QUANTITATIVE GENETICS OF SEXUAL SELECTION IN THE FIELD CRICKET,GRYLLUS INTEGER

Major theories of sexual selection predict heritable variation in female preferences and male traits and a positive genetic correlation between preference and trait. Here we show that female Texas field crickets, Gryllus integer, have heritable genetic variation for the male calling song stimulus level that produces the greatest phonotactic response. Approximately 34% of the variation in female preferences was due to additive genetic effects. Female choosiness, that is, the strength of the female response to her most preferred stimulus relative to her average response to all stimuli, did not show significant genetic effects. The male calling song character was not related to male size or age but did show significant genetic effects. Approximately 39% of the variation in the number of pulses per trill was due to additive genetic variation. The genetic correlation estimated for the field population was 0.51 ± 0.17. The number of pulses per trill produced by males is under stabilizing sexual selection.     

家蝇卵巢中保幼激素结合蛋白的研究(英语)

本文运用活性炭结合试验测定了家蝇卵巢中保幼激素结合蛋白(JHBP)的含量.卵巢中JHBP对JHⅢ有较高的结合活性,其结合常数为2.1×10~(-8)M,~3H保幼激素Ⅲ(~3H-JH)和JHBP的结合可被未标记的JH Ⅲ抑制,但保幼激素的类似物ZR 512或ZR515均无抑制效应.卵巢中JHBP的含量在家蝇羽化后48小时达最大值,是羽化后60小时、72小时家蝇卵巢中JHBP含量的6.5倍和15.5倍.羽化后24小时、36小时的家蝇卵巢无JHBP的结合活性.若刚羽化的家蝇用JHⅢ处理,36小时后,JHBP则可检测到.本实验的结果表明卵巢中JHBP浓度的变动可能和JH对家蝇卵黄发生的作用有关.     

JUVENILE HORMONE BINDING PROTEIN IN THE OVARIES OF HOUSEFLY MUSCA DOMESTICA VACINA

Abstract  By using charcocal binding assay, the juvenile hormone binding protein (JHBP) was determined in the ovaries of houseflies. This ovarian JHBP possesses high affinity with juvenile hormone III (JH III) and has a K_d of 2.1 III 10^--⁸ M. The binding of ³H-juvenile hormone III (³H-JH III) to this protein was inhibited by unlablled JH III, but not by juvenile hormone analog ZR 512 or ZR 515. The level of this ovarian JHBP reached the highest in houseflies 48 h after emergence, and was 6. 5-fold and 15. 5-fold higher than that in housefIies 60 h and 72 h after emergence, respectively. No binding activity was detected in the ovaries of houseflies 24 h or 36 h after emergence. The absence of JHBP in the ovaries of houseflies 36 h after emergence could be reversed by applying JH III to newly emerged houseflies. The data suggest that the fluctuation of the JHBP concentration might associate with the action of juvenile hormone (JH) on housefly vitellogenesis.     

THE GENETIC BASIS OF THE TRADE-OFF BETWEEN CALLING AND WING MORPH IN MALES OF THE CRICKET GRYLLUS FIRMUS

Wing dimorphisms exist in a wide range of insects. In wing-dimorphic species one morph is winged has functional flight muscles (LW), and is flight-capable, whereas the other has reduced wings (SW) and cannot fly The evolution and maintenance of wing dimorphisms is believed to be due to trade-offs between flight capability and fitness-related traits. Although there are well-established phenotypic trade-offs associated with wing dimorphism in female insects, there only exist two studies that have established a genetic basis to these trade-offs. The present study provides the first evidence for a genetically based trade-off in male insects, specifically in the sand cricket Gryllus firmus. Because they have to expend energy to maintain the flight apparatus (especially flight muscles), LW males are predicted to call less and therefore to attract fewer females. To be of evolutionary significance, call duration wing morph, and wing muscle condition (size and functionality) should all have measurable heritabilities and all be genetically correlated. Differences between morphs in male G. firmus in the likelihood of attracting a female were tested in the laboratory using a T-maze where females chose between a LW male and a SW male. Call duration for each male was recorded on the sixth day of adult life. A significant difference in call duration was found between SW and LW males (SW = 0.86 ± 0.01, LW = 0.64 ± 0.01 h). SW males attracted significantly more females than did LW males (63% vs. to 37%). All the traits involved in the trade-off had significant heritabilities (call = 0 75 ± 0 33; wing morph = 0.22 ± 007; muscle weight = 0.38 ± 0.09) and genetic correlations (call and wing morph = -0.46 ± 0.20 for SW, -0.68 ± 0.16 for LW; LW call and muscle weight = -0.80 ± 0.14). These results provide the first documented evidence that trade-offs between a dimorphic trait and a fitness-related character in males has a genetic basis and hence can be of evolutionary significance.     

