Flightless Morph Research Articles

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.

The hemolymph JH titer exhibits a large-amplitude, morph-dependent, diurnal cycle in the wing-polymorphic cricket, Gryllus firmus

The hemolymph juvenile hormone (JH) titer was measured in over 500 flight-capable and flightless, adult female Gryllus firmus at 3–6 h intervals during each of days 2–8 of adulthood. The flight-capable morph exhibited a large-amplitude daily cycle in the hemolymph JH titer, while the flightless morph exhibited a barely perceptible cycle. The JH titer cycle was observed on all days in the flight-capable morph, but the large amplitude cycle (>15–20 fold increase in mean titer; >100-fold increase in some individuals), began on day 5. For both the large and small amplitude cycles, the JH titer peaked near the end of the photophase–beginning of the scotophase. The hemolymph ecdysteroid titer did not exhibit a corresponding large amplitude daily cycle, although a low amplitude cycle (1–3-fold change) was seen in both morphs. The large magnitude rise in the JH titer in the flight-capable morph during the photophase was not due to decreased hemolymph volume or JH degradation. Daily cycles in the JH titer may be common, but may have gone unnoticed in other insect species due to restricted temporal sampling. Failure to identify these cycles can result in substantial errors in inferring biological roles for JH. Because JH regulates flight behaviors, morph-specific daily cycles in the JH titer may be especially common in dispersal-polymorphic insects, in which flight is restricted to one morph during a limited period of the day or night. However, because JH regulates numerous biological traits, analogous cycles may be common in insects exhibiting other types of complex (e.g. caste or phase) polymorphism, in which morphs differ in a biological characteristic that is restricted to a specific period of the photophase or scotophase.

The Endocrine Regulation of Wing Polymorphism in Insects: State of the Art, Recent Surprises, and Future Directions

The endocrine mechanisms controlling the development and reproduction of flight-capable (long-winged) and flightless (short-winged or wingless) morphs of wing-polymorphic insects have been intensively investigated. The "classical model," put forward in the early 1960s, postulates that morph-specific differences in development and reproduction are caused by variation in the titers of juvenile hormone (JH) and/or ecdysone. Despite decades of study, the importance of these hormones in regulating wing polymorphism in aphids and planthoppers remains uncertain. This uncertainly is largely a consequence of technical and size constraints which have severely limited the types of endocrine approaches that can be used in these insects. Recent studies in wing-polymorphic crickets (Gryllus) have provided the first direct evidence that the in vivo blood titers of juvenile hormone and ecdysone, and especially the activity of the JH regulator, juvenile hormone esterase, differ between nascent morphs. Morph differences are largely consistent with the classical model, although some types of data are problematic, and other explanations are possible. Adult morphs differ dramatically in the JH titer but titer differences are more complex than those proposed by the classical model. Detailed endocrine information is thus far available only for a few species of crickets, and the hormonal control of wing polymorphism for insects as a whole remains poorly understood. Future studies should continue to investigate the role of JH and ecdysteroids in morph development and reproduction, and should expand to include studies of morph-specific differences in hormone receptors and neurohormones.

Morph-dependent fatty acid oxidation in a wing-polymorphic cricket: implications for the trade-off between dispersal and reproduction

