the elevated triglyceride level in the LW morph, carbohydrate concentration was higher in the SW morph during early adulthood. Carbohydrate content also decreased with age in the SW but not in the LW adults. No differences were observed between morphs in (1) the total nitrogen or water contents of whole crickets, (2) the nitrogen content of wings or (3) the wet weight, dry weight, lipid content, or total nitrogen content of oviposited eggs.     

The physiological costs of flight capability in wing-dimorphic crickets

Nutritional indices, triglyceride levels and flight muscle developmental profiles were compared between long-winged (LW) and short-winged (SW; flightless) morphs of the cricketsGryllus rubens Scudder andG. firmus Scudder. This was done to identify potential physiological costs of flight capability in adults. The LW morph of each species converted a lower proportion of assimilated nutrients into biomass (reduced ECD) than did the SW morph. This documents increased respiratory metabolism in the LW morph. Triglyceride concentration was higher in LW vs. SW adults. This suggests that the elevated respiration in the LW morph may be at least partially due to the increased biosynthesis of this high energy substance. Preliminary data indicate higher respiration rates of LW functional vs. SW vestigial flight muscles. Collectively, these data suggest that the energetic cost of flight capability in adults results from biosynthesis of triglyceride flight fuel and flight muscle maintenance but not flight muscle growth. No flight muscle growth was observed in adults.     

Effects of suppressed oviposition activity and flight muscle histolysis on food consumption and ovarian development in a wing-dimorphic cricket: an explanation for sporadic conclusions related to physiological trade-offs

The effects of deprivation of oviposition substrate on food consumption and egg production were compared between the long-winged (LW) and the short-winged (SW) morph of a cricket, Modicogryllus confirmatus, to determine how suppressed oviposition activity would influence these traits in each wing morph. Food consumption was greatly suppressed in females deprived of oviposition substrate (-OS) compared to those given access to it (+OS) during the 2-week feeding trial in the SW morph but not in the LW morph. Some LW females shed their hindwings and histolyzed the flight muscles. Such de-alated LW (DLW) morphs tended to consume more food than intact LW (ILW) morphs. In all morphs, ovarian weight was heavier under -OS conditions than under +OS conditions during the second week of adulthood, although the differences were greater in SW morphs than in ILW morphs. In DLW morphs in which flight muscle histolysis was induced by artificial de-alation at adult emergence, the temporal changes in ovarian weight were similar to those of SW morphs.In SW morphs, food consumption was also significantly reduced when ovipositing females were deprived of oviposition substrate for 2 days compared to those allowed to oviposit continuously, but food consumption was not reduced in ILW or DLW morphs. SW females from which one ovary was extirpated at adult emergence, SW (-o), also showed a significant difference in food consumption when treated as above, indicating that food consumption was not determined simply by the number of ovarian eggs. The crop content was positively correlated to food consumption and smaller under -OS conditions than under +OS conditions. The 2-day deprivation of oviposition substrate caused no significant difference in the total number of deposited and ovarian eggs in any group, but the ovarian mass of developing oocytes tended to be smaller under -OS than under +OS conditions, particularly in SW morphs.These results indicate the possibility that some inconsistent results and conclusions discussed in recent studies, concerning the physiological trade-offs between flight capability and reproduction, were caused by the suppressed oviposition activity and failure to recognize the occurrence of flight muscle growth and histolysis in the test crickets.     

长颚斗蟋长翅和短翅型雄成虫营养物质的积累与分配差异

[目的]翅多型雄虫在繁殖方面的能量投入与雌虫相异,这种差异可能会导致雄虫飞行与繁殖权衡的生理机制发生改变.因此,本研究旨在探究翅二型长颚斗蟋Velarifictorus aspersus雄成虫在营养物质积累与分配方面是否存在飞行与繁殖的权衡关系.[方法]选取长颚斗蟋V.aspersus头幅相近的长翅和短翅型雄成虫,对羽...     

