A Single-Neuron Chemosensory Switch Determines the Valence of a Sexually Dimorphic Sensory Behavior

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Interactions between host sex and age of exposure modify the virulence–transmission trade‐off

The patterns of immunity conferred by host sex or age represent two sources of host heterogeneity that can potentially shape the evolutionary trajectory of disease. With each host sex or age encountered, a pathogen's optimal exploitative strategy may change, leading to considerable variation in expression of pathogen transmission and virulence. To date, these host characteristics have been studied in the context of host fitness alone, overlooking the effects of host sex and age on the fundamental virulence–transmission trade‐off faced by pathogens. Here, we explicitly address the interaction of these characteristics and find that host sex and age at exposure to a pathogen affect age‐specific patterns of mortality and the balance between pathogen transmission and virulence. When infecting age‐structured male and female Daphnia magna with different genotypes of Pasteuria ramosa, we found that infection increased mortality rates across all age classes for females, whereas mortality only increased in the earliest age class for males. Female hosts allowed a variety of trade‐offs between transmission and virulence to arise with each age and pathogen genotype. In contrast, this variation was dampened in males, with pathogens exhibiting declines in both virulence and transmission with increasing host age. Our results suggest that differences in exploitation potential of males and females to a pathogen can interact with host age to allow different virulence strategies to coexist, and illustrate the potential for these widespread sources of host heterogeneity to direct the evolution of disease in natural populations.     

Function‐related Drivers of Skull Morphometric Variation and Sexual Size Dimorphism in a Subterranean Rodent,Plateau Zokor (Eospalax baileyi)

Sexual dimorphism is prevalent in most living organisms. The difference in size between sexes of a given species is generally known as sexual size dimorphism (SSD). The magnitude of the SSD is determined by Rensch's rule where size dimorphism increases with increasing body size when the male is the larger sex and decreases with increasing average body size when the female is the larger sex. The unique underground environment that zokors (Eospalax baileyi) live under in the severe habitat of the Qinghai‐Tibetan Plateau (QTP) could create SSD selection pressures that may or may not be supported by Rensch's rule, making this scientific question worthy of investigation. In this study, we investigated the individual variation between sexes in body size and SSD of plateau zokors using measurements of 19 morphological traits. We also investigated the evolutionary mechanisms underlying SSD in plateau zokors. Moreover, we applied Rensch's rule to all extant zokor species. Our results showed male‐biased SSD in plateau zokors: The body‐ and head‐related measurements were greater in males than in females. Linear regression analysis between body length, body weight, and carcass weight showed significant relationships with some traits such as skull length, lower incisor length, and tympanic bulla width, which might support our prediction that males have faster growth rates than females. Further, the SSD pattern corroborated the assumption of Rensch's rule in plateau zokors but not in the other zokor species. Our findings suggest that the natural underground habitat and behavioral differences between sexes can generate selection pressures on male traits and contribute to the evolution of SSD in plateau zokors.     

SEXUAL SELECTION AND THE EVOLUTION OF SEXUAL SIZE DIMORPHISM IN THE WATER STRIDER,AQUARIUS REMIGIS

Sexual size dimorphism (SSD) is often attributed to sexual selection, particularly when males are the larger sex. However, sexual selection favoring large males is common even in taxa where females are the larger sex, and is therefore not a sufficient explanation of patterns of SSD. As part of a more extensive study of the evolution of SSD in water striders (Heteroptera, Gerridae), we examine patterns of sexual selection and SSD in 12 populations of Aquarius remigis. We calculate univariate and multivariate selection gradients from samples of mating and single males, for two sexually dimorphic traits (total length and profemoral width) and two sexually monomorphic traits (mesofemoral length and wing form). The multivariate analyses reveal strong selection favoring larger males, in spite of the female-biased SSD for this trait, and weaker selection favoring aptery and reduced mesofemoral length. Selection is weakest on the most dimorphic trait, profemoral width, and is stabilizing rather than directional. The pattern of sexual selection on morphological traits is therefore not concordant with the pattern of SSD. The univariate selection gradients reveal little net selection (direct + indirect) on any of the traits, and suggest that evolution away from the plesiomorphic pattern of SSD is constrained by antagonistic patterns of selection acting on this suite of positively correlated morphological traits. We hypothesize that SSD in A. remigis is not in equilibrium, a hypothesis that is consistent with both theoretical models of the evolution of SSD and our previous studies of allometry for SSD. A negative interpopulation correlation between the intensity of sexual selection and the operational sex ratio supports the hypothesis that, as in several other water strider species, sexual selection in A. remigis occurs through generalized female reluctance rather than active female choice. The implications of this for patterns of sexual selection are discussed.     

