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.     

Colour-pattern variation in relation to habitat in the grasshopper Chorthippus brunneus (Thunberg)

Abstract. 1. Chorthippus brunneus were collected from twenty-two different sites in Nottinghamshire, Derbyshire and Essex.
2. Discrete characters of the head, pronotum and forewings and hind femora were scored. The results were compared with the types of habitat from which the grasshoppers were collected.
3. Associations between striped and mottle-winged varieties were found.
4. The relationships between colour and habitat were examined.     

Spatial vision in binocular and monocular common field grasshoppers (Chorthippus brunneus)

Abstract The ability of the common field grasshopper Chorthippus brunneus to discriminate between different distances under binocular and monocular stimulus conditions is investigated based upon the peering‐jump behaviour. The results show that information obtained from only one eye is sufficient for the grasshopper to determine the jump direction and distance. However, information obtained from both eyes is advantageous for relative distance determination. It is hypothesized that the motion parallax signals from the left and right eye may be summed, thus improving performance. There is no behavioural evidence of a more complex correlation of the information from the two eyes.     

Timing of life cycles in a seasonal environment: the temperature-dependence of embryogenesis and diapause in a grasshopper (Chorthippus brunneus Thunberg)

Summary The durations of the stages in the embryonic development of Chorthippus brunneus (Thunberg) have been estimated at constant temperatures of 20° C and 30° C. There was stage-dependent variation in the temperaturedependence of morphogenesis-a pattern apparently not previously described. The earliest and latest stages of morphogenesis proceeded relatively rapidly at higher temperatures. But intermediate stages, which would tend to occur around the diapause stage in a temperate winter, proceeded relatively rapidly at lower temperatures, as did diapause development. At a given temperature, there was individual variation between eggs such that some developed consistently faster than others. However, these intrinsic differences could be altered by exposure to diapause-breaking conditions at different morphological stages. The results are discussed in terms of their adaptive significance, their effect on population dynamics in a seasonal environment, and their implications for the nature of diapause in insects generally.     

Mating signal variation and bimodality in a mosaic hybrid zone between Chorthippus grasshopper species

The grasshoppers Chorthippus brunneus and Chorthippus jacobsi are easily distinguished by male calling song and the number of stridulatory pegs on the hind femur, and form a mosaic hybrid zone in northern Spain. In this paper, we fit a two-dimensional cline to variation in male calling song characters, which are of particular interest as they are likely to be involved in mate choice by females. As with variation in peg number, local habitat makes only a small contribution in explaining deviations in mean song score from clinal expectations. However, the fitted width of the cline for song characters is significantly narrower than for peg number, suggesting that mating signals may be associated with reduced hybrid fitness in the field and that recombination rates are sufficient to allow clines for different characters to diverge in width. Despite this, estimates for the overall elevation in linkage disequilibrium at the zone center, based on covariance between peg and song characters, reveal a substantial overrepresentation of parental genotypes at the cline center relative to the expectations of a tension zone of similar width. Examination of covariance at individual sites reveals that this inflated estimate of linkage disequilibrium is caused by several sites where the distribution of phenotypes is effectively bimodal. This substantial variation in linkage disequilibrium at the cline center could result from local variation in the strength of assortative mating or selection against hybrids, or may reflect the long-distance colonization of empty habitat from outside the hybrid zone, which would continually create new contacts between parental genotypes at the cline center. Hybrid zones like this, in which strong linkage disequilibrium occurs in some situations but not in others, are of particular relevance to speciation research and allow investigation of the spread of combinations of alleles through different genetic and ecological backgrounds.     

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.

     

Assortative preferences and discrimination by females against hybrid male song in the grasshoppers Chorthippus brunneus and Chorthippus jacobsi (Orthoptera: Acrididae)

The grasshoppers Chorthippus brunneus and Chorthippus jacobsi are highly differentiated for male mating signals, and form a mosaic hybrid zone in northern Spain. At some sites within this zone, many hybrids are observed. At others, few hybrids are observed. Such bimodal sites may reflect recent contacts between parental genotypes, or local variation in levels of assortative mating or selection against hybrids. Playback of 12 parental and F1 male songs to 296 parental and hybrid females revealed positive assortative preferences in C. brunneus and C. jacobsi females, supporting a direct role of male mating signals in female choice. However, all female genotypic classes showed reduced responsiveness to F1 male songs. Such sexual selection against hybrids is consistent with the narrow cline width observed in the field for song characters relative to morphology. These results have implications for the genetic structure of the hybrid zone and for models of speciation by reinforcement.     

Grasshopper populations and weather: the effects of insolation on Chorthippus brunneus

ABSTRACT.

• 1 In nature, the internal body temperature of Chorthippus brunneus Thunberg is found to be raised substantially under the influence of direct insolation.
• 2 In the laboratory, similarly raised body temperatures, resulting from the radiant heat of a light bulb, lead to a 5.6-fold increase in the rate of development, and a similar or even higher increase in the rate of adult maturation. The rate of egg-production is even more greatly affected, being negligible in the absence of a radiant-heat source.
• 3 These physiological results provide an explanation for the frequently found correlation between grasshopper abundance and hot and/or dry weather.
• 4 The results are also discussed in terms of the problems they pose for the derivation of a physiological time-scale for basking insects, and the evolutionary forces and constraints that might give rise to them.

