Control of sexual behaviour pattern 'secondary defence' in the female grasshopper, Chorthippus curtipennis

The sexual behaviour of Chorthippus curtipennis females consists of ‘primary defence’, passive and active copulatory readiness, ‘secondary defence’, and egg-laying behaviour. Mechanical or chemical stimulation of the spermatheca by the spermatophore does not cause the release of ‘secondary defence‘; chemical stimuli deriving from the ovaries can also be excluded. ‘Secondary defence’ is triggered by the pressure of natural or artificial eggs against the walls of the lateral oviducts.     

Relationships between oviposition date, hatch date, and offspring size in the grasshopper Chorthippus brunneus

Abstract 1. The grasshopper Chorthippus brunneus has been shown to increase egg size with maternal age under constant laboratory conditions, such that late-laid eggs are larger than early laid eggs. In this study, an increase in the size of eggs (and hatchlings from those eggs) was recorded with date of oviposition in a field cage population. This suggests that the relationship between maternal age and egg size, observed previously in the laboratory, also occurs in the field. There was, however, some evidence that the behaviour of the maternal females in field cages was modified by the onset of autumnal weather at the end of the breeding season. Only a small number of eggs was laid in the last 3 weeks of the summer but these yielded relatively small hatchlings.
2. The date on which eggs were laid was correlated positively with their date of hatch in the following year. A consequence of this relationship, and that between oviposition date and egg size, was that size at hatch increased with date of hatch through most of the spring but then declined as the last few eggs hatched. Temporal variation in size at hatch parallelled temporal variation in the maternal females' investment in egg size in the previous year.
3. An undisturbed field population was monitored to assess whether the temporal variation in size at hatch resulted in size variation at eclosion in subsequent developmental stages. There were negative correlations between size at eclosion and date of eclosion at the third and fourth instars, suggesting that despite their smaller size at hatch, early hatchers experienced more favourable conditions for juvenile growth than late hatchers. The larger size at hatch of late hatchers may enhance their survival and compensate (albeit incompletely) for their reduced opportunity for growth. Female reproductive behaviour may represent an adaptive response to a predictable seasonal decline in offspring fitness at hatch.     

The genetics of colour-patterns in the grasshopper Chorthippus brunneus

Several alleles were found to determine the colour of the dorsal pronotum in Chorthippus brunneus; there was evidence for at least two loci (C and V). Brown (C^B)was the universal recessive and green (C^C) was dominant to all other colours. The white allele (C^W)was codominant with green(C^G)and purple (C^P). Wing-patterns were determined by a separate, probably linked locus (W). A dominant plain wing-pattern (W^P) was associated with colours other than brown. Striped(W^S)and mottled(W^mo) were codominant and a plain recessive allele (W^P) was also found. All three alleles were associated with the brown phenotype. A purple-sided allele (S^Pu) was sometimes obmd with C^pu.. S^Pu was dominant to brown sides (S^B), A series of markings on the dorsal and lateral pronotum (linea intermedia, fascia postocularis, linea media, carina media and zona lateralis) were investigated and found to be controlled at separate loci which may be linked to W. These characters were expressed by dominant alleles. Epistatic effects by modifier loci were shown to have an important effect on the determination of wing phenotype. Allele Wo⁺, for example, suppressed the stripe-wing pattern, linea media, carina media and zona lateralis. It was concluded that colour patterns appear to be under genetic control and that dominant alleles were rare in the wild. Changes in shades of colours were shown to be age-dependent and minor.     

