The effect of ethnicity on the vascular responses to cold exposure of the extremities

This is an accepted manuscript of an article published by Springer in European Journal of Applied Physiology on 01/08/2014, available online: https://doi.org/10.1007/s00421-014-2962-2 The accepted version of the publication may differ from the final published version.© 2014, Springer-Verlag Berlin Heidelberg. Purpose: Cold injuries are more prevalent in individuals of African descent (AFD). Therefore, we investigated the effect of extremity cooling on skin blood flow (SkBF) and temperature (Tsk) between ethnic groups.Methods: Thirty males [10 Caucasian (CAU), 10 Asian (ASN), 10 AFD] undertook three tests in 30 °C air whilst digit Tsk and SkBF were measured: (i) vasomotor threshold (VT) test—arm immersed in 35 °C water progressively cooled to 10 °C and rewarmed to 35 °C to identify vasoconstriction and vasodilatation; (ii) cold-induced vasodilatation (CIVD) test—hand immersed in 8 °C water for 30 min followed by spontaneous warming; (iii) cold sensitivity (CS) test—foot immersed in 15 °C water for 2 min followed by spontaneous warming. Cold sensory thresholds of the forearm and finger were also assessed.Results: In the VT test, vasoconstriction and vasodilatation occurred at a warmer finger Tsk in AFD during cooling [21.2 (4.4) vs. 17.0 (3.1) °C, P = 0.034] and warming [22.0 (7.9) vs. 12.1 (4.1) °C, P = 0.002] compared with CAU. In the CIVD test, average SkBF during immersion was greater in CAU [42 (24) %] than ASN [25 (8) %, P = 0.036] and AFD [24 (13) %, P = 0.023]. Following immersion, SkBF was higher and rewarming faster in CAU [3.2 (0.4) °C min−1] compared with AFD [2.5 (0.7) °C min−1, P = 0.037], but neither group differed from ASN [3.0 (0.6) °C min−1]. Responses to the CS test and cold sensory thresholds were similar between groups.Conclusion: AFD experienced a more intense protracted finger vasoconstriction than CAU during hand immersion, whilst ASN experienced an intermediate response. This greater sensitivity to cold may explain why AFD are more susceptible to cold injuries.Published versio

Role of cyclooxygenase in the vascular response to locally delivered acetylcholine in Caucasian and African descent individuals

This is an accepted manuscript of an article published by Elsevier in Microvascular Research on 17/01/2017, available online: https://doi.org/10.1016/j.mvr.2017.01.005 The accepted version of the publication may differ from the final published version.© 2017 Elsevier Inc. Introduction Individuals of African descent (AFD) are more susceptible to non-freezing cold injury (NFCI) compared with Caucasian individuals (CAU). Vasodilatation to acetylcholine (ACh) is lower in AFD compared with CAU in the non-glabrous foot and finger skin sites; the reason for this is unknown. Prostanoids are responsible, in part, for the vasodilator response to ACh, however it is not known whether the contribution differs between ethnicities. Methods 12 CAU and 12 AFD males received iontophoresis of ACh (1 w/v%) on non-glabrous foot and finger skin sites following placebo and then aspirin (600 mg, single blinded). Aspirin was utilised to inhibit prostanoid production by inhibiting the cyclooxygenase (COX) enzyme. Laser Doppler flowmetry was utilised to measure changes in skin blood flow. Results Not all participants could receive iontophoresis charge due to high skin resistance; these participants were therefore excluded from the analyses. Foot: ACh elicited greater maximal vasodilatation in CAU than AFD following placebo (P = 0.003) and COX inhibition (COXib) (P < 0.001). COXib did not affect blood flow responses in AFD, but caused a reduction in the area under the curve for CAU (P = 0.031). Finger: ACh elicited a greater maximal vasodilatation in CAU than AFD following placebo (P = 0.013) and COXib (P = 0.001). COXib tended to reduce the area under the curve in AFD (P = 0.053), but did not affect CAU. Conclusions CAU have a greater endothelial reactivity than AFD in both foot and finger skin sites irrespective of COXib. It is concluded that the lower ACh-induced vasodilatation in AFD is not due to a compromised COX pathway.Published versio

