Body size in the field cricket Gryllus integer (Orthoptera : Gryllidae) :heritability and male mating success

The influence of male body weight on the aggressive and mating behaviour of male Gryllus integer was studied under laboratory conditions. The relationship between adult age and weight was first determined; female weight increased and male weight decreased with age. Virgin males that had been isolated since the adult molt were paired for similar age and a difference in weight of greater than 200 mg. Paired males and a virgin female were observed in a glass arena for 24 minutes or until a mating occurred. Larger males mated significantly more often than smaller males. Larger males attacked more often, were more successful in aggressive encounters and had more contact with the female. Males that did not mate had lower rates of courtship and mounts than males that mated. Females in trials that did not result in a mating were signifcantly heavier than females in trials that resulted in a mating. Larger males that mated were significantly closer in weight to the weight of the female than larger males in trials that did not result in a mating. Larger males in trials that did not result in a mating had higher rates of aggressive stridulation than larger males that mated. Male weight is therefore important in mating success; fitness traits should theoretically show low genetic variability. However, significant heritability values were found for live weight, dry weight, head width, pronotum width and length, hind femur length and forewing length when estimated from the regression of offspring on mid-parent values, offspring and female and male values separately and full-sib correlations. The heritability of hind femur width was significant when estimated from the regression of offspring on male parent and from full-sib correlations. Heritability estimates of forewing length were significantly higher when estimated from the regression of offspring on female parent than when estimated from the regression of offspring on male parent. High phenotypic, genetic and environmental correlations were found between all pairs of traits. Data on male mating success and the heritability of fitness traits were discussed in terms of the maintenance of genetic variability.

Mating behaviours of the field cricket (Gryllus integer) at different levels of male density

The reproductive behaviour of the field cricket, Gryllus integer, was systematically observed in indoor arenas to determine the extent of female Choice and male-male competition at different sex ratios representing two male densities (12:6 and 6:6). The costs and benefits to males and females in those two densities were analyzed according to the theory of the evolution o£ leks. Observations were conducted during the dark hours when most calling occurred since hourly rates of courtship song and mating did not fluctuate significantly over a 24 h period. Female mating rates were not significantly different between densities, therefore males at high densities were not advantaged because of increased female tendencies to mate when social stimulation was increased. Mean rates of acoustical signalling (calling and courtin"g) did not differ significantly between densities. Mean rates of fighting by males at the high density were significantly greater than those of males at the low density. Mating benefits associated with callin~courting and fighting were measured. Mating rates did not vary with rates of calling at either density. Calling was not a prerequisite to mating. Courtship song preceded all matings. There was a significant power fit between male mating and courting rates, and male mating and fighting rates at the low, but not at the high, density. Density differences in the benefits associated with increased courting and fighting may relate, in part, to greater economic defensibility and monopoly of females due to reduced male competition at the low density. Dominant males may be preferentially chosen by females or better able to monopolize mating opportunities than subordinate males. Three criteria were used to determine whether dominant males were preferentially chosen by females. The number of matings by males who won fights (within 30 min of mating) was significantly greater than the number of matings by males who were defeated in such fights. Mating rates did not vary significantly with rates of winning at either density. There was a significant power fit between male mating rates and the percentage of fights a male won (irrespective of his fighting-frequency) at the low density. The mean duration a male guarded the female after mating did not vary significantly between densities. There was a significant linear relationship between the duration a spermatophore was retained and the duration a male guarded the female after mating. Courtship song apparently stimulated spermatophore removal. Male guarding involved inter-male aggression and reduced courtship attempts by other males. Males at the high density received no apparent reproductive benefits associated with increased social stimulation. Conclusive evidence for preferential choice of males by females, using the criteria examined here, is lacking. Males at the lower density had fewer competitors and could monopolize females more effectively.

Attractive silences in cricket courtship song : mate choice using a short-range signal /

Matings systems using signals for sexual communication have been studied extensively and results commonly suggest that females use these signals for locating males, species-identification, and mate choice. Although numerous mating systems employ multiple signals, research has generally focused on long-range signals perhaps due to their prominence and ease of study. This study focused on the short-range acoustic courtship song of crickets. The results presented here suggest this signal is under selection by female choice. Females mated preferentially with males having shorter silences between the two types of ticks within the song. The length of these silences (Gap 1) was correlated with male condition such that males having long silences were significantly lower in mass with respect to body size when compared to males having short silences. Both Gap 1 length and male condition were significantly repeatable within males over time suggesting the possibility these traits have a genetic basis. This study is the first empirical study to test female preferences within the natural variation of the courtship song. It now appears, at least in crickets, that both the longand short-range signals of a multi-signal mating system may contribute to male mating success.