Although a considerable amount of information is available on the ecology and physiology of wing polymorphism, much less is known about the biochemical–genetic basis of morph specialization for dispersal versus reproduction. Previous studies have shown that the dispersing morph of the wing-polymorphic cricket, Gryllus firmus, prioritizes the accumulation of triglyceride flight fuel over ovarian growth, while the opposite occurs in the flightless morph during the first week of adulthood. In this study, we compared the in vivo rate of lipid oxidation between genetic stocks of flight-capable versus flightless morphs to determine the role of lipid catabolism in morph specialization for flight versus reproduction. During the first five days of adulthood, in the absence of flight, fatty acid oxidation was substantially lower in the dispersing morph relative to the flightless morph, when either radiolabeled acetate or palmitate was used as a substrate. Differences between the morphs in fatty acid oxidation were genetically based, occurred co-incident with morph-specific differences in triglyceride accumulation and ovarian growth, and were observed on a variety of diets. A genetically based trade-off in the relative conversion of palmitate into CO 2 versus triglyceride was observed in morphs of G. firmus. Decreased oxidation of fatty acid and increased biosynthesis of triglyceride, both appear to play an important role in flight fuel accumulation, and hence morph specialization for flight. Conversely, increased oxidation of fatty acid likely fuels the enhanced ovarian growth in the flightless morph. The results of the present study on fatty acid catabolism, and previous studies on triglyceride and phospholipid biosynthesis, provide the first direct evidence that genetically based differences in in vivo flux through pathways of intermediary metabolism underlie a trade-off between flight capability and reproduction—a trade-off of central importance in insects.

Evolution of Cryptic Flight Morph and Life History Differences During Host Race Radiation in the Soapberry Bug, <I>Jadera haematoloma</I> Herrich-Schaeffer (Hemiptera: Rhopalidae)

The colonization of an introduced plant by soapberry bugs in Florida has resulted in the rapid evolution of a distinctive “host race.” Differences in the synchrony and persistence of local seed crops may have favored interracial differentiation in flight/life history tactics. To advance previous work, we quantified seed availability in individual native and introduced host plants. We tested the prediction that the new host’s relatively discrete period of annual seed production has selected for a higher frequency of a short-winged, flightless morph with a briefer generation time. Contrary to that prediction, short-winged bugs were not more common in the derived race, but further investigation revealed the unexpected presence of a long-winged morph, that like the short-winged, lacks flight muscle and exhibits the same rapid life cycle advantages. Consistent with prediction, the derived race, descended from volant long-winged colonists, shows an abundance of this “cryptic” flightless morph. In total, four flight/life history morphs were discovered, including two additional long-winged types that either histolyze or retain flight muscle. The morphs differed in life history traits both within and between host races. Morph frequency may be influenced by evolution in correlated characters: we found that beak length, which has rapidly evolved to be shorter on the smaller fruits of the introduced host, is discontinuously distributed between flight morphs, being greater in long-winged bugs. This study shows complex, unpredicted evolutionary relations between a mobility character, a trophic character, and the life history.

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.

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 covariance 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 × 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.

Differential lipid biosynthesis underlies a tradeoff between reproduction and flight capability in a wing-polymorphic cricket.

The biochemical basis of life-history tradeoffs is a poorly studied aspect of life-history evolution. We used radiotracer and endocrine approaches to investigate the extent to which morphs of a wing-polymorphic insect differ in the biosynthesis of lipid classes important for dispersal capability vs. reproduction (ovarian growth). The flight-capable genotype of Gryllus firmus biosynthesized a greater amount of total lipid and triglyceride (main flight fuel), which was preferentially allocated to somatic tissue during early adulthood. By contrast, the flightless genotype biosynthesized a significantly greater amount of phospholipid (important in egg development), which was preferentially allocated to ovaries. Topical application of a juvenile-hormone mimic to the flight-capable morph caused it to express all aspects of lipid metabolism seen in the flightless morph. Differences in biosynthesis between morphs (i) occur coincident with 100-400% greater ovarian growth in the flightless morph, (ii) result from alterations of both de novo biosynthesis of fatty acid and downstream partitioning of fatty acids into triglyceride vs. phospholipid, and (iii) possibly result from genetically polymorphic hormonal regulators with negative pleiotropic effects on lipid biosynthesis and ovarian growth. The present study provides direct documentation of genetically based alterations of in vivo flux through pathways of intermediary metabolism leading to the differential production of end products central to the specialization of phenotypes for alternate life histories.