Sexual Selection on male cuticular hydrocarbons via male-male competition and female choice

This is the final version of the article. Available from the publisher via the DOI in this record.Traditional views of sexual selection assumed that male-male competition and female mate choice work in harmony, selecting upon the same traits in the same direction. However, we now know that this is not always the case and that these two mechanisms often impose conflicting selection on male sexual traits. Cuticular hydrocarbons (CHCs) have been shown to be linked to both social dominance and male attractiveness in several insect species. However, although several studies have estimated the strength and form of sexual selection imposed on male CHCs by female mate choice, none have established whether these chemical traits are also subject to sexual selection via male-male competition. Using a multivariate selection analysis, we estimate and compare sexual selection exerted by male-male competition and female mate choice on male CHC composition in the broad-horned flour beetle Gnatocerus cornutus. We show that male-male competition exerts strong linear selection on both overall CHC abundance and body size in males, while female mate choice exerts a mixture of linear and nonlinear selection, targeting not just the overall amount of CHCs expressed but the relative abundance of specific hydrocarbons as well. We discuss the potential implications of this antagonistic selection with regard to male reproductive success

Pre and Post-copulatory Selection Favor Similar Genital Phenotypes in the Male Broad Horned Beetle.

PublishedJournal ArticleSexual selection can operate before and after copulation and the same or different trait(s) can be targeted during these episodes of selection. The direction and form of sexual selection imposed on characters prior to mating has been relatively well described, but the same is not true after copulation. In general, when male-male competition and female choice favor the same traits then there is the expectation of reinforcing selection on male sexual traits that improve competitiveness before and after copulation. However, when male-male competition overrides pre-copulatory choice then the opposite could be true. With respect to studies of selection on genitalia there is good evidence that male genital morphology influences mating and fertilization success. However, whether genital morphology affects reproductive success in more than one context (i.e., mating versus fertilization success) is largely unknown. Here we use multivariate analysis to estimate linear and nonlinear selection on male body size and genital morphology in the flour beetle Gnatocerus cornutus, simulated in a non-competitive (i.e., monogamous) setting. This analysis estimates the form of selection on multiple traits and typically, linear (directional) selection is easiest to detect, while nonlinear selection is more complex and can be stabilizing, disruptive, or correlational. We find that mating generates stabilizing selection on male body size and genitalia, and fertilization causes a blend of directional and stabilizing selection. Differences in the form of selection across these bouts of selection result from a significant alteration of nonlinear selection on body size and a marginally significant difference in nonlinear selection on a component of genital shape. This suggests that both bouts of selection favor similar genital phenotypes, whereas the strong stabilizing selection imposed on male body size during mate acquisition is weak during fertilization.This work was supported by a Leverhulme Early Career Fellowship (ECF/2010/0067) [to C.M.H.] and a NERC and a Leverhulme Research Fellowship (NE/G005303/1) [to D.J.H.]. The Society for Integrative and Comparative Biology provided assistance for attending the symposium at which this research was presented

Operational sex ratio and density predict the potential for sexual selection in the broad-horned beetle

This is the final version. Available from Elsevier Masson via the DOI in this record.Sexual selection can act on all aspects of the phenotype and the opportunity for selection (I s ) sets its maximal strength. Popular approaches to alter I s include the manipulation of the operational sex ratio (OSR) and/or density, with an increase in I s predicted with a male-biased OSR and at higher density. However, debate continues regarding the utility of I s to measure meaningful changes in the strength of selection, as changes in I s with OSR and density may only reflect stochastic processes. Here we tested whether the manipulation of OSR and density alters I s in the broad-horned flour beetle, Gnatocerus cornutus, a species where males are under intense sexual selection and the targets of selection are known. We also recorded the average number of fights and mating behaviour of individuals in our competitive arenas. We found significant main effects of OSR and density on I s , with the opportunity for selection being highest in male-biased high-density treatments. There were also significant effects of OSR and density on the average number of matings, whereas only density influenced the average number of fights. These results suggest that manipulation of OSR and density influence the opportunity for sexual selection in G. cornutus and our observations of fighting and mating behaviour provide a proximate mechanism for the change in I s .NERCAR