Sexual selections in the northern fall field cricket, Gryllus pennsylvanicus: a manipulation of the adult sex ratio

The operational sex ratio has long been considered an important constraint on the structure of mating systems. The effects of an experimentally manipulated sex ratio on mating behavior and selection were investigated in a polygynous species, Gryllus pennsylvanicus, where the potential exists for spatial/temporal fluctuations in sex ratio of field populations. Four different sex ratios (males: females, 5:0, 5:2, 5:5, 5:10) were investigated. Observations were conducted in late summer over two field seasons, from 2400 h , to 1000 h EST. Several male characters thought to be associated with male reproduc.tive success were studied: calling duration, searching distance, weight, fighting behavior, courtship frequency, and mating success. Variance in male mating success was used as the indicator for the opportunity for sexual selection. Total selection was estimated as the univariate regression coefficient between relative fitness and the character of interest, while direct selection was estimated as standardized partial regression coefficients generated from a multiple regression of relative fitness on each character. The opportunity for sexual selection was highest at 5:2 and lowest at 5:10. The frequency of fighting behavior was highest at 5:2 and 5:5. Fighting ability (% wins) was determined to be an important correlate of male body weight. Direct selection for increased male body weight was detected at 5:2, while total selection for body weight was seen at 5:5. Selection on male body weight was not detected at 5: 10. Calling duration decreased as sex ratio became more female-biased. Total and direct selection were detected for increased calling at 5:2, only total selection for calling was seen at 5:5, whereas direct selection against calling was detected at 5: 10. Searching distance also decreased as sex ratio became more female-biased, however no form of selection was detected for searching at any of the sex ratios. Data are discussed in terms of sexual selection on male reproductive tactics, the mating system and maintenance of genetic variation in male reproductive behavior.

Effects of female postmating odour on male sexual behaviour, in Heliconius butterflies /

The effects of the female postmating odour on male sexual behaviour were examined in Heliconius erato and H. charithonia (Lepidoptera: Nymphalidae). Predictions from the antiaphrodisiac hypothesis were tested using the two reproductive strategies of these species. Within the pupal mating strategy, results from behavioural experiments quantified and statistically tested dispersal rates of pupal-perched males to the presence of stimuli with and without the postmating odour. Results do not support an antiaphrodisiac function to the postmating odour. Similarly, within the adult courtship strategy, behavioural test results indicate that males do not alter their expenditure of energy in terms of either the duration or frequency of courtship behaviours elicited by females with and without the postmating odour. The data from both experiments did not support the antiaphrodisiac hypothesis for the function of the female postmating odour. A novel hypothesis predicting that the postmating female odour acts as an oviposition-deterring pheromone is presented.

Individual differences and factors affecting male behaviour in field crickets

Individual differences in male sexual behav~our and the factors influencing calling behaviour were studied in the field crickets Gryllus 2 integer and Q. veletis. In a large (13m) outdoor arena individually numbered adult male ~~ integer started calling at three to five days of age but thereafter the age of individual G. integer males did not affect nightly calling duration. Calling also did not correlate with individual weight. In this study individual male calling was continuously distributed from 0 hrs. per night to 3.5 hrs. per night, on average. A temporal effect on the number of G. integer males calling was observed. The number of males calling through the night was uniform, but a sharp increase in the number calling was observed in the early morning. No difference in calling times was observed between the night and dawn callers. AlsC)' males calling at dawn usually didnotc'all during the preceeding night. Calling and reproductive success in 1979 demonstrated a negative logarithmic relationship while in the 1980(initial) population a negative linear relationship was observed. No relationship was seen in the 1980 high density population. The ratio of non-callers to callers also affected the mating of individuals in the 1979 and1980(initial) densities:-non~callers (males calling .5 hrs. per night, on average, or less) obtained more females when the population contained a high number of callers, this being a negative logarithmic relationship to, No such relationship was observed in the 1980 high density population. Individual displacement varied nightly and was not correlated to amount of calling or reproductive success of individual G. integer males. G. integer males were displa~ed more when in a higher density in the outdoor arena Male G. integer and G. veletis behaviours were also observed in an indoor arena at different densities and, in G. veletis, with respect to female presence. When females were present in the arena, in G. veletis, male calling was reduced. Males of both species called less, on average, when in ~ higher density, than when they were in a lower density. Male displacement of both species increased on average when in a higher density as compared to displacement in a lower density. Aggression was measured by aggressive call-ing and fighting and was studied in regards to density.G. integer demonstrated less aggression in all but one comparison at higher density. No difference was observed in the ratio of aggressive calling to f.ighting comparison in G. integer. G. veletis demonstrated mixed results. No difference in aggression between densities was observed in comparisons. Less.aggression did occur in higher densities when comparisons invol.ved fighting behaviour. Male behaviour represents a competitive strategy against ot~er males, strategy being defined as a genetic (in part) alternative to other strategies. In this sense, the factors of time, density, male-male aggression, and female presence are conditions demonstrated to affect male behaviour in G. integer and G. veletis. Individual male differences and other considerations suggest that alternative male behaviours are represented by at least two conditional strategies. This possibility, and the transient 'or stable nature of genetic polymorphisms in field cricket behaviour are considered.