Enzymological and radiotracer studies of lipid metabolism in the flight-capable and flightless morphs of the wing-polymorphic cricket, Gryllus firmus

The flight-capable morph of the wing-polymorphic cricket, Gryllus firmus, exhibited significantly higher activities of each of five lipogenic enzymes compared with the obligately flightless morph on a standard and a high-carbohydrate diet during early adulthood. Similarly, the rate of incorporation of [ 14 C] -acetate into total lipid was higher in the flight-capable morph during this time. By contrast, activities of lipogenic enzymes and rates of lipid biosynthesis, in general, did not differ between morphs on a low nutrient diet during early adulthood. Differences in lipid biosynthesis account for previously documented differences in lipid reserves between morphs on some, but not all, diets. Results of the present and previous studies indicate that increased lipid biosynthesis in the flight capable morph on standard and high-carbohydrate diets constitutes an important adaptation for flight (production of lipid flight fuel). Lipid biosynthesis is negatively correlated with ovarian growth, and may be an important biochemical component of the trade-off between flight capability and ovarian growth in G. firmus. Morphs also differed in activities of three enzymes of lipid catabolism. However, the extent to which variation in activities of these enzymes between morphs results in variation in lipid catabolism is unclear. Finally, the flight-capable morph had a substantially higher activity of alanine aminotransferase in the fat body. Amino acids may be utilized for lipid biosynthesis or energy production to a greater degree in the dispersing morph compared with the oligately flightless morph. This study is the first to document differences in intermediary metabolism that underlie adaptations of morphs of a dispersal-polymorphic species for flight vs. egg production.

The metabolic basis of life history variation: genetic and phenotypic differences in lipid reserves among life history morphs of the wing-polymorphic cricket, Gryllus firmus

The flight-capable morph of the wing-polymorphic cricket, Gryllus firmus, accumulated a substantially greater quantity of total lipid and triglyceride, compared with the obligately flightless morph, during the first five days of adulthood. Increased lipid accumulation in the flight-capable morph was genetically based, and was produced when ovarian growth is substantially reduced in that morph. Temporal changes in lipid levels suggest that the higher triglyceride reserves in the flight-capable morph fed a high-nutrient diet were produced by elevated lipid biosynthesis. By contrast, on a low-nutrient or high carbohydrate diet, increased lipid levels in the flight-capable morph appeared to result primarily from decreased lipid utilization. Increased biosynthesis or retention of triglyceride (the major flight fuel in Gryllus) by the flight-capable morph may significantly divert nutrients from egg production and hence may be an important physiological cause of its reduced ovarian growth. The obligately flightless morph allocated a greater proportion of total lipid to phospholipid than did the flight-capable morph. No functionally-significant differences in total lipid or triglyceride were produced between morphs during the last nymphal stadium. A second flightless morph, derived from the flight-capable morph by histolysis of flight muscles during adulthood, also had reduced amounts of total lipid and triglyceride but increased ovarian growth compared with the flight capable morph on the standard (high-nutrient) diet. Important qualitative and quantitative aspects of lipid metabolism differ genetically between the flight-capable and flightless morphs of G. firmus and likely contribute importantly to their respective adaptations for flight capability vs. reproduction. This is the first study to document genetically-based differences in energy reserves between morphs of a complex (phase, caste, flight) polymorphism in which morphs also differ genetically in key life history traits.