沙蟋翅多型性的调控机理

沙蟋Gryllus firmus Zera&Denno成虫的后翅有长翅和短翅2种类型,是翅多型性机理研究的极佳模式昆虫。长翅成虫从第5日龄开始迁飞,而短翅成虫的主要特点是繁殖。除了翅的表型差异外,长翅成虫的飞行肌发达,呈褐色;卵巢幼小,直到飞行停止后(大约在10d以后)才开始发育。而短翅成虫的飞行肌退化并呈乳白色;卵巢在第4日龄就发育成熟,表现为卵巢硕大。对翅多型性机理的深入研究,将有利于了解沙蟋迁飞和扩散的内在机理,为准确地预测预报该虫的发生提供重要的理论和实际依据。文章概述沙蟋翅多型性与外界环境的相互关系,以及体内生化代谢和内分泌激素等的变化对该虫迁飞和生殖的影响和作用,进而探讨翅多型的遗传机制和进化意义等问题。     

Physiological trade-offs between reproduction,flight capability and longevity in a wing-dimorphic cricket,Modicogryllus confirmatus

Physiological and morphological comparisons were made between the short-winged (SW) and long-winged (LW) morphs of a cricket, Modicogryllus confirmatus, to determine the cost of the flight capacity and the physiological mechanisms underlying trade-offs between different life history traits related to migration and reproduction. Both wingmorphs grew at a similar rate and no consistent correlation was found between nymphal development and adult body size. The metathoracic muscles at adult emergence represented 4.2% of the wet body weight in the SW morph and 10.5% in the LW morph. Fat content of the body at adult emergence was positively correlated to dry body weight, but no significant difference was found in mean fat content between the two wingmorphs or between sexes after body weight was adjusted. SW females fed ad libitum produced significantly more eggs than LW females during the first 20days of adult life. Egg production was not correlated to either body size or the duration of nymphal development. LW adults lived longer than SW ones when kept with water alone or when given various amounts of food on day 1 and otherwise kept with water alone. In females, a highly significant correlation was found between longevity and egg production, indicating the presence of a trade-off. LW females mainly allocated the energy from food to flight muscle development and general maintenance of the body rather than to egg production, whereas SW females used it for egg production and longevity. LW females that had been de-alated at adult emergence histolyzed the flight muscle and used the energy from food for egg production almost exclusively. These results suggest that energy allocation and trade-offs after adult emergence may play crucial roles in the functional differentiation of the two wingmorphs.     

The role of juvenile hormone and juvenile hormone esterase in wing morph determination in Modicogryllus confirmatus

The role of juvenile hormone (JH) and juvenile hormone esterase (JHE) in regulating wing morph determination was studied in the cricket Modicogryllus confirmatus. JHE activities were significantly higher in nascent long-winged (LW) vs short-winged (SW) crickets during the latter half but not during the first half of the last stadium. The magnitude and direction of the activity differences were similar to those previously documented between wing morphs of the cricket, Gryllus rubens. In contrast, activities of general esterase, an enzyme or group of enzymes with no demonstrated role in regulating the JH titer in insects, showed no or only minor differences between morphs. The magnitude and direction of the JHE activity variation is consistent with a regulatory role for this enzyme in some aspect of wing dimorphism. However, the timing of the differences (exclusively during the last half of the last stadium) argue against a role in regulating wing length development per se. Single or multiple applications of juvenile hormone-III to nascent LW individuals during the first few days of the last stadium significantly redirected development from long to short wings. Multiple applications of acetone, by itself, also increased the production of short-winged adults. For most treatments, all individuals with shortened wings also had undeveloped flight muscles. These data suggest that JH may play a role in wing morph determination in M. confirmatus but that it affects a different aspect of the polymorphism from JHE.     

Differential allocation of resources underlies the dispersal-reproduction trade-off in the wing-dimorphic cricket,Gryllus rubens

The cricket, Gryllus rubens (Orthoptera, Gryllidae), exists in natural populations as either a fully-winged (LW), flight-capable morph or as a short-winged (SW) morph that cannot fly. The SW morph is substantially more fecund than the LW morph. In this study we report on the physiological basis of this trade-off between flight capability and fecundity. Results from gravimetric feeding trials indicate that LW and SW morphs are equivalent in their consumption and digestion of food. However, during the adult stage, the LW morph is less efficient in converting assimilated nutrients into biomass. This may be a consequence of the respired loss of assimilated nutrients due to the maintenance of functional flight muscles in the LW morph. In both morphs the gross biomass devoted to flight muscles does not change significantly during the first 14 days of adult growth while there is a significant biomass gain in ovarian tissue mass during the same period. SW morphs have vestigial flight muscles and gain substantially more ovarian mass relative to the LW morphs. These data are consistent with a trade-off between flight muscle maintenance in the LW morph and ovarian growth in the SW form. This is the first evidence for a life-history trade-off that has a physiological basis which is limited to the allocation of acquired and assimilated nutrients within the organism.     