The effect of social condition and diet on the frequency of male-male agonistic displays in the field cricket, gryllus bimaculatus /

The frequency and type of agonistic displays involved in male-male encounters should be significantly influenced by the presence of females. Discrete agonistic displays vary in energy expenditure and risk, and therefore should be dependent on available resources. The influence of live females and the scent of females, on the frequency of male agonistic displays was observed in a laboratory terrarium using the field cricket Gryllus bimaculatus. The effect of energy constraints on display frequency was also determined. Half the males were fed a diet high in protein and fet; the other males were fed a lower quality diet, for a 7-11 day period. The frequency of five individual displays and mating frequency were recorded using an Event Recorder and notebook. Each group of males was presented with three experimental conditions, over three days, involving the presence or absence of live females and female scent. The presence of females elicited an increase in all displays except antennation; female scent increased the frequency of antennations, mandible flares and grapples, but to a lesser extent than did live females. The frequency of grapples significantly increased for males fed the high quality diet; however diet did not influence the other displays. The combined influence of diet and condition was significant for mandible flare only. Mating frequency was not influenced by diet. However, the frequency ofthe displays were positively correlated with mating frequency for high quality fed males. Escalated displays involving high costs, such as grapple and mandible flare, increased in frequency when the benefits of winning contests were high in G.bimaculatus. Escalation to grapple behaviour was less evident for males fed the lower quality diet as this imposed energy constraints on high cost displays.

Isolation and characterization of the female sex pheromone of the sugar beet cyst nematode, heterodera schachtii, and an analysis of male precopulatory behaviour /

The sugar beet cyst nematode, Heterodera schachtii, is a major agricultural pest. The disruption of the mating behaviour of this plant parasite in the field may provide a means of biological control, and a subsequent increase in crop yield. The H. schachtii female sex pheromone, which attracts homospecific males, was collected in an aqueous medium and isolated using high performance liquid chromatography. Characterization of the male-attractive material revealed that it was heat stable and water soluble. The aqueous medium conditioned by female H. schachtii was found to be biologically active and stimulated male behaviour in a concentration dependent manner. The activity of the crude pheromone was specific to males of H. schachtii and did not attract second stage juveniles. Results indicated that vanillic acid, a putative nematode pheromone, is not an active component of the H. schachtii sex pheromone. Male H. schachtii exhibited stylet thrusting, a poorly understood behaviour of the male, upon exposure to the female sex pheromone. This behaviour appeared to be associated with mate-finding and was used as a novel indicator of biological activity in bioassays. Serotonin, thought to be involved in the neural control of copulatory behaviour in nematodes, stimulated stylet thrusting. However, the relationship between stylet thrusting induced by the sex pheromone and stylet thrusting induced by serotonin is not clear. Extracellular electrical activity was recorded fi-om the anterior region of H. schachtii males during stylet thrusting, and appeared to be associated with this behaviour. The isolation of the female sex pheromone of H. schachtii may, ultimately, lead to the structural identification and synthesis of the active substance for use in a novel biological control strategy.