The genetics of colour-patterns in the grasshopper Chorthippus brunneus

Several alleles were found to determine the colour of the dorsal pronotum in Chorthippus brunneus; there was evidence for at least two loci ( C and V ). Brown ( C^B )was the universal recessive and green ( C^C ) was dominant to all other colours. The white allele ( C^W )was codominant with green( C^G )and purple ( C^P ). Wing-patterns were determined by a separate, probably linked locus ( W ). A dominant plain wing-pattern ( W^P ) was associated with colours other than brown. Striped( W^S )and mottled( W^mo ) were codominant and a plain recessive allele ( W^P ) was also found. All three alleles were associated with the brown phenotype. A purple-sided allele ( S^Pu ) was sometimes obmd with C^pu.. S ^Pu was dominant to brown sides ( S^B ), A series of markings on the dorsal and lateral pronotum ( linea intermedia, fascia postocularis, linea media, carina media and zona lateralis ) were investigated and found to be controlled at separate loci which may be linked to W . These characters were expressed by dominant alleles.
Epistatic effects by modifier loci were shown to have an important effect on the determination of wing phenotype. Allele Wo⁺ , for example, suppressed the stripe-wing pattern, linea media, carina media and zona lateralis .
It was concluded that colour patterns appear to be under genetic control and that dominant alleles were rare in the wild. Changes in shades of colours were shown to be age-dependent and minor.     

Enzyme variation in the grasshopper Chorthippus brunneus (Thunberg)

Forty-two populations of the grasshopper Chorthippus brunneus from four different areas were scored for polymorphism at 15 enzyme loci. The areas sampled were Nottinghamshire, inland Essex, the Essex coast and Dovedale in Derbyshire. Five loci were found to be polymorphic (isocitrate dehydrogenase (2), malate dehydrogenase, glyceroI-3-phosphate dehydrogenase and phosphoglucomutase) but levels of heterozygosity were generally low. Differences in enzyme variation between different areas were demonstrated using multivariate analysis.     

Postglacial expansion and genome subdivision in the European grasshopper Chorthippus parallelus

A noncoding nuclear DNA marker sequence (Cpnl-1) was used to investigate subdivision in the grasshopper Chorthippus parallelus and deduce postglacial expansion patterns across its species range in Europe. Investigation of the spatial distribution of 71 Cpnl-1 haplotypes and estimation of levels of genetic differentiation (K_ST values) between populations and geographic regions provided evidence for subdivision of C. parallelus into at least five major geographic regions and indicated that the French form of C. parallelus originated after range expansion from a Balkan refugium, Further evidence for subdivision of C. parallelus between Italy and northern Europe suggests that the Alps may have formed a significant barrier to gene flow in this grasshopper.     

Population density, phenotype and mortality in the grasshopper Chorthippus brunneus

ABSTRACT.

• 1 The patterns of mortality in laboratory populations of the grasshopper Chorthippus brunneus Thunberg are examined at a range of densities and at different stages of nymphal development with particular reference to the phenotypes of nymphs that die or survive.
• 2 Total mortality during nymphal development rises from density-independence at the lowest densities to approximately compensating density-dependence at the highest densities.
• 3 The greatest proportion of total mortality occurs in the first two instars. The stage broadly covering the second instar is the ‘key-stage’ determining adult density. The mortality during these stages is largely density-independent. There is relatively little mortality during stages broadly covering the final instar, but it is density-dependent and also plays a significant part in determining final adult density.
• 4 As the mortality becomes more density-dependent, either with density itself or with cohort age, the smallest individuals become increasingly disproportionately prone to mortality.
• 5 Males, which are smaller, are more susceptible to mortality than females, but their susceptibility relative to their size is less than that of females.

     

Limited condition dependence of male acoustic signals in the grasshopper Chorthippus biguttulus

In many animal species, male acoustic signals serve to attract a mate and therefore often play a major role for male mating success. Male body condition is likely to be correlated with male acoustic signal traits, which signal male quality and provide choosy females indirect benefits. Environmental factors such as food quantity or quality can influence male body condition and therefore possibly lead to condition-dependent changes in the attractiveness of acoustic signals. Here, we test whether stressing food plants influences acoustic signal traits of males via condition-dependent expression of these traits. We examined four male song characteristics, which are vital for mate choice in females of the grasshopper Chorthippus biguttulus. Only one of the examined acoustic traits, loudness, was significantly altered by changing body condition because of drought- and moisture-related stress of food plants. No condition dependence could be observed for syllable to pause ratio, gap duration within syllables, and onset accentuation. We suggest that food plant stress and therefore food plant quality led to shifts in loudness of male grasshopper songs via body condition changes. The other three examined acoustic traits of males do not reflect male body condition induced by food plant quality.     