Dimorphic ergatoid males and their reproductive behavior in the ponerine antHypoponera bondroiti

Summary We studied the reproductive behavior of the ponerine antHypoponera bondroiti from Okinawa, Japan. This species has dimorphic wingless ergatoid males (major and minor), dimorphic reproductive females (alate queens and wingless reproductive intercastes), and workers. Workers have neither ovarioles nor spermatheca. Major ergatoid males are the largest colony members. Two major males fought one another in the nest until one disappeared, leaving the other to occupy the nest chambers where queens emerge and mate. Minor ergatoid males also fought one another, although they seemed to be less pugnacious, resulting in occasional cohabitation of multiple minor males in the same nest chamber. Major males never attacked minor ones, allowing them to coexist in the same nest chamber. Minor males seemed to mimic females. Both major and minor males mated with both alate queens and intercastes within the nest. After mating, some alate queens shed their wings and remained in the nest, while the others left the nest for dispersal in the laboratory. Intercastes remained in the nest.     

Statistical discrimination of sex in Melanoides tuberculata (Gastropoda: Thiaridae)

Systematists may rely on morphometric differences among samples of specimens for the recognition of living and fossil species, even though morphometric differentiation may be caused by non-genetic factors, such as ecophenotypy, differential growth rates and taphonomic mixing. When genetic differences between sexes or among closely related species are expressed as differences in the morphology of the individual or population, potentially valuable information becomes available to the systematist for a variety of genetic and ecological investigations. We have studied the morphology of the freshwater snail Melanoides tuberculata (Muller, 1774) in Israel, where males occur in what would otherwise be normally parthenogenetic (all female) populations. In modern M. tuberculata, sex may be determined by observation of gonadal tissue; in fossil specimens, any classification according to sex must be accomplished using only preservable features of the mineralized shell. Previous research confirmed that in large samples, mean shell shape of male and female snails differed significantly, but the degree of difference was too small to identify the sex of any individual specimen. We apply a three stage process that results with a high degree of accuracy in the discrimination of individual M. tuberculata specimens by sex on the basis of continuous morphological characters: (1) measurement of many aspects of shell morphology of individuals of known sex, and stepwise discrimination to discover which of the variables, if any, contribute to the morphometric differentiation of males from females (one time only, for the species); (2) use of these selected variables in a clustering procedure to make a preliminary assignment of each specimen to sex; (3) use of cluster assignments in a discrimination procedure to optimally predict sex. For species that exhibit morphometric differences between two groups, and for which continuous morphometric variation precludes the a priori recognition of discrete clusters, this sequential procedure may be of broad applicability. These objective methods may be applied to the discrimination within any set of specimens for which the hypothesis of two, and only two, constituent groups may be entertained.     

Geographic variation in seed dormancy among populations of Echinochloa crus-galli

In 1991–1993, we investigated the incidence of seed dormancy in 25 local populations of barnyard grass, Echinochloa crus-galli (L.) P.Beauv., in the western Czech Republic. The percentage of germination after 4 months afterripening of dry seeds at 25°C varied between 0.0 and 83.6%. Although there were significant annual differences in the percentage of germination at some localities, typical proportions of dormant seeds persisted over 3 years at field sites where the seed bank was not disturbed. One-way ANOVA (using data from 14 cultivated or abandoned fields) revealed that 73.0% of variance in seed dormancy incidence could be attributed to the effect of locality (P<0.001). Incidence of dormancy was not correlated with mother plant stature (dry above-ground biomass, number of tillers, maximal stem height) nor seed mass. There was a significant correlation (r ²=0.403, P<0.005) between dormancy incidence at natural localities in 1991 and in F₁ offspring sown at experimental grounds at Praha-Ruzyn in 1992. The results indicate that heredity is important in maintaining local variation in seed dormancy, probably favoured by the self-pollinating reproduction of barnyard grass.     