Male reproductive competition in the field crickets : gryllus veletis and G. pennsylvanicus

Sexual behavior in the field crickets, Gryllus veletis and G. pennsylvanicus , was studied in outdoor arenas (12 m2) at high and low levels of population density in 1983 and 1984. Crickets were weighed, individually marked, and observed from 2200 until 0800 hrs for at least 9 continuous nights. Calling was measured at 5 min intervals, and movement and matings were recorded hourly. Continuous 24 hr observations were also conducted,·and occurrences of aggressive and courtship songs were noted. The timing of males searching, calling, courting, and fighting for females should coincide with female movement and mating patterns. For most samples female movement and matings occurred at night in the 24 hr observations and were randomly distributed with time for both species in the 10 hr observations. Male movement for G. veletis high density only was enhanced at night in the 24 hr observations, however, males called more at night in both species at high and low densities. Male movement was randomly distributed with time in the 10 hr observations, and calling increased at dawn for the G. pennsylvanicus 1984 high density sample, but was randomly distributed in other samples. Most courtship and aggression songs in the 24 hr observations were too infrequent for statistical testing and generally did not coincide with matings. Assuming residual reproductive value, and costs attached to a male trait in terms of future reproductive success decline with age, males should behave in more costly ways with age; by calling and moving more with age. Consequently, mating rates should increase with age. Female behavior may not change with age. G. veletis , females moved more with age at both low density samples, however, crickets moved less with age at high density. G. pennsylvanicus females moved more with age in the 1984 low density sample, whereas crickets moved less with age in the 1983 high density sample. For both species males in the 1984 high density samples called less with age. For G. pennsylvanicus in 1983 calling and mating rates increased with age. Mating rates decreased with age for G. veletis males in the high density sample. Aging may not affect cricket behavior. As population density increases fewer calling sites become available, costs of territoriality increase, and matings resulting from non-calling behavior should increase. For both species the amount of calling and in G. veletis the distance travelled per night was not different between densities. G. pennsylvanicus males and females moved more at low density. At the same deneity levels there were no differences in calling, mating, and, movement rates in G. veletis , however, G. pennsylvanicus males moved more at high density in 1983 than 1984. There was a positive relationship between calling and mating for the G. pennsylvanicus low density sample only, and selection was acting directly to increase calling. For both species no relationships between movement and mating success was found, however, the selection gradient on movement in the G. veletis high density population was significant. The intensity of selection was not significant and was probably due to the inverse relationship between displacement and weight. Larger males should call more, mate more, and move less than smaller males. There were no correlations between calling and individual weight, and an inverse correlation between movement and size in the G. veletis high density population only. In G. pennsylvanicus , there was a positive correlation between individual weight and mating, but, some correlate of weight was under counter selection pressure and-prevented significance of the intensity of selection. In contrast, there was an inverse correlation in the G.·veletis low density B sample. Both measures of selection intensities were significant and showed that weight only was under selection pressures. An inverse correlation between calling and movement was found for G. veletis at low density only. Because males are territorial, females are predicted to move more than males, however, if movement is a mode of male-male reproductive competition then males may move more than females. G. pennsylvanicus males moved more than females in all samples, however, G. veletis males and females moved similar distances at all densities. The variation in relative mating success explained by calling scores, movement, and weight for both species and all samples were not significant In addition, for both species and all samples the intensity of selection never equalled the opportunity for selection.

The effect of gregarine parasite infections, age, and diet on calling song structure and mating behaviour in the Texas field cricket, Gryllus integer

In the past ten years, many researchers have focussed their attention on parasites regarding the role they may play in causing variations in male secondary sexual traits and subsequent effects on female choice. Male age has also been suggested to be an important factor in female choice if old age reflects superior genes. This study investigated the effects that gregarine gut parasites, age, and diet have on the calling and mating behaviour of the male Texas field cricket, Gryllus integer. Male calling songs were recorded in the laboratory using a Digital Signal Processing Network. The song parameters measured were: pulse rate, pulse width, burst duration, pulses per burst, interburst interval, and percent missing pulses. The effects of parasite load and age on the various calling song parameters was investigated in crickets that were fed two different diets varying in nutritional quality. None of the calling song parameters were affected by either parasite load or age in either diet grou p. Courtship behaviour was ob served and recorded using an Eventlog recorder on an IBM computer in the laboratory. Females mated equally with paras(tized and unparasitized males and with old and young males The total duration and proportion of time spent performing each of 9 courtship displays were recorded for males on each diet. Only one display was affected by parasite load. Highly parasitized males fed the nutritionally inferior diet juddered for a proportionately shorter time than males with low parasite loads. Also, older males performed juddering and shaking antennae proportionally longer and juddering and raising wings for longer durations than younger males. Males that successfully mated were observed for performance of 8 post-copulatory guarding behaviour displays. None of the guarding behaviours were affected by parasite load. However, one display was affected by age, with older males performing guard turning for shorter durations than younger males. Results are discuss,ed in terms of the influence of parasites and age on female choice.

