Insect herbivory and ontogeny: How do growth and development influence feeding behaviour,morphology and host use?

Herbivorous insects exploit many different plants and plant parts and often adopt different feeding strategies throughout their life cycle. The conceptual framework for investigating insect–plant interactions relies heavily on explanations invoking plant chemistry, neglecting a suite of competing and interacting pressures that may also limit herbivory. In the present paper, the methods by which ontogeny, feeding strategies and morphological characters inhibit herbivory by mandibulate holometabolous insects are examined. The emphasis on mechanical disruption of plant cells in the insect digestive strategy changes the relative importance of plant ‘defences’, increasing the importance of leaf structure in inhibiting herbivory. Coupled with the implications of substantial morphological and behavioural changes in ontogeny, herbivores adopt a range of approaches to herbivory that are independent of plant chemistry alone. Many insect herbivores exhibit substantial ontogenetic character displacement in mandibular morphology. This is tightly correlated with changes in feeding strategy, with changes to the cutting edges of mandibles increasing the efficiency of feeding. The changes in feeding strategy are also characterized by changes in feeding behaviour, with many larvae feeding gregariously in early instars. Non‐nutritive hypotheses considering the importance of natural enemies, shelter‐building and thermoregulation may also be invoked to explain the ontogenetic consequences of changes to feeding behaviour. There is a need to integrate these factors into a framework considering the gamut of potential explanations of insect herbivory, recognizing that ontogenetic constraints are not only viable explanations but a logical starting point. The interrelations between ontogeny, size, morphology and behaviour highlight the complexity of insect–plant relationships. Given the many methods used by insect herbivores to overcome the challenges of consuming foliage, the need for species‐specific and stage‐ specific research investigating ontogeny and host use by insect herbivores is critical for developing general theories of insect–plant interactions.     

Why do multiple traits determine mating success? Differential use in female choice and male competition in a water boatman

Mating success is often determined by multiple traits, but why this occurs is largely unknown. Much attention has been paid to female preferences for multiple traits, but surprisingly few researchers have addressed the possibility that multiple traits are important because they serve different functions in female choice and male-male competition. Differential trait function could result from a conflict of interest between the sexes or from constraints forcing the sexes to pay attention to different traits. I show that traits determined at distinct life-history stages differ in their importance in female choice and male-male competition in a water boatman Sigara falleni. Juvenile conditions determined body and foreleg pala size and were the main determinants of mating success under female choice, whereas adult conditions determined body mass and influenced mating success when male competition was included. This differential use of condition-dependent traits under the two selection regimes appeared to arise partly from a conflict between the sexes, since the two selection forces (female choice and male competition) conflict for selection on pala size, and partly from constraints, as females appeared unable to assess adult condition.     

Modeling intra-sexual competition in a sex pheromone system: how much can female movement affect female mating success?

Mating disruption theory predicts that high concentrations of female pheromone, and/or large numbers of release sites, should confuse males orienting to "calling" females, reduce the number of successful matings, and decrease the reproductive potential of the population. In this scenario, females are regarded as stationary point sources of pheromone. Past behavioral observations, however, have shown virgin female grape root borers, Vitacea polistiformis Harris, significantly alter their behavior in mating disruption treatments. Treated females call at different heights, move less before call initiation, and move more after call initiation than control females. Pheromone gland dragging and wing fanning also increase significantly during pheromone treatments. These behavioral differences are significant only if they alter the mating success of females. Because long-term field studies are impractical, we used known behavior of male and female GRB to build a Fortran language time step model, adding the effects of female movement to past models of male pheromone plume following. Females were distributed randomly, and then assigned a conditional movement strategy. If females were within the competitive portion of another female's plume, the downwind female moved. Except in the lowest population density tested, females moving upwind and crosswind when in a competing female's pheromone plume mated significantly more often than females remaining stationary. In all population simulations, mating success was significantly reduced when females moved downwind. These field and simulation studies provide strong evidence for female movement as a previously overlooked potential mechanism for resistance to mating disruption treatments, as well as a shaping behavior in the evolution of pheromone communication systems.     

