Analysis of the importance of genotypic variation, metabolic rate, morphology, sex and development time on immune function in the cricket, Gryllus firmus

Immune defence is hypothesized to be a trait that bears significant fitness costs as well as benefits in that mounting a defence depreciates the value of other life‐history traits. Thus the cost of mounting an immune response could affect the evolution of both the immune system and correlated life history traits. In this study we examined, by means of a diallel cross of four inbred lines, the genetic basis of two measures of immune function, metabolic rate and several traits in the sand cricket, Gryllus firmus. We specifically addressed the following questions: (1) is immune function determined primarily by genetic constitution or correlations with phenotypic traits that could reduce the effectiveness of the immune response; (2) do the two measures of immune function covary; (3) What are the contributions of additive, nonadditive and maternal effects to the immune function? As estimates of immune function, we used lytic activity and encapsulation rate. We found that inbred crickets were smaller than individuals from the crossed lines and took longer to develop. However, inbred lines did not differ from the crossed lines in immune function nor metabolic rates, suggesting that increased homozygosity has little or no effect on these traits in G. firmus. We found that both immune parameters showed significant genetic variation but no consistent relationships with the other phenotypic traits (metabolic rate, head width, body mass, development time and activity). There was significant additive genetic variation only in encapsulation rate, but, with the exception of the activity measure, significant nonadditive and reciprocal variances were found in all traits. Metabolic rate of crickets was heritable, but there was neither phenotypic nor genetic association between metabolic rate and the two parameters of immune function. Further, there was no correlation between these two measures. Females showed a higher encapsulation response than males, but there was no sex differences in lytic activity. Our study indicates that genetic variation in immune parameters can be a very significant contributor to phenotypic variation in immune function.     

The effects of sex hormones on immune function: a meta‐analysis

The effects of sex hormones on immune function have received much attention, especially following the proposal of the immunocompetence handicap hypothesis. Many studies, both experimental and correlational, have been conducted to test the relationship between immune function and the sex hormones testosterone in males and oestrogen in females. However, the results are mixed. We conducted four cross‐species meta‐analyses to investigate the relationship between sex hormones and immune function: (i) the effect of testosterone manipulation on immune function in males, (ii) the correlation between circulating testosterone level and immune function in males, (iii) the effect of oestrogen manipulation on immune function in females, and (iv) the correlation between circulating oestrogen level and immune function in females. The results from the experimental studies showed that testosterone had a medium‐sized immunosuppressive effect on immune function. The effect of oestrogen, on the other hand, depended on the immune measure used. Oestrogen suppressed cell‐mediated immune function while reducing parasite loads. The overall correlation (meta‐analytic relationship) between circulating sex hormone level and immune function was not statistically significant for either testosterone or oestrogen despite the power of meta‐analysis. These results suggest that correlational studies have limited value for testing the effects of sex hormones on immune function. We found little evidence of publication bias in the four data sets using indirect tests. There was a weak and positive relationship between year of publication and effect size for experimental studies of testosterone that became non‐significant after we controlled for castration and immune measure, suggesting that the temporal trend was due to changes in these moderators over time. Graphical analyses suggest that the temporal trend was due to an increased use of cytokine measures across time. We found substantial heterogeneity in effect sizes, except in correlational studies of testosterone, even after we accounted for the relevant random and fixed factors. In conclusion, our results provide good evidence that testosterone suppresses immune function and that the effect of oestrogen varies depending on the immune measure used.     

Forceps size and immune function in the earwig Forficula auricularia L.

Females of many species select their mates on the basis of the size or intensity of sexual ornaments, and it has been suggested that these provide reliable signals of a male's ability to resist parasites and pathogens. European earwigs, Forficula auricularia , are sexually dimorphic in forceps shape and length. Male forceps are used as weapons in male contests for access to females, but recent findings suggest that females also choose males on the basis of their forceps length. In the present study, we tested the hypotheses that in the European earwig, F. auricularia , the size of forceps is correlated with immune function and that immune function differs between the sexes. We found that encapsulation rate was not correlated with the length of forceps in either sex, but was negatively correlated with body size. By contrast, lytic activity of the haemolymph increased with overall body size in both sexes but, in females, lytic activity increased with relative forceps length whereas, in males, it decreased with relative forceps length. After accounting for effects of body size, there was no remaining significant correlation in females but the negative correlation in males remained. Furthermore, we found that males had higher encapsulation rate and higher lytic activity than females, suggesting that males have stronger immune defence. The results of the study indicate that the size of forceps in male earwigs does not reliably reflect male immune defence.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 509–516.     