     

Oviposition date and pattern of embryogenesis in the grasshopper Chorthippus brunneus (Orthoptera, Acrididae)

The morphological development of eggs of the grasshopper Chorthippus brunneus , laid during successive periods through a single season, was closely monitored in the field at a site in NW England. Eggs within the same clutch developed synchronously. However, there were significant differences between clutches laid at the same time, reflecting spatial variation in microclimate (and possibly maternal effects and/or genetic differences). Eggs laid early in the season entered diapause and hatched relatively synchronously in the following spring. Eggs laid later failed to reach the diapause-stage before winter, but hatched, albeit late and with greater variance. Diapause was averted in these eggs. Plasticity in the nature of diapause has been documented from laboratory studies of many insect species (including C. brunneus ,) but this study directly relates the incidence of such developmental plasticity to oviposition date and seasonal temperature cycles in the field. The adaptive significance of diapause aversion and the evolutionary consequences for female reproductive behaviour are considered.     

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.     

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.     

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.     

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.     

Pathfinding of peripheral neurons in the central nervous system of an embryonic grasshopper (Chorthippus biguttulus)

The peripheral leg nerves of grasshoppers are initially formed by a set of pioneer neurons and guidepost cells. These cells are used as guiding structures for later-arising axons of sensory neurons. The development of the central projections of the pioneer cells, the guidepost cells and some sensory cells is shown with Lucifer Yellow injection or with DiI application. The axons of the pioneer cells Ti1 enter the central nervous system at 38% of embryonic development. They turn anteriorly close to the midline and ascend with no major branching to the brain. The axons of the guidepost cells Fe1 and Tr1 follow the same path but do not ascend to the brain. Sensory axons of the subgenual organ and the femoral organ probably do not follow the central path pioneered by the former neurons. They end ipsilaterally in the respective thoracic neuromere, as is found in the adult.     

Within- and between-individual sequence variation among ITS1 copies in the meadow grasshopper Chorthippus parallelus indicates frequent intrachromosomal gene conversion

Sequencing multiple copies of the ITS1 region revealed the coexistence of two or more haplotypes within the genome of Chorthippus parallelus. Using a PCR-RFLP approach, the ITS1 numbers and frequencies of haplotypes present in each of 40 individuals were investigated, revealing a consistent lack of homogeneity. For each individual, the level of intra-individual variation was estimated from a sample of 20 ITS1 copies. The level of differentiation in haplotype frequency among individuals was then estimated by maximum likelihood using models based on the Dirichlet distribution. This confirmed the existence of significant levels of variation among individuals within each population studied. The most likely turnover mechanism that could generate this pattern of variation is gene conversion, operating at the intrachromosomal level. Furthermore, the discovery of linkage disequilibrium among the ITS1 haplotypes of C. parallelus suggests that intrachromosomal gene conversion occurs more frequently than interchromosomal recombination. Subspecies of C. parallelus showed significantly different haplotype distributions following about 0.5 Myr of divergence. With respect to the process of concerted evolution, we show that homogenization of repeats is slow relative to speciation, and the standing variation among individuals is sufficient for selection to operate.     

Genetic divergence in the Chorthippus parallelus species group (Orthoptera: Acrididae)

Specimens of Chorthippus parallelus parallelus, C. p. erythropus, C. montanus, C. curtipennis and C. dorsatus were examined electrophoretically. This analysis indicates that C. curtipennis is very closely related to C. p. parallelus and C.p. erythropus (Nei's I = 0.97) while C. montanus is more distantly related ( I = 0.88). The values are consistent with divergence of C. p. erythropus from C. p. parallelus since its separation by the Pyrenean ice sheet during the Pleistocene and of C. curtipennis since invasion of North America by C. parallelus at the same time. A much earlier divergence of C. montanus is implied.     

Ecotypic differentiation in the grasshopper Chorthippus brunneus: life history varies in relation to climate

Life history variations among 27 populations of the grasshopper Chorthippus brunneus from around the British Isles were examined under laboratory conditions over three generations. Multiple-regression analysis was used to examine the relationship between grasshopper life histories and the climates of their ancestral sites. Grasshoppers from cooler sites were heavier at hatching. Grasshoppers from northern sites grew faster and developed through fewer instars, attaining adulthood earlier, at the expense of adult size. Depending on the measure of adult size used, adults were larger in warmer, sunnier or more southerly locations. Ecotypic differentiation is probably widespread among animals as it is among plants, though it is more rarely demonstrated by zoological studies, especially over the wide geographical scale covered here. Evidence from regression analysis supports the hypothesis that ecotypic differentiation in C. brunneus is an evolutionary response to climatic variation. The existence of intraspecific genetic diversity for climatic adaptations has implications for biodiversity conservation and the understanding of biotic responses to climatic change. It deserves wider recognition. Received: 5 October 1998 / Accepted: 30 June 1999