Male mating success and risk of predation in a wolf spider: a balance between sexual and natural selection?

1. Traits that benefit males through sexual selection are simultaneously expected to impair males by provoking costs through natural selection. If we consider the two male fitness components, mating success and viability, then we may expect that the increase in male mating success resulting from a larger trait size will be counterbalanced by an increase in viability costs.
2. We studied the benefits and costs of male mate searching and sexual signalling activity in the wolf spider Hygrolycosa rubrofasciata . In the field, males search females actively and court them by drumming dry leaves with their abdomen. Females have been shown to prefer males with high drumming rate. Male moving and especially drumming is energetically highly demanding and drumming results in significant mortality costs.
3. Our objective in this study was to determine whether male mate-searching activity or drumming activity affect male mating success and the risk of males being predated.
4. It was evident that both higher mate-searching activity and higher drumming activity benefited males by increasing their mating success. Higher mate-searching activity clearly impaired males by causing direct increase in predation risk. There was also a slight tendency that more actively drumming males had higher risk of predation and from all of the predated males 13.3% were caught directly after they had drummed. Furthermore, male drumming activity decreased drastically in the presence of the predator.
5. We conclude that in H. rubrofasciata both increased mate-searching activity and drumming activity benefit males through sexual selection, but at the same time natural selection provokes direct balancing costs on the same traits.     

The transvestite serpent: why do male garter snakes court (some) other males?

In large mating aggregations of red-sided garter snakes, Thamnophis sirtalis parietalis, in Manitoba, male courtship is directed not only to females, but also to other males with female-like skin lipids ('she-males'). We show that 'she-maleness' is an intrinsic property of a male rather than an artefact of lipid transfer from females, and that male-male courtship is very common in the field. She-males were distinctive in terms of appearance (they were heavier than other males and more often covered with mud), behaviour (they were inactive and rarely courted females) and performance (they were slow crawlers, ineffective courters and easily outcompeted by other males in mating trials). 'She-maleness' was not a characteristic of a particular subset of males, as envisaged in previous work; instead, it was a transitory phase that most (perhaps all) male snakes passed through soon after they first emerged from the winter den. Recently emerged males spent their first day or two relatively inactive, while restoring physiological functions (including locomotor performance and courtship ability). Experimental application of female skin lipids on to males dramatically decreased courtship levels of the recipient snakes. Thus, recently emerged males may derive two kinds of benefit from mimicking female skin lipids. First, female mimicry 'switches off' the male's own (energetically expensive) courtship at a time when that courtship would be unproductive. Second, it may disadvantage his rivals by distracting them from females, and increasing their energy expenditure. Copyright 2000 The Association for the Study of Animal Behaviour.     

Sexual selection in cane toads Rhinella marina： A male＇s body size affects his success and his tactics

Abstract Male body size can play an important role in the mating systems of anuran amphibians. We conducted labora- tory-based trials with cane toads Rhinella （Bufo） marina from an invasive population in the wet-dry tropics of northern Australia, to clarify the effects of a male＇s body size on his reproductive success and behavior （mate choice）. Males were stimulated with a synthetic hormone to induce reproductive readiness. Larger body size enhanced a male toad＇s ability to displace a smaller rival from amplexus, apparently because of physical strength： more force was required to dislodge a larger than a smaller amplectant male. A male＇s body size also affected his mate-choice criteria. Males of all body sizes were as likely to attempt amplexus with another male as with a female of the same size, and preferred larger rather than smaller sexual targets. However, this size prefe- rence was stronger in larger males and hence, amplexus was size-assortative. This pattern broke down when males were given access to already-amplectant male-female pairs： males of all body sizes readily attempted amplexus with the pair, with no size discrimination. An amplectant pair provides a larger visual stimulus, and prolonged amplexus provides a strong cue for sex iden- tification （one of the individuals involved is almost certainly a female）. Thus, a male cane toad＇s body size affects both his ability to defeat rivals in physical struggles over females, and the criteria he uses when selecting potential mates, but the impacts of that selectivity depend upon the context in which mating occurs     