Juvenile hormone titer and morph-specific reproduction in the wing-polymorphic cricket,Gryllus firmus

Juvenile hormone titers and reproductive characteristics were measured in adult wing and flight-muscle morphs of the wing-polymorphic cricket, Gryllus firmus, during the first week of adulthood. This species has three morphs: one flight capable morph with fully-developed wings and fully-developed flight muscles [LW(F)], one flightless morph with fully-developed wings and histolyzed (non-functional) flight muscles [LW(H)], and another flightless morph with underdeveloped (short) wings and underdeveloped flight muscles (SW). Both flightless morphs [LW(H) and SW] had larger ovaries which contained a greater number of postvitellogenic eggs compared with the flight capable [LW(F)] morph. The juvenile hormone titer was significantly higher in SW compared with LW(F) females on days 3-7 of adulthood. On these days, the JH titer also was significantly higher in the other flightless morph, LW(H), compared with flight-capable [LW(F)] females as determined by one statistical test, but did not differ significantly by another test. The JH titer was positively correlated with ovarian mass or terminal oocyte length, but not with the number of post-vitellogenic eggs. This study is the first direct comparison of juvenile hormone titers in adult wing morphs of a wing-polymorphic insect. Results indicate that an elevated juvenile hormone titer may be at least partly responsible for one of the most distinctive features of wing-polymorphic species, the increased early fecundity of flightless females.     

Critical Flight Time for Switch from Flight to Reproduction in the Wing Dimorphic Cricket Velarifictorus aspersus

Flight dimorphism has been considered to result from a balance of costs and benefits between flight capability and reproduction. The validity of the hypothesis, however, has been challenged recently. In this study, we examined the effect of flight time on trade-off between flight capability and reproductive development in Velarifictorus aspersus and we found that flight of 5 min did not promote reproductive development of long-winged (LW) adults, but flight of 30, 60, or 120 min could promote reproductive development both in female and male crickets. The results indicate that flight time may serve as a signal for LW V. aspersus to switch from migration to reproduction, and trade-off between flight ability and reproduction may be attenuated when flight time reaches a critical threshold. In addition, rapid reproductive development occurred before dealation when LW insects were allowed to fly for 30 min, which indicates that flight may influence reproductive development directly. Food consumption of short-winged adults was significantly higher than that of unflown LW adults or LW adults with 5 min flight, but similar to that of LW adults with 30, 60, or 120 min flight, suggesting that difference of reproductive development may be positively correlated with their food consumption.     

Juvenile Hormone and the endocrine regulation of wing polymorphism in insects: new insights from circadian and functional‐genomic studies in Gryllus crickets

For decades, Juvenile Hormone (JH) has been a major focus of studies investigating the endocrine regulation of wing‐polymorphism. The most general model postulates a single threshold, above which JH causes the expression of traits that define the short‐winged morph (SW), and below which JH causes the expression of traits that define the long‐winged morph (LW). Early studies in aphids and crickets reported ambiguous results as a result of the small size of aphids or the very low JH titre in nymphal crickets. Detailed studies in wing morphs of adult Gryllus firmus Scudder uncovered an unexpected and novel morph‐specific JH titre circadian cycle (cycling in LW but not in SW) in both the laboratory and field. This finding clearly contradicts the classic model. Morph‐specific daily rhythms in global gene expression are strongly associated with (and are possibly caused by) the morph‐specific JH titre rhythm. Daily rhythms for hormonal traits and gene expression, which are largely ignored in studies of life‐history evolution, may be common and play an important role in adaptation. Juvenile Hormone has likely evolved a specialized within‐morph function in G. firmus, regulating aspects of daily flight in the LW morph, which exhibits circadian flight. Other hormones, such as insulin‐like peptides and ecdysteroids, possibly regulate the expression of chronic (long‐term, noncircadian) differences between LW and SW morphs. Future studies should aim to investigate JH titres in more detail, as well as other hormones, most notably peptides and biogenic amines, which are largely ignored in endocrine studies of wing polymorphism.     