The Influence of Male Diet on Life History Traits of Female Mosquitoes

In the field, mosquitoes characteristically feed on sugars soon after emergence and intermittently during their adult lives. Sugar meals are commonly derived from plant nectar and homopteran honeydew, and without them, adults can only survive for a few days on larval reserves. In addition to sugar, females of most species rely on blood for the initiation and maintenance of egg development; thus their reproductive success depends to some extent on the availability of blood hosts. Males, on the other hand, feed exclusively on sugars. Consequently, their sexual maturation and reproductive success is largely dependent upon access to sugar sources. Plant nectar and homopteran honeydew are the two main sugar sources utilized by mosquitoes in the wild. Previous laboratory studies had shown that differences between nectar sources can affect the survivorship and biting frequency of disease vectoring mosquitoes. However, little is known on how sugar composition influence the reproductive processes in male mosquitoes. Male mosquitoes transfer accessory gland proteins and other hormones to their mates along with sperm during mating. In the female, these seminal fluid constituents exert their influence on reproductive genes that control ovulation and vitellogenesis. The present study tests the hypothesis that the mates of males consuming different sugar meals will exhibit varying levels of induction of vitellogenin (a gene which regulates the expression of egg yolk precursor proteins). Real-time quantitative RT-PCR was used to investigate how each sugar meal indirectly influences vitellogenin mRNA abundance in female Anopheles stephensi following mating. Results indicate that mates of nectar-fed males exhibit 2-fold greater change in vitellogenin expression than the mates of honeydew-fed males. However, this response did not occur in non-blood fed controls. These findings suggest that the stimulatory effect of mating on vitellogenesis in blood meal-reliant (i.e. anautogenous) mosquitoes may only be synergistic in nature. The present study also sought to compare the potential fitness costs of mating incurred by females that do not necessarily require a blood meal to initiate a reproductive cycle (i.e., exhibit autogeny). Females of the facultatively autogenous mosquito, Culex molestus were allowed to mate with males sustained on either nectar or honedyew. Mean lifetime fecundity and survivorship of females under the two different mating regimes were then recorded. Additionally, one-dimensional gel electrophoresis was used to verify the transfer of male accessory gland proteins to the sperm storage organs of females during mating.While there was no significant difference in survival between the test treatments, the mates of nectar-fed males produced 11% more eggs on average than mates of honeydew-fed males. However, additional data are needed to justify the extrapolation of these findings to natural settings. These findings prompt further investigation as the differences caused by diet variation in males may be reflected across other life history traits such as mating frequency and insemination capacity.

A Phenomenologically Sensitized Analysis of Images Depicting Stressed Embodiment in an Adolescent Male with Autism Spectrum Disorder (ASD)

Each person with Autism Spectrum Disorder (ASD) comes with unique characteristics (idiosyncratic) that give clues to the world they know (Connolly, 2008). It is through their body that they (a) know the world they are experiencing, (b) make meaning, and (c) express certain behaviours. I used Laban’s Movement Analysis (LMA) to practice an attuned and appreciative approach to describing and understanding the body movement in one severe manifestation of autism in an adolescent male. LMA observes human movement across many disciplines and can be applied in many contexts providing a body honoring discourse for description (Connolly, 2008). The framework examines movement in body, space, quality, and relation. Each theme provides a detailed description of the individual’s movement, thus, giving us a richer understanding of patterns and possible triggers to self-injurious behaviours (SIB). During the summer of August 2013, I participated in Brock University’s annual Autism Camp and worked with a 15 year old male named “Aaron” who manifests with low functioning autism. The purpose of my research project was to code and analyze a series of photos taken to help gain insight into movement patterns associated with stressed embodiment and self-injury in “Aaron”. As I understood more about these embodied expressions, I uncovered valuable information on how to read patterns and discover what triggers these events, thus providing strategies on how to help people do more refined observations and make meaning of the behaviour. Laban’s movement analysis provided a sensitized discourse appropriate to the embodied expressions depicted in the photos.