Quantitative genetics of immune function and body size in the house cricket,Acheta domesticus

Female house crickets are attracted to male calling song containing a relatively high number of syllables per ‘chirp’, which tends to be produced by large males. In a previous study, we showed that this song characteristic is also positively and independently correlated with haemocyte load, an important determinant of the ability to produce an encapsulation response in insects. Females will therefore tend to select males with high encapsulation ability (and large body size) as mates. The present study demonstrates that variation in haemocyte load and body size, together with a second parameter of immune function (the ability to encapsulate a synthetic substrate), is heritable in the same population. Moreover, all three traits are shown to be positively genetically correlated. In favouring males that produce calling song with the preferred characteristics, females should therefore also tend to produce larger offspring with a greater ability to produce an encapsulation response.     

Queen recruitment and split sex ratios in polygynous colonies of the ant Formica exsecta

Sex ratios in social insects have become a general model for tests of inclusive fitness theory, sex ratio theory and parent–offspring conflict. In populations of Formica exsecta with multiple queens per colony , sex ratios vary greatly among colonies and the dry-weight sex ratio is extremely male-biased, with 89% of the colonies producing males but no gynes (reproductive females). Here we test the queen-replenishment hypothesis, which was proposed to explain sex ratio specialization in this and other highly polygynous ants (i.e. those with many queens per nest). This hypothesis proposes that, in such ants, colonies produce gynes to recruit them back into the colony when the number of resident queens falls below a given threshold limiting colony productivity or survival. We tested predictions of the queen-replenishment hypothesis by following F. exsecta colonies across two breeding seasons and relating the change in effective queen number with changes in sex ratio, colony size and brood production. As predicted by the queen-replenishment hypothesis, we found that colonies that specialized in producing females increased their effective queen number and were significantly more likely to specialize in male production the following year. The switch to male production also coincided with a drop in productivity per queen as predicted. However, adoption of new queens did not result in a significant increase in total colony productivity the following year. We suggest that this is because queen production comes at the expense of worker production and thus queen production leads to resource limitation the following year, buffering the effect of greater queen number on total productivity.     

Individual variation in nest size and nest site features of the Bornean orangutans (Pongo pygmaeus)

Nest construction is a daily habit of independent orangutans for sleeping or resting. Data on their nests have been used in various ecological studies (e.g., density estimation, ranging behavior, evolution of material culture) because they are the most observable field signs. We investigated nest size and nest site features of Bornean orangutans in the wild during 10 months' fieldwork at three sites in East Kalimantan, Indonesia: Kutai National Park, Birawa, and Meratus. To examine individual variation, we followed 31 individual orangutans and recorded the 92 nests they made for nest size (diameter) and nest site features (height of nest above ground, tree species used for the nest site, the diameter and height of the tree, whether the nest was new or reused, and nest location within the tree). Analyses taking age–sex classes of the focal individuals into consideration showed significant age–sex differences in nest size and location, but not in nest height or nest tree features (diameter, height of tree, and height of lowest branch). Mature orangutans (adult females, unflanged and flanged males) made larger nests than immatures (juveniles and adolescents). Flanged male orangutans with larger nests used stable locations for nesting sites and reused old nests more frequently than immatures. The overall proportion of nests in open (exposed) locations was higher than in closed (sheltered) locations. Flanged males and immatures frequently made open nests, whereas adult females with an infant preferred closed locations. The good correspondence between nest size and age–sex classes indicates that nest size variation may reflect body size and therefore age–sex variation in the population. Am. J. Primatol. 71:393–399, 2009. © 2009 Wiley‐Liss, Inc.     