Effect of a juvenile hormone analogue on lipid metabolism in a wing-polymorphic cricket: implications for the endocrine-biochemical bases of life-history trade-offs

The wing-polymorphic cricket, Gryllus firmus, has a flight-capable morph (LW[f]: long winged with functional flight muscles) and a flightless morph (SW: short winged with reduced nonfunctional flight muscles) that differ genetically in many aspects of lipid metabolism. To determine whether these differences result from genetically based alterations in endocrine regulation, the juvenile hormone mimic, methoprene, was applied to the LW(f) morph. This hormone manipulation converted the LW(f) morph into a SW phenocopy with respect to all aspects of lipid metabolism studied; that is, methoprene application decreased in vivo biosynthesis of total lipid and triglyceride, increased absolute and relative biosynthesis of phospholipid, increased oxidation of fatty acids, and decreased in vitro specific activities of each of six lipogenic enzymes and a transaminase. Furthermore, methoprene increased ovarian growth and decreased fat body mass and flight muscle mass in the LW(f) morph. Differences in each of these biochemical, morphological, or reproductive traits between hormone-treated and control LW(f) females were similar in magnitude to differences between unmanipulated LW(f) and SW females. Variation in endocrine regulation contributes significantly to genetically based differences in lipid metabolism between LW(f) and SW females. This is the first evidence for endocrine regulation of a genetically based life-history trade-off operating via hormonal effects on specific metabolic pathways and enzymes of intermediary metabolism.     

丽斗蟋翅二型雄虫食物消化能力及消化酶活性比较

【目的】丽斗蟋Velarifictorus ornatus具明显的翅二型现象,为探讨翅型分化对丽斗蟋翅二型雄虫消化能力及中肠内消化酶活性产生的影响,对长翅型与短翅型雄虫食物消化能力及中肠内消化酶活性进行了检测比较。【方法】我们采取重量营养指数测定了羽化后12 d内丽斗蟋两型雄成虫增长量、相对增长率、取食量、食物利用率、近似消化率和食物转化率。为进一步明确丽斗蟋翅二型成虫食物消化能力与中肠内消化酶活性的关系,我们采用4种专用底物测定了中肠内用于分解蛋白质、脂肪和碳水化合物的总蛋白酶、胰蛋白酶、脂肪酶和淀粉酶的活性。【结果】结果表明,丽斗蟋两型雄虫取食量、食物转化率、食物利用率与增长量均无统计差异,但中肠内消化酶活性变化规律不同。成虫羽化后4 d时,长翅型雄虫中肠内总蛋白酶与胰蛋白酶活性显著高于短翅型雄虫,相反,羽化后0 d时,短翅型雄虫中肠内总蛋白酶与胰蛋白酶活性则显著高于长翅型雄虫,而羽化后12 d时,虽然短翅型雄虫总蛋白酶活性高于长翅型雄虫,但胰蛋白酶活性在两型雄虫间并无差异。成虫羽化后0 d时,两型雄虫脂肪酶活性无差异,但无论是羽化后4 或 12 d,长翅型雄虫中肠内脂肪酶活性皆显著大于短翅型雄虫。成虫羽化后4 d时,短翅型雄虫中肠内淀粉酶活性显著高于长翅型雄虫,而羽化后0与12 d时,两型雄虫间无显著差异。【结论】丽斗蟋翅二型雄虫食物消化能力无显著差异,但羽化后不同时间,中肠内消化酶活性存在差异,该差异可能与成虫羽化后不同时期,翅二型雄虫在飞行与繁殖投资中对不同能源物质的需求有关。     

长颚斗蟋长翅和短翅型雌成虫飞行肌发育、生殖力及寿命的比较

为探讨长颚斗蟋Velarifictorus asperses (Walker)翅型分化的生态学意义, 对室内饲养获得的长翅和短翅型雌成虫飞行肌和卵巢的发育, 以及长、 短翅型雌成虫的生殖力和寿命进行了比较研究。结果表明： 羽化当日, 长翅型雌成虫飞行肌重38.68±9.15 mg, 显著高于短翅型的17.53±4.44 mg （P<0.05）; 而二者卵巢重量无显著差异（P>0.05）, 分别为4.69±1.04 mg和4.88±0.97 mg。羽化后8 d内, 长翅型雌成虫飞行肌重量增加了48.9%, 短翅型雌成虫飞行肌重量无明显增加; 而短翅型雌成虫卵巢的重量增加至93.5±11.7 mg, 约为长翅型雌成虫的4.5倍。短翅型雌成虫的产卵前期显著短于长翅型, 其早期产卵量及总产卵量亦显著高于长翅型; 而两翅型雌成虫中后期产卵量及寿命无显著差异（P>0.05）。此外, 长翅型雌成虫在羽化后12 d开始发生飞行肌的降解, 飞行肌降解个体的卵巢重量显著高于未降解个体, 与短翅型相似。结果提示, 飞行肌与生殖系统的发育之间存在资源分配的权衡关系（trade-off), 且这种资源分配的差异可能会导致长翅型与短翅型个体在生活史策略上出现分化, 即长翅型个体具有飞行能力, 而短翅型个体则在生殖方面获得更高的收益, 且飞行肌的降解可能是长翅型个体由飞行转向生殖发育的生理信号。     