QUANTITATIVE GENETICS OF SHAPE IN CRICKET WINGS: DEVELOPMENTAL INTEGRATION IN A FUNCTIONAL STRUCTURE

The role of developmental and genetic integration for evolution is contentious. One hypothesis states that integration acts as a constraint on evolution, whereas an alternative is that developmental and genetic systems evolve to match the functional modularity of organisms. This study examined a morphological structure, the cricket wing, where developmental and functional modules are discordant, making it possible to distinguish the two alternatives. Wing shape was characterized with geometric morphometrics, quantitative genetic information was extracted using a full‐sibling breeding design, and patterns of developmental integration were inferred from fluctuating asymmetry of wing shape. The patterns of genetic, phenotypic, and developmental integration were clearly similar, but not identical. Heritabilities for different shape variables varied widely, but no shape variables were devoid of genetic variation. Simulated selection for specific shape changes produced predicted responses with marked deflections due to the genetic covariance structure. Three hypotheses of modularity according to the wing structures involved in sound production were inconsistent with the genetic, phenotypic, or developmental covariance structure. Instead, there appears to be strong integration throughout the wing. The hypothesis that genetic and developmental integration evolve to match functional modularity can therefore be rejected for this example.     

沙蟋飞行肌中保幼激素滴度的昼夜节律

沙蟋Gryllus firmus是一种翅多型性的昆虫, 是研究种内迁飞和生殖调控的模式生物。本研究应用高效液相色谱仪（HPLC）、 气相色谱-质谱联用仪(GC-MS)对沙蟋长翅（有飞行能力）和短翅（无飞行能力）雌虫飞行肌内保幼激素（juvenile hormone, JH）和脂肪酸进行了定性定量分析。结果表明： 在温度28℃, 光周期16L∶8D条件下, 第5和第7日龄的沙蟋长翅雌虫飞行肌中JH的滴度具有明显的昼夜节律, 在飞行前（即关灯前）4 h, JH的滴度分别由386.52±68.40 ng/g和630.36±37.26 ng/g增加至1 327.53±277.98 ng/g和1 685.77±143.95 ng/g, 与短翅型SW相比分别增加了约3.4倍与2.7倍 (P＜0.05)。而相同日龄的短翅雌虫及第1日龄的两型雌虫均无明显的节律变化。进一步在第5和7日龄的长翅雌虫中发现了一个16C的脂肪酸--14-甲基十五烷酸, 也具有节律变化且与JH节律出现的时间相吻合, 而在无飞行能力的沙蟋中没有发现这种现象。实验还证明JH滴度的增加和节律不是由飞行肌的重量或者飞行肌重量比的变化所致。这些发现有助于探讨和了解保幼激素对飞行调控的内在机理。     

Coevolution of traits determining migratory tendency: correlated response of a critical enzyme,juvenile hormone esterase,to selection on wing morphology