BENEFITS OF DISPERSAL IN PATCHY ENVIRONMENTS: MATE LOCATION BY MALES OF A WING-DIMORPHIC INSECT

Dispersal dimorphisms, in which both flight-capable and flightless adults occur in the same species, are commonplace in insects. Such dimorphisms are seen as reflecting a balance between the benefits and costs of flight and wing reduction or loss. In heterogeneous habitats, theory predicts that fitness trade-offs can favor the evolution of a dispersal dimorphism in which both flight-capable and flightless morphs are retained in the same population. Despite the wealth of theory, however, there has never been an explicit field assessment of how habitat heterogeneity directly influences the reproductive success of the flight-capable and flightless wing forms of an insect species. The objective of this research was to investigate how variation in habitat heterogeneity (vegetation structure) and female density influence mate location and thus the potential for reproductive success by the male wing forms of a salt marsh inhabiting insect, the planthopper Prokelisia dolus. By placing unmated females in the field at different densities and in sparse and contiguous vegetation, we were able to compare the ability of male wing forms to locate stationary mates. Our data show that both vegetation structure and female density differentially influenced the ability of the male wing forms to locate mates. Flight-capable males located females and acquired matings far more frequently in sparse vegetation, whereas flightless males discovered females more often in contiguous vegetation. Flight-capable males located females more effectively at low female densities, whereas flightless males discovered females more efficiently at higher female densities. Thus, natural variation in vegetation structure and female density are two important factors which combine to influence the mating success of each male wing form. We conclude that habitat heterogeneity in concert with the known reproductive penalties imposed by flight capability (reduced siring ability) interact to favor the persistence of the dispersal polymorphism in males of this planthopper species.

Benefits of Dispersal in Patchy Environments: Mate Location by Males of a Wing-Dimorphic Insect

Dispersal dimorphisms, in which both flight-capable and flightless adults occur in the same species, are commonplace in insects. Such dimorphisms are seen as reflecting a balance between the benefits and costs of flight and wing reduction or loss. In heterogeneous habitats, theory predicts that fitness trade-offs can favor the evolution of a dispersal dimorphism in which both flight-capable and flightless morphs are retained in the same population. Despite the wealth of theory, however, there has never been an explicit field assessment of how habitat heterogeneity directly influences the reproductive success of the flight-capable and flightless wing forms of an insect species. The objective of this research was to investigate how variation in habitat heterogeneity (vegetation structure) and female density influence mate location and thus the potential for reproductive success by the male wing forms of a salt marsh inhabiting insect, the planthopper Prokelisia dolus. By placing unmated females in the field at different densities and in sparse and contiguous vegetation, we were able to compare the ability of male wing forms to locate stationary mates. Our data show that both vegetation structure and female density differentially influenced the ability of the male wing forms to locate mates. Flight-capable males located females and acquired matings far more frequently in sparse vegetation, whereas flightless males discovered females more often in contiguous vegetation. Flight-capable males located females more effectively at low female densities, whereas flightless males discovered females more efficiently at higher female densities. Thus, natural variation in vegetation structure and female density are two important factors which combine to influence the mating success of each male wing form. We conclude that habitat heterogeneity in concert with the known reproductive penalties imposed by flight capability (reduced siring ability) interact to favor the persistence of the dispersal polymorphism in males of this planthopper species.

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.

The endocrine-genetic basis of life-history variation: the relationship between the ecdysteroid titer and morph-specific reproduction in the wing-polymorphic cricket Gryllus firmus.

The hormonal basis of variation in life-history traits is a poorly studied topic in life-history evolution. An important step in identifying the endocrine-genetic causes of life-history variation is documenting statistical and functional associations between hormone titers and genotypes/phenotypes that vary in life-history traits. To this end, we compared the blood ecdysteroid titer and the mass of the ovaries during the first week of adulthood among a flight-capable morph and two flightless morphs of the wing-polymorphic cricket Gryllus firmus. Ecdysteroids are a group of structurally related hormones that regulate many important aspects of reproduction in insects. Both the ecdysteroid titer and ovarian mass were significantly higher in each of two flightless morphs compared with the flight-capable morph throughout the first week of adulthood. Genetically based differences in the ecdysteroid titer and ovarian mass between morphs from different selected lines were similar to phenotypically based differences among morphs from the same control (unselected) lines. By day 7 of adulthood, ovaries were typically 200-400% larger and the ecdysteroid titer was 60-300% higher in flightless versus the flight-capable morph. In addition, highly significant, positive, phenotypic correlations were observed between the ecdysteroid titer and ovarian mass in pooled samples of the two flightless and flight-capable crickets from control lines or from selected lines. The ecdysteroid titer was sufficiently elevated in the flightless morphs to account for their elevated ovarian growth. This is the first direct documentation that naturally occurring phenotypes/genotypes that differ in early fecundity, a key life-history trait, also differ phenotypically and genetically in the titer of a key reproductive hormone that potentially regulates that trait.