Testicular dysfunction in hybrids between parapatric subspecies of the grasshopper Chorthippus parallelus

Two subspecies of Chorthippus parallelus meet in a narrow hybrid zone in the Pyrenees. Laboratory Fl hybrids between the two races had atrophied testes and were virtually sterile. Backcross males had variable levels of dysfunction but were generally intermediate between Parental and Fl males; there were some reciprocal differences. No other effects on fertility and viability of males and females were apparent, and Fl males had normal accessory glands. These data provide another clear example of Haldane's rule and have implications for the structure of hybrid zones, the evolution of mating isolation and mating strategies.     

Chromosomal differentiation through an Alpine hybrid zone in the grasshopper Chorthippus parallelus

The grasshopper Chorthippus parallelus is genetically differentiated over its range into at least five major geographical subdivisions. Two of these subdivisions, designated as subspecies, meet and form a hybrid zone in the Pyrenees. These subspecies differ for a wide range of morphological, behavioural and chromosomal characters, which vary clinally across the zone. A further geographical subdivision exists within Italy. Here we present the first evidence for chromosomal divergence between populations of the grasshopper north and south of the Alps. Across two Alpine cols (Col de Larche, between France and Italy; Passo de Resia, between Austria and Italy), these populations differ in both structural and functional components of the X-chromosome. Northern Alpine individuals possess an active nucleolar organizing region (NOR) at the distal end of the X-chromosome and an associated region of heterochromatin (C-band). Both these features are absent from individuals from the south of the Alps. However, all individuals examined carry distally located rDNA on the X-chromosome. Clinal transition was examined in the distal C-band in transects through the two cols. The cline centres are roughly coincident with the tops of the cols. Both clines were of similar widths (Col de Larche, 21.88 km; Passo de Resia, 24.05 km), and therefore much wider than those for an X-linked distal C-band in the Pyrenean hybrid zone. This suggests that there are different selective pressures on the cytogenetic characters in the Alps. The results are discussed in the context of the historical population dynamics of the species in relation to the climatic changes associated with the Pleistocene ice ages.     

Population density, phenotype and reproductive output in the grasshopper Chorthippus brunneus

ABSTRACT. 1. The effects of population density on the reproductive output of individual female grasshoppers ( Chorthippus brunneus Thunberg) and the relationships between reproductive output and component elements of each female's phenotype, were investigated.
2. Reproductive output was primarily determined by the rate of egg-pod production. Increased density led to significant reductions in the rate of egg-pod production and reproductive output.
3. Exoskeleton size (reflecting nymphal experience) was positively correlated with clutch size at both high and low densities, but condition (reflecting adult experience) showed no such correlation. At high density, exoskeleton size was more strongly correlated with the rate of egg-pod production and reproductive output than condition. It is concluded that the nymphal aspects of size are more important than the adult aspects.
4. At low density, females with small exoskeletons compensated for their smaller clutches by producing egg-pods at a faster rate. Thus, there was no overall relationship between reproductive output and any of the phenotypic characteristics.
5. The effects of competition were weakly asymmetric in high density populations. The weakness of the asymmetry suggests scramble-like interactions for resources. It is concluded that not only are the effects of competition influenced by individual differences, but also that competition may reveal differences that would not otherwise be apparent.     

Recombination within extra segments: evidence from the grasshopper Chorthippus jucundus

A detailed analysis, using a modified silver staining technique which permits the visualization of the chromatid cores in bivalents at metaphase I, shows that some so-called chromosome associations are indeed true chiasmata. This study employs a supernumerary segment on the longest of the autosome pairs in the grasshopper Chorthippus jucundus that produces clear asymmetric bivalents in heterozygotes. Clear evidence of crossing over within the supernumerary segment explains part of the polymorphism of this segment, since unequal crossing over can produce different-sized segments. The origin of this segment and its mode of inheritance are also considered in the light of these results.     