Interactive effects of social environment,age and sex on immune responses in Drosophila melanogaster

Social environments have been shown to have multiple effects on individual immune responses. For example, increased social contact might signal greater infection risk and prompt a prophylactic upregulation of immunity. This differential investment of resources may in part explain why social environments affect ageing and lifespan. Our previous work using Drosophila melanogaster showed that single‐sex social contact reduced lifespan for both sexes. Here, we assess how social interactions (isolation or contact) affect susceptibility to infection, phagocytotic activity and expression of a subset of immune‐ and stress‐related genes in young and old flies of both sexes. Social contact had a neutral, or even improved, effect on post‐infection lifespan in older flies and reduced the expression of stress response genes in females; however, it reduced phagocytotic activity. Overall, the effects of social environment were complex and largely subtle and do not indicate a consistent effect. Together, these findings indicate that social contact in D. melanogaster does not have a predictable impact on immune responses and does not simply trade‐off immune investment with lifespan.     

Male mating success in the Azorean rock-pool blenny: the effects of body size, male behaviour and nest characteristics

Larger male Azorean rock-pool blennies Parablennius sanguinolentus parvicornis received more female visits, were courted more often by females and received more spawnings. Larger males also received a higher number of male intrusions, attacked more conspecifics and defended larger territories. Larger males showed more nest cleaning behaviour and a marginally non-significant trend for higher egg fanning rate. Male courtship, male attack rate against conspecifics and parental behaviour were all correlated with the frequency of female spawnings received by each male even when controlling for male size, suggesting that these behaviour patterns influence male mating success. On the other hand, a positive partial correlation was found between female courtship and the frequency of female spawnings, controlling for the number of female visits, which suggests a role for female mate choice on male mating success. Finally, males nesting in chambers in the bottom of pools received more spawnings than males nesting either in crevices or under boulders. However, nest opening area was associated significantly negatively with male mating success, when controlling for male size. Thus, the present data suggest strongly that male characteristics overrule nest characteristics in determining male mating success in the Azorean rock-pool blenny.     

Testing the predictions of sex allocation hypotheses in dimorphic,cooperatively breeding riflemen

Evolutionary theory predicts that parents should invest equally in the two sexes. If one sex is more costly, a production bias is predicted in favour of the other. Two well‐studied causes of differential costs are size dimorphism, in which the larger sex should be more costly, and sex‐biased helping in cooperative breeders, in which the more helpful sex should be less costly because future helping “repays” some of its parents’ investment. We studied a bird species in which both processes should favor production of males. Female riflemen Acanthisitta chloris are larger than males, and we documented greater provisioning effort in more female‐biased broods indicating they are likely costlier to raise. Riflemen are also cooperative breeders, and males provide more help than females. Contrary to expectations, we observed no male bias in brood sex ratios, which did not differ significantly from parity. We tested whether the lack of a population‐wide pattern was a result of facultative sex allocation by individual females, but this hypothesis was not supported either. Our results show an absence of adaptive patterns despite a clear directional hypothesis derived from theory. This appears to be associated with a suboptimal female‐biased investment ratio. We conclude that predictions of adaptive sex allocation may falter because of mechanistic constraint, unrecognized costs and benefits, or weak selection.     

Inter- and intra-sexual variation in immune defence in the cabbage white butterfly, Pieris rapae L. (Lepidoptera: Pieridae)

Abstract.  1. Immune defence imposes fitness costs as well as benefits, so organisms are expected to optimise, not maximise, their immune responses. This should result in variation in immune responses under varying environmental conditions.
2. Males and females are expected to exhibit different immune responses because life-history differences between the sexes affect optimal immune response. These life-history differences should usually result in a greater female, than male, immune defence. In this study, intra- and inter-sexual variation in one component of immune defence, the encapsulation response, in cabbage white butterflies ( Pieris rapae L.), was examined.
3. Encapsulation decreased with increasing age and in response to reduced diet quality.
4. Contrary to predictions, males generally had greater immune responses than females, although this pattern varied with age.
5. These patterns of inter- and intra-sexual variation in encapsulation may result from resource-based trade-offs with components of reproductive effort and/or because of sexual dimorphism in melanin-based wing patterns.     