Human male mating strategies: I. Courtship tactics of the “quality” and “quantity” alternatives

Human males may adopt the “quality” strategy, a long-term pair bond with considerable paternal investment, or the “quantity” reproductive strategy, short-term pair bonds with little paternal investment. We hypothesized that (1) the two strategies require different courtship tactics, which can be derived from their different goals, and (2) the behavior used by quality courters is perceived as honest while the quantity strategist is often exploitative. Act nominations of 79 subjects and the investigators generated a set of 71 dating behaviors. Sixty-four other subjects classified each behavior by the theoretically derived tactics. Fifty-four males rated the likelihood of usage of each behavior by a man interested in marriage (quality courter) and by men interested in sexual relationships (quantity courters) who typically deceive, manipulate, or coerce women. Fifty-three other males rated each behavior on ethical scales of honesty, deception, manipulation, and coercion. Both kinds of ratings indicated that (1) tactics of quality courtship involve honest advertisement through mutual assessment, resource expenditure, and a delay in sexual relations, and (2) tactics of quantity courtship involve indirect or direct threats, psychological pressure, and talking about sex. Ethical ratings and the range of behaviors likely to be used by quality and quantity courters confirmed the hypothesis about ethical perceptions of the two strategies and suggested that quantity courters are opportunistic as well as exploitative.     

May random processes explain mating success in leks?

The object of this paper is to verify whether in specific cases the variance of mating success among lekking males may be due exclusively to a random mechanism, as opposed to the adaptive mechanisms of mate choice which are usually postulated in the literature in the framework of sexual selection theory. In fact, some studies attempted to compare observed distributions of male mating success with a Poisson ‘null’ distribution based on the conjecture of random mating; the conjecture is usually rejected. In this paper we construct a plausible model (the ‘null’ hypothesis) for a strictly random non-adaptive pattern of social behaviour of lekking males and females and we perform several simulations for reasonable choices of parameter values. It should be observed that some of the simulations based on our random model lead to a distribution of male mating success which is Poisson-like. However, contrary to predictions, in several simulations a random process of mate choice lead to non-Poissonian distributions. Accordingly, the fact that, when performing a statistical test on several sets of field data, we find both cases which are in agreement with Poisson distribution, or a normal one, and cases which are not, does not allow us to reject the assumption of random male reproductive success. Thus it is legitimate to conjecture that in many cases the inter-individual variability of male mating success might indeed be determined by random processes. If this conjecture were to be confirmed by further studies, the actual significance of sexual selection in the evolution of lekking species should be reassessed, and a novel approach in the analysis of field data would be called for.     

Plasticity of boldness in rainbow trout,Oncorhynchus mykiss: do hunger and predation influence risk-taking behaviour?

Boldness, a measure of an individual's propensity for taking risks, is an important determinant of fitness but is not necessarily a fixed trait. Dependent upon an individual's state, and given certain contexts or challenges, individuals may be able to alter their inclination to be bold or shy in response. Furthermore, the degree to which individuals can modulate their behaviour has been linked with physiological responses to stress. Here we attempted to determine whether bold and shy rainbow trout, Oncorhynchus mykiss, can exhibit behavioural plasticity in response to changes in state (nutritional availability) and context (predation threat). Individual trout were initially assessed for boldness using a standard novel object paradigm; subsequently, each day for one week fish experienced either predictable, unpredictable, or no simulated predator threat in combination with a high (2% body weight) or low (0.15%) food ration, before being reassessed for boldness. Bold trout were generally more plastic, altering levels of neophobia and activity relevant to the challenge, whereas shy trout were more fixed and remained shy. Increased predation risk generally resulted in an increase in the expression of three candidate genes linked to boldness, appetite regulation and physiological stress responses - ependymin, corticotrophin releasing factor and GABA(A) - but did not produce a significant increase in plasma cortisol. The results suggest a divergence in the ability of bold and shy trout to alter their behavioural profiles in response to internal and exogenous factors, and have important implications for our understanding of the maintenance of different behavioural phenotypes in natural populations.     