Nutrient absorption and utilization by wing and flight muscle morphs of the cricket Gryllus firmus: implications for the trade-off between flight capability and early reproduction

Absorption efficiency (AD, approximate digestibility, assimilation efficiency) of various macronutrients and conversion of absorbed nutrients to biomass (ECD) were compared among the two types of flightless morph and the flight-capable morph of the cricket, Gryllus firmus. No biologically significant phenotypic or genetic difference in AD for carbohydrate, protein or lipid was observed among morphs fed either a high-nutrient (100%) or a low-nutrient (25%) diet. Thus, previously-documented differences among adult morphs in carbohydrate and lipid content must be caused by processes other than variation in nutrient absorption by morphs during adulthood. Relative absorption efficiency of total dry mass of food by morphs of G. firmus appears to be a valid indicator of relative AD of total calories. Morphs did not differ phenotypically or genetically in the excretion of end products of nitrogen metabolism (uric acid, hypoxanthine plus xanthine) on either the high nutrient or the low nutrient diet. Nutritional indices corrected for excreted nitrogenous metabolites were very similar to uncorrected indices, and the pattern of variation among the morphs was the same for corrected or uncorrected values. Each of the two types of flightless morph converted a greater proportion of absorbed nutrients into body mass, mainly ovaries, and allocated a smaller proportion of assimilated nutrients to respiration than did the flight-capable morph. Moreover, the trade-off between respiration and early reproduction was substantially magnified on the low nutrient diet. These results extend previous findings of a trade-off between flight capability and early reproduction in wing-polymorphic Gryllus species (1) to diets of very different nutrient quantity, and (2) to flightlessness arising from different causes: blockage of flight muscle development in juveniles vs histolysis of fully-developed flight muscles in adults.

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.

Predicting correlated responses in natural populations: changes in JHE activity in the bermuda population of the sand cricket, gryllus firmus

Quantitative genetic methods have been used to examine selection responses in domesticated organisms but there are few cases of their application to predict changes in natural populations: there are, to our knowledge, no cases in which correlated responses to selection have been predicted. In the present paper we use quantitative genetic parameters estimated from a half-sib experiment to predict the changes expected in juvenile hormone esterase (JHE) activity in the Bermuda population of the wing dimorphic cricket, Gryllus firmus. JHE activity is genetically correlated with wing form in this cricket and hence changes in the proportion of macroptery (volant morph) are predicted to bring about a correlated response in JHE activity. The Bermuda population has a higher proportion of macropterous individuals (95%) compared to the stock (35%, originally from Florida) from which the heritabilities and correlations were estimated. The quantitative genetic analysis makes three predictions which were tested both qualitatively and quantitatively. In all cases the null hypothesis that the observed results correspond to those predicted cannot be rejected. As predicted, in the Bermuda population there is: (i) an increase in the population mean JHE activity; (ii) a leftwards shift in the curve relating the probability of microptery (flightless morph) to JHE activity; and (iii) a decrease in the mean JHE activity within each morph.