Variation in body mass index among Polish adults: effects of sex, age, birth cohort, and social class

Remains of 15 hominids were recovered within a Mousterian archaeological context in the cave of Qafzeh, Israel. Dated to ca. 95 kyr BP, this skeletal material has been crucial for understanding biological, chronological, and cultural aspects of anatomically modern ancient Homo sapiens. The high proportion of children (N = 8) in Qafzeh Cave is unique among Middle Palaeolithic sites and encourages the search for skeletal evidence of disease and trauma. We report on the case of one child, Qafzeh 12, ca. 3 years old (according to modern human reference standards), who manifests some outstanding skeletal abnormalities that indicate hydrocephalus.     

Rapid evolution of larval life history,adult immune function and flight muscles in a poleward‐moving damselfly

Although a growing number of studies have documented the evolution of adult dispersal‐related traits at the range edge of poleward‐expanding species, we know little about evolutionary changes in immune function or traits expressed by nondispersing larvae. We investigated differentiation in larval (growth and development) and adult traits (immune function and flight‐related traits) between replicated core and edge populations of the poleward‐moving damselfly Coenagrion scitulum. These traits were measured on individuals reared in a common garden experiment at two different food levels, as allocation trade‐offs may be easier to detect under energy shortage. Edge individuals had a faster larval life history (growth and development rates), a higher adult immune function and a nearly significant higher relative flight muscle mass. Most of the differentiation between core and edge populations remained and edge populations had a higher relative flight muscle mass when corrected for latitude‐specific thermal regimes, and hence could likely be attributed to the range expansion process per se. We here for the first time document a higher immune function in individuals at the expansion front of a poleward‐expanding species and documented the rarely investigated evolution of faster life histories during range expansion. The rapid multivariate evolution in these ecological relevant traits between edge and core populations is expected to translate into changed ecological interactions and therefore has the potential to generate novel eco‐evolutionary dynamics at the expansion front.     

蛾类昆虫雄性信息素及其功能

昆虫性信息素是两性通讯系统的基础,其中雄性信息素的研究相对较少。本文综述了蛾类昆虫雄性信息素的研究进展。迄今已鉴定出40余种蛾类昆虫的雄性信息素,其行为学功能主要有对雌性的引诱和激欲、对同种雄性的抑制及种间隔离等。     

Age, sex, and dominance-related mushroom body plasticity in the paperwasp Mischocyttarus mastigophorus

Social Hymenoptera are important models for analyzing functional brain plasticity. These insects provide the opportunity to learn how individuals' social roles are related to flexible investment in different brain regions. We assessed how age, sex, and individual behavior influence brain development in a primitively eusocial paper wasp, Mischocyttarus mastigophorus. Previous research in other species has demonstrated experience-dependent changes in central and primary sensory centers in the brain. The mushroom body (MB) calyx is a central processing region involved in sensory integration, learning and memory and may be particularly relevant to social behavior. We extend earlier cross-sectional studies of female brain/behavior associations by measuring sex- and age-related differences in MB calyx volume, and by quantifying optic lobe and antennal lobe development. Age did predict MB development: calyx neuropils increased in volume with age. We show that MB development differs between the sexes. Males, who frequently depart to seek mating opportunities, have larger MB calyx collars (which receive optic input) than females. In contrast, females have augmented predominantly antenna-innervated MB calyx lips, which may be useful for nestmate recognition and interactions on the nest. Sex differences in MB development increased with age. After accounting for age and sex effects, social aggression was positively correlated with MB calyx volume for both sexes. We found little evidence for relationships among sex, age, or behavior and the volumes of peripheral sensory processing structures. We discuss the implications of gender- and age-related effects on brain volume in relation to male and female life history and reproductive success.     