Migratory tendency in insects is a complex trait, composed of a suite of correlated behavioural, physiological, morphological and life history traits. We investigate the genetic and physiological basis of the coevolution of this suite of traits using laboratory lines of the wing dimorphic cricket, Gryllus firmus, selected for increasing and decreasing incidence of macroptery. Selection on wing morphology has produced strong direct responses in proportion macropterous as well as correlated (indirect) responses in wing muscle histolysis, flight propensity and fecundity. We investigate the hypothesis that these responses have been mediated by changes in the metabolism of juvenile hormone (JH) during the final nymphal stadium (the critical period for wing morph determination). Previous studies of Gryllus sp. have established that JH titre in this period is determined primarily by the activity of the degradative enzyme, juvenile hormone esterase (JHE). Assays of JHE activity in the final nymphal stadium of the replicated control and selected lines demonstrate highly significant differences in both mean activity and the probability of macroptery for a given level of activity (i.e., the threshold activity required to induce wing formation). These correlated responses in JH metabolism support the general hypothesis that the correlations among traits determining migratory tendency result at least in part from the common influence of JH during the final nymphal stadium. We discuss these results in the context of the quantitative genetic model for the evolution of polygenic, dichotomous traits (the threshold model), and present four general predictions concerning the coevolution of traits associated with ecological (i.e., trophic, life history, behavioural) dimorphisms.     

CHARACTERIZATION AND FUNCTIONAL STUDY OF A PUTATIVE JUVENILE HORMONE DIOL KINASE IN THE COLORADO POTATO BEETLE Leptinotarsa decemlineata (Say)

Juvenile hormone diol kinase (JHDK) is an enzyme involved in JH degradation. In the present article, a putative JHDK cDNA (LdJHDK) was cloned from the Colorado potato beetle Leptinotarsa decemlineata. The cDNA consists of 814 bp, containing a 555 bp open reading frame encoding a 184 amino acid protein. LdJHDK reveals a high degree of identity to the previously reported insect JHDKs. It possesses three conserved purine nucleotide‐binding elements, and contains three EF‐hand motifs (helix‐loop‐helix structural domains). LdJHDK mRNA was mainly detected in hindgut and Malpighian tubules. Besides, a trace amount of LdJHDK mRNA was also found in thoracic muscles, brain–corpora cardiaca–corpora allata complex, foregut, midgut, ventral ganglia, fat body, epidermis, and hemocytes. Moreover, LdJHDK was expressed throughout all developmental stages. Within the first, second, and third larval instar, the expression levels of LdJHDK were higher just before and right after the molt, and were lower in the intermediate instar. In the fourth larval instar, the highest peak of LdJHDK occurred 56 h after ecdysis. Ingestion of double‐stranded RNA (dsRNA) against LdJHDK successfully knocked down the target gene, increased JH titer, and significantly upregulated LdKr‐h1 mRNA level. Knockdown of LdJHDK significantly impaired adult emergence. Thus, we provide a line of experimental evidence in L. decemlineata to support that LdJHDK encodes function protein involved in JH degradation.     

THE RELATIONSHIP BETWEEN INSECT GROWTH REGULATORS WITH JUVENILE HORMONE ACTIVITY AND INSECT FEEDING BEHAVIOURAL RESPONSE

Abstract  Five insect growth regulators (IGRs) with juvenile hormone (JH) activity including fenoxy-carb, methoprene, NC-170, NC-184 and NC-196 were selected to study on the relationship between IGRs and feeding behaviour of the fourth instar larvae of Calospilos suspecta (Warren) and the fifth instar larvae of Ostrinia furnacalis Guenee, by topical and dietary applications as well as "diet column" methods. The results indicated that the antifeeding indices of methoprene, fenoxycarb, NC-170, NC-184, NC-196 and toosendanin against the fifth instar larvae of Ostrinia furnacalis in no-choice test were -37. 7, 14. 6, -12. 3, 10. 9, 14. 7 and 62. 2% respectively. Methoprene could stimulate the two species of insects to feed, whereas fenoxycarb had some antifeeding activity, and the effects of all the tested chemicals were different. Therefore, disturbances of IGRs with JH activity on insect feeding behavioural response may be one of the mechanisms of action not to be ignored. Moreover, by comparing "diet column" method with "faeces weight" method, it was suggested that the two methods might be applied in bioassay of antifeeding behaviour.     