Body size and fecundity in the waterstrider Aquarius remigis: a test of Darwin's fecundity advantage hypothesis

The general female bias in body size of animals is usually attributed to fecundity selection. While many studies have demonstrated a positive relationship between body size and fecundity, the most common interpretation of fecundity selection is that larger females have larger abdomens and can hold more eggs, yet the relationship between abdomen size and fecundity has rarely been examined. For the waterstrider, Aquarius remigis, we find a significant relationship between body size and fecundity and demonstrate that the target of fecundity selection is abdomen size. Thus, larger females have higher fecundities because they have larger abdomens and not because of their total size per se. The rate at which fecundity increases with increasing abdomen size exceeds that which would be expected due to a simple volume constraint and suggests that other factors, such as increased ability to obtain resources, may contribute to the increase in fecundity with body size. Selection intensities estimated from our data indicate that fecundity selection could be a significant selective force on both total and abdomen lengths. Previous studies have found that abdomen size increased faster than body size and thus, larger females had relatively larger abdomens. The relationship of abdomen length and thorax length in A. remigis is hypoallometric and indicates that larger females have relatively smaller abdomens. We hypothesize that this may reflect conservation of abdomen size in females developing under poor conditions. Finally, while egg size is not directly related to body size, we find a trade-off between egg size and number when female abdomen length is held constant, suggesting that selection on egg size may influence abdomen length only indirectly through its effects on fecundity.     

In-vitro evidence for feed-back regulation of β-ketoacyl-acyl carrier protein synthase III in medium-chain fatty acid biosynthesis

The cerulenin-insensitive -ketoacyl-acyl carrier protein (ACP) synthase III (KAS III, EC 2.3.1.41) catalyzes the first condensing step of the fatty-acid synthase (FAS) reaction in plants and bacteria, using directly acetyl-CoA as substrate for condensation with malonyl-ACP. In order to identify a possible site for regulation of the biosynthesis of medium-chain fatty acids, the influence of acyl-ACPs of different chain-lengths (C₄,C₆,C₈ and C₁₀) on the activity of KAS III was investigated in vitro using an FAS preparation from seeds of Cuphea lanceolata Ait. (a crop accumulating up to 90% decanoic acid into triacylglycerols) that had been treated with 100 M cerulenin. All acyl-ACPs investigated led to a decrease in the activity of KAS III towards acetyl-CoA, an effect apparently related to the length of the acyl chain. Analysis of the reaction products of the assay revealed that short-chain acyl-ACPs elongated to a very small extent simultaneously with acetyl-CoA. This extent of elongation did not correlate with the decrease in KAS III-activity levels. These data excluded the possibility of competition between acetyl-CoA and acyl-ACPs, but indicated that acyl-ACPs inhibited the enzyme. Decanoyl-ACP caused the highest decrease in enzyme activity (IC₅₀ = 0.45 M), thus being a potent inhibitor of KAS III. Michaelis-Menten kinetics revealed that the inhibition of KAS III by decanoyl-ACP was non-competitive in relation to malonyl-ACP and uncompetitive in relation to acetyl-CoA. Moreover, our data indicate that KAS III has a strict specificity for the elongation of acetyl-CoA. An inhibition of KAS III by acyl-ACPs was observed in experiments using FAS preparations from rape seeds and spinach leaves, but the inhibition of KAS III from C. lanceolata seeds by decanoyl-ACP was approximately 1.5-fold higher. The data provide evidence that acyl-ACPs are involved in the modulation of plant fatty-acid biosynthesis by a feed-back mechanism.Abbreviations ACP acyl carrier protein - DTT dithiothreitol - TCA trichloroacetic acid - ecACP acyl carrier protein from Escherichia coli - FAS fatty-acid synthase - IC₅₀ concentration causing 50% inhibition - KAS -ketoacyl-ACP synthase - NEM N-ethylmaleimide In honour of Professor Hartmut K. Lichtenthaler's sixtieth birthdayThis work was supported by a grant from the German Ministry of Research and Technology (BMFT) and in part by the Fonds der Chemischen Industrie and the Ministry of Science and Research of the State Northrhine-Westfalia. The authors wish to thank Prof. G. Röbbelen (University of Göttingen, Göttingen, Germany) for kindly providing the plant material. This paper is part of the doctoral thesis of Fritzi Maike Brück.