Individual variation and the endocrine regulation of behaviour and physiology in birds: a cellular/molecular perspective

Investigations of the cellular and molecular mechanisms of physiology and behaviour have generally avoided attempts to explain individual differences. The goal has rather been to discover general processes. However, understanding the causes of individual variation in many phenomena of interest to avian eco-physiologists will require a consideration of such mechanisms. For example, in birds, changes in plasma concentrations of steroid hormones are important in the activation of social behaviours related to reproduction and aggression. Attempts to explain individual variation in these behaviours as a function of variation in plasma hormone concentrations have generally failed. Cellular variables related to the effectiveness of steroid hormone have been useful in some cases. Steroid hormone target sensitivity can be affected by variables such as metabolizing enzyme activity, hormone receptor expression as well as receptor cofactor expression. At present, no general theory has emerged that might provide a clear guidance when trying to explain individual variability in birds or in any other group of vertebrates. One strategy is to learn from studies of large units of intraspecific variation such as population or sex differences to provide ideas about variables that might be important in explaining individual variation. This approach along with the use of newly developed molecular genetic tools represents a promising avenue for avian eco-physiologists to pursue.     

Host instar, body size and fitness in the koinobiotic parasitoid Aphidius nigripes

In aphidiine parasitoids, resources for growth and adult body size increase with host instar used by ovipositing females, but the fitness consequences of body size on fitness are poorly documented. We compared the fitness of male and female A. nigripesadults that varied in size as a consequence of developing in different instars of their host Macrosiphum euphorbiae. When reproductive fitness was measured without considering time, female wasps from small and large hosts performed similarly, contributing 125–175 foundresses plus 100–180 sons to the next generation. However, when expressed as the innate capacity for increase (r _m), female fitness correlated with host-induced variation of wasp size, indicating that micropopulations initiated by large wasps would increase faster. In a wind-tunnel, a sex pheromone plume from large female wasps induced more males to fly upwind when released at a distance of 50 cm downwind than small females, indicating that large females were sexually more attractive. With respect to male body size effects on fitness, large individuals performed similar to small ones, whether fitness was measured by lifetime mating frequency, fertile inseminations, or proportion of daughters among progeny born from their mates. When young naive males of unequal size were directly competing for mating with a virgin female, small and large males had equal mating success, and large individuals were no more successful than small ones at displacing a competitor already positioned on a receptive female. In a wind-tunnel test where males were scored on their ability to reach a female pheromone source, small and large males were equally affected by wind speed but reached the source located 50 cm downwind in equal proportions, suggesting similar capacity for finding mates by flying upwind. Our results indicate that despite host resources not being fixed at the time of attack for the koinobiont A. nigripes, fitness consequences of resource limitation by the mother may be perceived to be greater for daughters than sons, which would explain male-biased sex ratio in early-instar hosts.     

性别、季节和体型大小对吐鲁番沙虎巢域的影响

2008年6月份至2009年5月份对吐鲁番沙虎的巢域进行调查:2008年6月份至2008年8月份为繁殖季节(RS),2008年9月份至2009年5月份(冬眠期除外)为非繁殖季节(NRS)。利用截趾标志重捕法研究吐鲁番沙虎的巢域,共标记283只吐鲁番沙虎,累计繁殖季节24只,非繁殖季节43只重捕超过3次(其中13只个体在繁殖季节和非繁殖季节均够3次以上捕捉次数,为重复个体),可以用于计算个体巢域面积数据。利用软件MapGis计算最小凸多边形法(MCP)巢域面积,并分析性别、体型大小、季节等因素对巢域的影响。结果表明:吐鲁番沙虎非繁殖季节雄性、雌性与幼体各组间的巢域面积差异均显著,繁殖季节巢域面积差异不显著;雌雄个体不同季节或全年合并比较巢域面积差异性均不显著;非繁殖季节面积与吻肛长(SVL)显著相关、全年成体组的巢域面积与吻肛长显著相关;成体巢域面积季节差异显著(U=41,P=0.046),幼体则没有季节差异(U=159,P=0.537)。因而,吐鲁番沙虎的巢域大小受性别因素影响不大,体型大小对巢域面积有显著影响,由于繁殖、食物资源等的季节变化是影响吐鲁番沙虎巢域最重要的因素。