具保幼激素活性昆虫生长调节剂与昆虫摄食行为反应的关系(英文)

本文以烯虫酯、双氧威、ＮＣ－１７０、ＮＣ－１８４和ＮＣ－１９６等具保幼激素活性的昆虫生长调节剂（ＩＧＲｓ）为药剂,以丝绵木金星尺蠖 Ｃａｌｏｓｐｉｌｏｓ ｓｕｓｐｅｃｔａ４龄幼虫和亚洲玉米螟 Ｏｓｔｒｉｎｉａ　ｆｕｒｎａｃａｌｉｓ５龄幼虫为试材,采用点滴法和混药饲喂法,结合非选择性饲料柱称重法,研究了ＩＧＲｓ与昆虫摄食行为之间的关系。结果表明,烯虫酯、双氧威、ＮＣ－１７０、ＮＣ－１８４、ＮＣ－１９６和川楝素对亚洲玉米螟５龄幼虫的非选择性拒食率依次为－３７．７、１４．６、－１２．３、１０．９、１４．７和６２．２％,烯虫酯刺激两种试虫取食,而双氧威则有一定的拒食作用,不同药剂表现出的影响不尽一致。因此,外源具保幼激素活性的活性物质对昆虫摄食行为反应的干扰破坏,可能是ＩＧＲs作用机理中不可忽视的重要方面之一。对饲料柱称重法和虫粪称重法进行了比较,证实两种方法结果相近,趋势一致,均可用于昆虫摄食行为的生测。     

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.     

THE EVOLUTION OF ALTERNATE MORPHOLOGIES: FITNESS AND WING MORPHOLOGY IN MALE SAND CRICKETS

Many organisms show distinct morphological types. We argue that the evolution of these alternate morphologies depends upon both fitness differences between morphs within each sex and the genetic correlation between sexes. In this paper, we examine the evolution of alternate morphologies using wing dimorphism in insects as a model system. Many insect species are wing dimorphic, one morph having wings and being capable of flight, the other lacking functional wings. While there is a well established trade-off in females between macroptery and reproduction, there are few data on the possible costs in males. We examine trade-offs between macroptery and life-history traits in male sand crickets, Gryllus firmus, and estimate the genetic correlation of wing dimorphism between the sexes. Macropterous males develop faster than micropterous males and are either larger or the same size depending upon rearing conditions. There is no difference in absolute or relative testis size at eclosion or 7 d thereafter. Finally, there is no difference between macropterous and micropterous males in relative success at siring offspring. Thus, with respect to the above traits, there are no costs associated with being winged in male G. firmus. It is possible that there may be a trade-off between calling rate and macroptery. A comparison of the relative frequency of macroptery between males and female across different orders of insects supports this hypothesis. The genetic correlation of wing dimorphism between the sexes is high (r₈ = 0.86), and hence the frequency of macroptery in males may be strongly influenced by selection acting on females.     

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。     

家蚕中的5-氧脯氨酸酶及保幼激素类似物对它的影响

本工作首次在家蚕Bombyx mori的马氏管、中肠、丝腺及脂肪体等组织中测到了γ-谷氨酰循环中一个关键酶——5-L-氧脯氨酸酶的活力.该酶以马氏管中活力最高,在蚕的中肠、血淋巴中均存在有游离的5-氧脯氨酸.观察了保幼激素类似物(JHA)处理后,家蚕中肠、丝腺和脂肪体中5-L-氧脯氨酸酶活力的变化,同时观察了血淋巴中5-氧脯氨酸含量的变化.对该酶及γ-谷氨酰循环在蚕体中氨基酸转运上的可能作用进行了讨论.     

保幼激素类似物对家蚕丝腺与脂肪体蛋白质合成的调节作用

应用示踪原子法,研究了家蚕Bombyx mori 5龄幼虫丝腺与脂肪体细胞内蛋白质合成的变化规律及保幼激素类似物(JHA738)的调节作用。从5龄初到龄末,家蚕丝腺细胞内蛋白质合成持续升高,5龄中期、后期的蛋白质合成活性分别是前期的1.60倍和2.86倍;全龄出现2个合成高峰,一个是在5龄72h,为细胞固有蛋白质合成高峰,另一个是在5龄192h,为丝蛋白合成高峰。脂肪体细胞内蛋白质合成作用呈现脉冲式的变化。在5龄前期和中期用JHA处理家蚕(剂量为4μg/条),对丝腺细胞固有蛋白质合成和脂肪体细胞蛋白质合成均表现出抑制作用,而对丝蛋白合成则表现出促进作用。本实验结果为进一步阐明JHA增丝机理提供了直接证据。     