Biochemical aspects of flight and flightlessness in Gryllus: flight fuels, enzyme activities and electrophoretic profiles of flight muscles from flight-capable and flightless morphs

Female Gryllus assimilis subjected to 4.5–7.7 h continuous tethered flight had significantly lower amounts of total lipid, triglyceride and total soluble carbohydrate compared with unflown controls. A much greater amount of total lipid (6.3 mg) was used during flight compared with carbohydrate (0.14 mg). Flown individuals also had substantially reduced amounts of injected, radiolabeled [ 14C]-oleic acid. Activities of lipid, carbohydrate and amino acid catabolizing enzymes in flight muscles of G. assimilis and its wing-polymorphic congener, G. firmus, were very similar to activities in insects which primarily utilize lipid to power flight. By contrast, enzyme activities were very different from those in insects which primarily or exclusively use carbohydrate or proline as a flight fuel. These results strongly implicate lipid as the major flight fuel in Gryllus. Previous studies have shown that lipid levels are higher in flight-capable (long-winged) G. firmus that have small ovaries compared with flightless (short-winged) females that have large ovaries. Results of the present and previous studies collectively indicate that elevated lipid in long-winged G. firmus represents an energetic cost of flight capability which reduces (trade-offs with) reproduction in Gryllus. In G. firmus, mass-specific activities of nearly all enzymes were considerably reduced in underdeveloped, and to a lesser degree in histolyzed muscle, compared with fully-developed flight muscle. An important exception was alanine aminotransferase, whose activity was the highest in histolyzed muscle, and which may be involved in the catabolism of amino acids derived from muscle degradation. Despite the dramatic differences in enzyme activity, electrophoretic profiles of soluble flight-muscle proteins differed only subtly between fully-developed and underdeveloped or histolyzed flight muscles.

Antagonistic and reinforcing pleiotropy: a study of differences in development time in wing dimorphic insects

AbstractMorphological dimorphisms are found in many different taxa. Wing dimorphism in insects, in which some individuals possess wings and associated flight muscles and are thus volant while others lack a functional flight apparatus and are thus flightless, is a typical example of such types of dimorphisms. It has been extensively studied and such studies have demonstrated that the volant form, although possessing the advantage of flight capability, suffers a fitness cost in a delay in the onset of reproduction after emergence into the adult form and a reduced fecundity. Previous comparative analyses have suggested that there is no consistent trend for development time (hatching to adult) to differ between the two morphs. The present study analyses the phenotypic and genetic correlations between development time and wing morph in the cricket Gryllus firmus. It is shown that the macropterous (volant) morph develops faster than the micropterous (flightless morph). This trade‐off is manifested at both thephenotypic and genetic level. Further, a comparative analysis shows that the same phenotypic trade‐off is generally found in other Orthopteran species so far studied, but in other orders the micropterous morph develops faster. Provided that the phenotypic trade‐off is genetically based, in the Orthoptera the fitness advantage of the earlier onset of reproduction in micropterous females is offset by the extended development time (antagonistic pleiotropy). However, in other orders there is reinforcing pleiotropy in that the micropterous females develop faster and reproduce sooner than the macropterous morph. These results highlight the complexity of fitness interactions and the need to study a phenomenon across several taxa.

EVIDENCE THAT THE MAGNITUDE OF THE TRADE-OFF IN A DICHOTOMOUS TRAIT IS FREQUENCY DEPENDENT.

Many traits, such as wing dimorphism, paedomorphosis, and cyclomorphosis vary dichotomously. Such dimorphisms are maintained in part because of a trade-off between components of fitness: for example, in insects, the flightless morph cannot migrate but has a greater fecundity than the flight-capable morph. Several recent theoretical studies have analyzed the evolution of dichotomous traits, assuming that each morph can be characterized by a discrete syndrome of characters. Consideration of the genetic basis of dimorphism suggests that this assumption is incorrect. In this paper, I report a test of this assumption using the wing-dimorphic sand cricket, Gryllus firmus. It is predicted that, rather than remaining constant, the fecundity of macropterous (long-winged, flight-capable) females will decrease as the proportion of macropterous females in the population or family increases. This prediction is supported by fecundity data from lines selected for high and low proportions of macroptery and by sib analysis. Thus, models that seek to predict the evolution of dichotomous traits should take into account the likelihood that values of components of fitness, such as fecundity, may be related to their frequencies in the population.