Environmental correlates of phenotypic variation: do variable tidal regimes influence morphology in intertidal seaweeds?

Seaweed morphology is often shaped by the hydrodynamic environment. However, exposure to air at low tide represents an additional factor potentially affecting the morphology of intertidal species. Here, we examined the relationships between the morphology of Hormosira banksii, an important intertidal habitat‐forming seaweed in southern Australia, and environmental factors across multiple spatial scales around the island of Tasmania, Australia. Tasmania is surrounded by a diverse coastline with differences in wave exposure, tidal parameters, and temperature. We sampled Hormosira from four regions (100s km apart), three sites (10s km apart) within each region, and two zones (meters apart; eulittoral and sublittoral) at each site, and measured multiple morphological variables to test for differences in morphology at those different spatial scales. Thirteen environmental variables reflecting wave exposure, tidal conditions, and temperature for each site were generated to assess the relationship between Hormosira morphology and environmental variation. Morphology varied at all spatial scales examined. Most notably, north coast individuals had a distinct morphology, generally having smaller vesicles and shorter fronds, compared to other regions. Tidal conditions were the main environmental factors separating north coast sites from other sites and tidal regime was identified as the best predictor of morphological differences between regions. In contrast to other studies, we found little evidence that wave exposure was associated with morphological variation. Overall, our study emphasizes the role of tidal conditions, associated with emersion stress during low tide, in affecting the morphology of intertidal seaweeds.     

Does part reproductive history predict competitive interactions and male mating success in pupfish?

Two experiments were performed to determine the effect of past territoriality and mating success on subsequent dominance and ability to attract females. First, in 14 staged, pair-wise encounters, the development of nuptial coloration and agonistic behaviour was examined in two types of males: previously territorial (PT) and previously non-territorial (PNT) males. Previously territorial males developed a more intense nuptial coloration and won all contests. Second, to test the males' ability to attract females, other factors were controlled that could have affected female mate choice, such as male size, quality of breeding substrate and male-male interactions. In 40 trials, PT males were more active, developed a more intense nuptial coloration, courted females more vigorously, spawned sooner and had a higher mating success than PNT males. Thus, in pupfish, Cyprinodon pecosensis, previous reproductive history is a good indicator of both dominance and mating success. Only males in good physical condition achieve high mating success, and both intra- and inter-sexual selection are important in maintaining the close correlation between expression of nuptial coloration, a second sexual trait, and other male attributes, such as physical condition and vigour, that allow a male to secure and defend a territory against rivals and to attract and spawn with females.     

Flight morphology along a latitudinal gradient in a butterfly: do geographic clines differ between agricultural and woodland landscapes?

Bergman and converse Bergman rules, amongst others, describe latitudinal variation in size of organisms, including flying ectotherms like butterflies. However, geographic clines in morphological traits of functional significance for flight performance and thermoregulation may also exist, although they have received less attention within a biogeographical context. Variation in flight‐related morphology has often been studied relative to landscape structure. However, the extent to which landscape effects interact with latitudinal clines of phenotypic variation has rarely been tested. Here we address the effect of latitude, landscape type and the interaction effect on body size and flight‐related morphology in the speckled wood butterfly Pararge aegeria. Male adult butterflies were collected from two replicate populations in each agricultural and woodland landscape types along a 700 km cline in six latitudinal zones. Overall size, adult body mass and wing area increased with latitude in line with Bergmann's rule. Forewing length, however, decreased with latitude. As predicted from thermoregulatory needs in ectotherms, the basal wing part was darker to the north. Latitudinal trends for flight‐related morphological traits were opposite to predictions about flight endurance under cooler conditions that were observed in some non‐lepidopteran insects, i.e. wing loading increased and wing aspect ratio decreased with latitude. Opposite trends can, however, be explained by other aspects of butterfly flight performance (i.e. mate‐location behaviour). As predicted from differences in environmental buffering in woodland landscapes along the latitudinal gradient, significant landscape×latitude interaction effects indicated stronger latitudinal clines and stronger phenotypic variation for size and flight morphology in the agricultural landscape compared to the woodland landscape. In agreement with significant interaction effects, morphological differentiation increased with latitude and was higher between population pairs of agricultural landscape than between population pairs of woodland landscape. These results demonstrate that landscape, latitude and their interaction contribute to the understanding of the complex geographic variation in P. aegeria adult phenotypes across Europe.     