THE EVOLUTION OF THRESHOLD TRAITS: A QUANTITATIVE GENETIC ANALYSIS OF THE PHYSIOLOGICAL AND LIFE-HISTORY CORRELATES OF WING DIMORPHISM IN THE SAND CRICKET

Many traits are phenotypically discrete but polygenically determined. Such traits can be understood using the threshold model of quantitative genetics that posits a continuously distributed underlying trait, called the liability, and a threshold of response, individuals above the threshold displaying one morph and individuals below the threshold displaying the alternate morph. For many threshold traits the liability probably consists of a hormone or a suite of hormones. Previous experiments have implicated juvenile hormone esterase (JHE), a degratory enzyme of juvenile hormone, as a physiological determinant of wing dimorphism in the crickets Gryllus rubens and G. firmus. The present study uses a half-sib experiment to measure the heritability of JHE in the last nymphal stadium of G. firmus and its genetic correlation with fecundity, a trait that is itself genetically correlated with wing morph. The phenotypic and genetic parameters are consistent with the hypothesis that JHE is a significant component of the liability. Comparison of sire and dam estimates suggest that nonadditive effects may be important. Two models have been proposed to account for the fitness differences between morphs: the dichotomy model, which assumes that each morph can be characterized by a particular suite of traits, and the continuous model, which assumes that the associated fitness traits are correlated with the liability rather than the morphs themselves. The latter model predicts that the fitness differences will not be constant but change with the morph frequencies. Variation in fecundity and flight muscle histolysis are shown to be more consistent with the continuous model. Data from the present experiment on JHE are inconclusive, but results from a previous selection experiment also suggest that variation in JHE is consistent only with the continuous model.     

家蚕中的5-氧脯氨酸酶及保幼激素类似物对它的影响

本工作首次在家蚕Bombyx mori的马氏管、中肠、丝腺及脂肪体等组织中测到了γ-谷氨酰循环中一个关键酶——5-L-氧脯氨酸酶的活力.该酶以马氏管中活力最高,在蚕的中肠、血淋巴中均存在有游离的5-氧脯氨酸.观察了保幼激素类似物(JHA)处理后,家蚕中肠、丝腺和脂肪体中5-L-氧脯氨酸酶活力的变化,同时观察了血淋巴中5-氧脯氨酸含量的变化.对该酶及γ-谷氨酰循环在蚕体中氨基酸转运上的可能作用进行了讨论.     

HORMONAL CONTROL OF THE VITELLOGENESIS IN THE JAPANESE OAK SILKWORM, ANTHERAEA YAMAMAZ (LEPIDOPTERA: SATURNIIDAE)

Abstract Effects of ecdysteroid and juvenile hormone (JH) on vitellogenesis of the Japanese oak silkworm, Antheraea yamami are reported in this article. After topical treatment with 20-hydroxyecdysone alone or JH analog (i.e. methoprene) alone and combined treatment with these two chemicals, vitellogenin (Vg) titers in the fat body and haemolymph at the pupal stage were mostly higher than those of the control, indicating that both ecdysteroid and JH exerted a promoting effect on the synthesis of Vg. In contrast, the Vg uptake was markedly inhibited by JH while stimulating effect of the ecdysteroid could be shown that vitellin (Vt) titer in the ovary was lower after methoprene treatments, but higher after 20-hydroxyecdyson treatments. Meanwhile, effects of these two hormones on Vg synthesis in the fat body were also tested with the incubation in vitro with Grace medium containing H-leucine and the hormones. The results demonstrated that Vg synthesis was stimulated after treating with methoprene alone or 20-hydroxyecdysone alone and combined treating with these two chemicals, and particularly ecdysteroid had more marked positive effect. To comprehensively concluded our results, it could be regarded that ecdysteroid play the main role in the regulation of vitellogenesis for the Japanese oak silkworm.