Morph-specific differences in metabolism related to flight in the wing-dimorphic Aphis gossypii

Abstract The cotton aphid, Aphis gossypii Glover, is a wing-dimorphic species, which causes globally important agricultural losses. In this present study, we compared the biochemical basis of wing polymorphism in A. gossypii with respect to trade-off of energy resources, including glycogen, trehalose, lipids (total lipid, triglyceride and phospholipid), free fatty acids, and soluble protein between dispersal and reproduction morphs during the wing-bud nymph and adulthood. Total lipid, triglyceride and free fatty acids were significantly higher in winged versus wingless morphs at 12 h of adulthood, the period during which alates are able to fly. By contrast, the wingless morph contained more glycogen than the winged morph from the 4th nymphal stage to adulthood. Trehalose content in the wingless morph was also higher than that in the winged morph during the 3rd and 4th nymphal stages, but vice versa at 12 h of adulthood. Finally, soluble protein content increased from nymphs to adults and was higher during adulthood in aptera versus alate. Whole-body water content in 12-h adults was significantly higher in apterae than that in alatae. These results indicate significant physiological differences between morphs related to specialization for flight.     

A morph-specific daily cycle in the rate of JH biosynthesis underlies a morph-specific daily cycle in the hemolymph JH titer in a wing-polymorphic cricket

A previous study documented a high amplitude, morph-specific daily cycle in the hemolymph JH titer in the wing-polymorphic cricket, Gryllus firmus. The JH titer rose and fell 10-20 fold in the flight-capable [LW(f), long-winged] morph during the late-photophase-early scotophase, while it was relatively constant during that time in the flightless (SW, short-winged) morph. In the present study we documented a dramatic morph-specific daily cycle in the in vitro rate of juvenile hormone (JH) biosynthesis that was tightly correlated with the hemolymph JH titer on days 5-7 of adulthood. Biosynthetic rates rose and fell 1-2 fold between the late photophase-early scotophase on each of days 5-6 and 6-7 of adulthood in the LW(f) morph, while biosynthetic rates were relatively constant during this period in the flightless, short-winged morph (SW), except for a slight dip in the rate of biosynthesis late in the photophase on these days. Similar morph-specific patterns of JH biosynthesis were observed whether rates were measured on corpora allata attached to corpora cardiaca in males or females, or on corpora allata alone. Hemolymph juvenile hormone esterase activity was significantly higher in the LW(f) vs. the SW morph during the beginning of scotophase, when the JH titer is decreasing rapidly in the LW(f) morph. Results indicate that the morph-specific daily cycle in the JH titer in G. firmus is primarily regulated by a morph-specific daily cycle in the rate of JH biosynthesis and to a lesser degree by hemolymph JH esterase activity. This is the first documentation of a diurnal cycle in the rate of JH biosynthesis in any insect, or a daily cycle in the rate of JH biosynthesis that is correlated with a specific morph in a polymorphic species. Results have important implications for the endocrine regulation of dispersal polymorphism, circadian rhythms of insect hormone titers and their regulators, and general studies of the JH titer and its regulation in insects.     

Potential Reproductive Advantage of Short-over Long-Winged Adult Males of the Cricket <Emphasis Type="Italic">Velarifictorus ornatus</Emphasis>

A trade-off between flight capability and reproduction is well known in adult females of the wing-dimorphic cricket Velarifictorus ornatus, but it is not clear whether such a trade-off exists in adult males of the species. In the present study, we investigated sexual maturation time, mating frequency, and the fertilization success of spermatophores after sequential mating in long-winged (LW) and short-winged (SW) adult males of V. ornatus to evaluate the potential reproductive advantage of the SW over the LW male morph. We found that the SW males of V. ornatus attained sexual maturity earlier and produced heavier spermatophores during the early stage after adult emergence than their LW counterparts. Additionally, within a 24-h mating period, the SW males showed a higher mating frequency, greater spermatophore weight, and shorter intermating time interval compared with their LW counterparts. Although females copulated with the two male morphs produced eggs of similar size, fertilization success by SW males was significantly higher than by the LW males. These results provide support for a trade-off between dispersal capability and reproduction success in wing-dimorphic males of V. ornatus.     