Why do males emerge before females? protandry as a mating strategy in male and female butterflies

Summary The reproductive strategy of butterfly males can be defined as being to maximize the number of females mated. We have earlier shown that, if the eclosion period of females is regarded as given, males should emerge before females to achieve maximal reproductive success. However, females may also be considered to have a reproductive strategy with respect to the issue when to emerge. In this paper we assume that females are selected to minimize the time spent unmated (to minimize prereproductive death), and analyze when females should optimally emerge in relation to males to achieve this end. We show that there is no conflict between the sexes with respect to the timing of eclosion when the length of the eclosion period is approximately equal for males and females. Thus, protandry should be considered a reproductive strategy of both males and females.     

Age-specific reproductive success in a long-lived bird: do older parents resist stress better?

In many vertebrates, reproductive performance increases with advancing age but mechanisms involved in such a pattern remain poorly studied. One potential mechanism may be the hormonal stress response, which shifts energy investment away from reproduction and redirects it towards survival. In birds, this stress response is achieved through a release of corticosterone and is also accompanied by a decrease in circulating prolactin, a hormone involved widely in regulating parental cares. It has been predicted that, when the value of the current reproduction is high relative to the value of future reproduction and survival, as it is expected to be in older adults, the stress response should be attenuated to ensure that reproduction is not inhibited. We tested this hypothesis by measuring the corticosterone and prolactin responses of known-age (8-36 years old) incubating snow petrels (Pagadroma nivea) to a standardized capture/handling stress protocol. We also investigated whether an attenuation of the stress responses will correlate with a lower occurrence of egg neglect, a frequently observed behaviour in snow petrels. The probability of successfully fledging a chick increased from 6 years to 12 years before stabilizing after 12 years of age. Corticosterone response to stress was unaffected by age. Prolactin response to stress, however, was influenced clearly by age: in both sexes older breeders had higher stress-induced prolactin levels than younger ones. This was due to an increasing attenuation of the prolactin response to stress with advancing age in females, and in males this was due to a probably higher intrinsic capacity of older males to secrete prolactin. Moreover, higher stress-induced prolactin levels were correlated with a lower probability of neglecting the egg. In young breeders, the combination of a robust corticosterone increase with a lower ability to maintain prolactin secretion during acute stress is probably one of the functional causes of their lower incubation commitment. We suggest that the ability to maintain a threshold level of prolactin during a stressful situation may be an important physiological mechanism involved in the improvement of reproductive performance with advancing age in long-lived birds.     

Mechanosensitive channels: what can they do and how do they do it?

While mechanobiological processes employ diverse mechanisms, at their heart are force-induced perturbations in the structure and dynamics of molecules capable of triggering subsequent events. Among the best characterized force-sensing systems are bacterial mechanosensitive channels. These channels reflect an intimate coupling of protein conformation with the mechanics of the surrounding membrane; the membrane serves as an adaptable sensor that responds to an input of applied force and converts it into an output signal, interpreted for the cell by mechanosensitive channels. The cell can exploit this information in a number of ways: ensuring cellular viability in the presence of osmotic stress and perhaps also serving as a signal transducer for membrane tension or other functions. This review focuses on the bacterial mechanosensitive channels of large (MscL) and small (MscS) conductance and their eukaryotic homologs, with an emphasis on the outstanding issues surrounding the function and mechanism of this fascinating class of molecules.