Circadian rhythmicity of prolactin secretion in Huntington's chorea

Prolactin circadian secretion has been studied in 16 males and in 8 females with Huntington's Chorea, in order to evaluate changes of the hormone's titres and of their periodicity in a condition where alterations of neurotransmission have been documented. Prolactin levels have been found normal, enhanced and low levels have been observed in one male case. Sleep connected surge has been found to be present in all cases. A harmonic analysis of periodical changes has been performed by a Fourier's cosines series. The changes in the occurence of different harmonic frequencies in respect to normal controls are not significant. No correlation could be drawn among clinical indices, hormone levels and the feature of the occurring harmonics.     

Simple inheritance of color and pattern polymorphism in the steppe grasshopper Chorthippus dorsatus

The green–brown polymorphism of grasshoppers and bush-crickets represents one of the most penetrant polymorphisms in any group of organisms. This poses the question of why the polymorphism is shared across species and how it is maintained. There is mixed evidence for whether and in which species it is environmentally or genetically determined in Orthoptera. We report breeding experiments with the steppe grasshopper Chorthippus dorsatus, a polymorphic species for the presence and distribution of green body parts. Morph ratios did not differ between sexes, and we find no evidence that the rearing environment (crowding and habitat complexity) affected the polymorphism. However, we find strong evidence for genetic determination for the presence/absence of green and its distribution. Results are most parsimoniously explained by three autosomal loci with two alleles each and simple dominance effects: one locus influencing the ability to show green color, with a dominant allele for green; a locus with a recessive allele suppressing green on the dorsal side; and a locus with a recessive allele suppressing green on the lateral side. Our results contribute to the emerging contrast between the simple genetic inheritance of green–brown polymorphisms in the subfamily Gomphocerinae and environmental determination in other subfamilies of grasshoppers. In three out of four species of Gomphocerinae studied so far, the results suggest one or a few loci with a dominance of alleles allowing the occurrence of green. This supports the idea that brown individuals differ from green individuals by homozygosity for loss-of-function alleles preventing green pigment production or deposition.Subject terms: Quantitative trait loci, Quantitative trait     

Heritability estimates for characters under sexual selection in the grasshopper,Chorthippus brunneus

Mating success of male grasshoppers, Chorthippus brunneus, has previously been shown to be highest for intermediate, and lowest for extreme individuals with respect to at least one parameter of the male calling song and probably also one morphological character. The heritabilities of these and other song and morphological characters have been estimated in an experiment utilizing the North Carolina I design. Heritability estimates for song parameters are all low (<0·28). Estimates for morphological characters are higher, particularly for features of the stridulatory file, despite the fact that it was just these characters that were shown to be under stabilizing sexual selection. These results are related to sexual selection theory.     

The origins of premating reproductive isolation: testing hypotheses in the grasshopper Chorthippus parallelus

There are many proposed routes for the origin of premating reproductive isolation, but few systematic studies aimed at testing their relative importance. Accumulated information about the biogeographical history of the European meadow grasshopper, Chorthippus parallelus, has allowed us to make a planned series of comparisons among populations aimed at distinguishing the contributions of some of these hypotheses. We have compared the effects on assortative mating of long-term isolation in glacial refugia, founder events during postglacial colonization, and sympatry with a closely related species. A likelihood-based analysis allowed us to separate effects of variation in male and female mating propensity among populations from variation in mate choice leading to assortative mating. All three effects contributed significantly to the overall variation in mating pattern in a set of 21 pairwise comparisons among seven populations. Male cuticular composition, but not other candidate signals, was significantly associated with the level of assortative mating. Of the hypotheses for the origin of reproductive isolation, only the predictions of the founder hypothesis explained a significant amount of the variation in assortative mating. This does not rule out the possiblity that there may be some other explanation. Having established the pattern of divergence, it is possible to generate hypotheses that explain our results at least as well as the founder hypothesis. However, because many such post hoc hypotheses are possible, they cannot be tested with this dataset. On this basis, our results favor the hypothesis that some aspect of the colonization process tends to accelerate divergence in mating signals leading to premating reproductive isolation. This could be accomplished through any one of several mechanisms. Colonization involves many bottlenecks as new populations are established at the edge of the range by long-distance migrants. Genetic effects may be important, but these bottlenecks may also alter the conditions under which mates are found and chosen, as suggested by Kaneshiro. At the same time, the colonizing populations may encounter novel environmental challenges.