Genetic and diurnal variation in the juvenile hormone titer in a wing-polymorphic cricket: implications for the evolution of life histories and dispersal

The wing-polymorphic cricket, Gryllus firmus, contains (1) a flight-capable morph (LW(f)) with long wings and functional flight muscles, (2) a flightless morph with reduced wings and underdeveloped flight muscles (SW), and (3) a flightless morph with histolyzed flight muscles but with fully developed wings (LW(h)). The LW(f) morph differed genetically from the SW morph and phenotypically from the LW(h) morph in the size of flight muscles, ovarian growth during the first week of adulthood, and the hemolymph titer of juvenile hormone (JH). This is the first study to document that phenotypes that differ genetically in morphological aspects of dispersal capability and in ovarian growth also differ genetically in the titer of a hormone that potentially regulates those traits. The JH titer rose 9-100-fold during the photophase in the flight-capable LW(f) morph but did not change significantly during this time in either flightless morph. Prolonged elevation of the in vivo JH titer in flight-capable females, by topical application of a hormone analogue, caused a substantial increase in ovarian growth and histolysis of flight muscles. The short-term, diurnal rise in the JH titer in the dispersing morph may be a mechanism that allows JH to positively regulate nocturnal flight behavior, while not causing maladaptive histolysis of flight muscles and ovarian growth. This is the first demonstration of naturally occurring, genetically based variation for diurnal change in a hormone titer in any organism.     

Life-history evolution and the microevolution of intermediary metabolism: activities of lipid-metabolizing enzymes in life-history morphs of a wing-dimorphic cricket

Although a considerable amount of information is available on the ecology, genetics, and physiology of life-history traits, much more limited data are available on the biochemical and genetic correlates of life-history variation within species. Specific activities of five enzymes of lipid biosynthesis and two enzymes of amino acid catabolism were compared among lines selected for flight-capable (LW[f]) versus flightless (SW) morphs of the cricket Gryllus firmus. These morphs, which exist in natural populations, differ genetically in ovarian growth (100-400% higher in SW) and aspects of flight capability including the size of wings and flight muscles, and the concentration of triglyceride flight fuel (40% greater in LW[f]). Consistently higher activity of each enzyme in LW(f) versus SW-selected lines, and strong co-segregation between morph and enzyme activity, demonstrated genetically based co-variance between wing morph and enzyme activity. Developmental profiles of enzyme activities strongly paralleled profiles of triglyceride accumulation during adulthood and previous measures of in vivo lipid biosynthesis. These data strongly imply that genetically based elevation in activities of lipogenic enzymes, and enzymes controlling the conversion of amino acids into lipids, is an important cause underlying the elevated accumulation of triglyceride in the LW(f) morph, a key biochemical component of the trade-off between elevated early fecundity and flight capability. Global changes in lipid and amino-acid metabolism appear to have resulted from microevolutionary alteration of regulators of metabolism. Finally, strong genotype x environment (diet) interactions were observed for most enzyme activities. Future progress in understanding the functional causes of life-history evolution requires a more detailed synthesis of the fields of life-history evolution and metabolic biochemistry. Wing polymorphism is a powerful experimental model in such integrative studies.     

Morph‐specific differences in biochemical composition related to flight capability in the wing‐polyphenic Sitobion avenae

Flight performance at various times after emergence in the alate morph and age‐dependent changes in biochemical composition of winged and wingless morphs were evaluated in the wing‐polyphenic aphid Sitobion avenae (Fabricius) (Hemiptera: Aphididae). Alates exhibited the highest flight activity at 18–36 h after adult emergence. Throughout the nymphal and adult development, the whole‐body content of total lipid was significantly higher in the winged vs. wingless morph, whereas the content of water, soluble sugar, glycogen, phospholipid, and soluble protein showed significantly higher levels in the wingless vs. winged morph. There were no significant differences in the content of triglyceride and free fatty acid during nymphal and adult stages in both morphs. However, triglyceride content was significantly higher in the winged vs. wingless morph during adulthood. Differences in biochemical composition between morphs indicate that there is an energetic cost of flight capability. Our results from S. avenae adults showed that total lipid and triglyceride for the winged morph accumulated significantly to a maximum, and water content decreased significantly to a minimum, on days 1 and 2 after the final molt, exactly when the highest flight activity was reached. This study suggests that flight activity is associated with triglyceride and water content.