Same-sex sexual behaviour as a dominance display

Same-sex sexual behaviour (SSB) is widespread across taxa. One adaptive hypothesis to explain the occurrence and maintenance of SSB is that it acts to intensify or diminish aggression by providing males with a means to reinforce or resolve dominance. However, evidence for this hypothesis is very limited across taxa and the possibility that SSB acts as an extension of intrasexual competition remains contentious. We investigated the role of SSB in intensifying or diminishing aggression in the broad-horned flour beetle, Gnatocerus cornutus. We tested the hypothesis that SSB is an extension of male-male competition by observing how the occurrence of SSB and the stability of SSB courtship roles (i.e. whether males switched between mounting and being mounted) influenced levels of aggression within pairs. We found that, typically, males rapidly establish fixed SSB roles and moreover that the occurrence of SSB and the stability of SSB roles had a highly significant effect on levels of aggression observed within pairs. Pairs in which one male consistently mounted the other showed significantly lower levels of aggression than pairs in which neither male exhibited SSB or in which males continuously switched SSB roles and attempted to mount each other. Furthermore, males that were consistently on the receiving end of SSB demonstrated lower propensity to court females and had a lower mating success than active males. This pattern was analogous to that found in loser males as a result of fighting. Males that lost fights also courted less and had lower mating success than males that won fights. Our findings provide the first empirical support for the hypothesis that SSB is an extension of male-male competition. Furthermore, our results suggest that SSB may act as a display, allowing males to resolve dominance hierarchies without escalating into an injurious fight

Genotype-by-sex-by-diet interactions for nutritional preference, dietary consumption, and lipid deposition in a field cricket

This is the author accepted manuscript. The final version is available from Springer Nature via the DOI in this recordChanges in feeding behaviour, especially the overconsumption of calories, has led to a rise in the rates of obesity, diabetes, and other associated disorders in humans and a range of animals inhabiting human-influenced environments. However, understanding the relative contribution of genes, the nutritional environment, and their interaction to dietary intake and lipid deposition in the sexes still remains a major challenge. By combining nutritional geometry with quantitative genetics, we determined the effect of genes, the nutritional environment, and their interaction on the total nutritional preference (TP), total diet eaten (TE), and lipid mass (LM) of male and female black field crickets (Teleogryllus commodus) fed one of four diet pairs (DPs) differing in the ratio of protein to carbohydrate and total nutritional content. We found abundant additive genetic variance for TP, TE, and LM in both sexes and across all four DPs, with significant genetic correlations between TE and TP and between TP and LM in males. We also found significant genotype-by-DP and genotype-by-sex-by-DP interactions for each trait and significant genotype-by-sex interactions for TE and LM. Complex interactions between genes, sex, and the nutritional environment, therefore, play an important role in nutrient regulation and lipid deposition in T. commodus. This finding may also help explain the increasing rate of obesity and the maintenance of sex differences in obesity observed across many animal species, including humans.H was funded by a University Royal Society Fellowship and Equipment Grant and by NERC (NE/G00949X/1) and AJW by a BBSRC Fellowship. JR was funded by a NERC studentship (NERC/1200242) awarded to JH

Constrained evolution of the sex comb in Drosophila simulans

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this record.Male fitness is dependent on sexual traits that influence mate acquisition (pre-copulatory sexual selection) and paternity (post-copulatory sexual selection), and while many studies have documented the form of selection in one or the other of these arenas, fewer have done it for both. Nonetheless, it appears that the dominant form of sexual selection is directional, although theoretically, populations should converge on peaks in the fitness surface, where selection is stabilizing. Many factors, however, can prevent populations from reaching adaptive peaks. Genetic constraints can be important if they prevent the development of highest fitness phenotypes, as can the direction of selection if it reverses across episodes of selection. In this study, we examine the evidence that these processes influence the evolution of the multivariate sex comb morphology of male Drosophila simulans. To do this, we conduct a quantitative genetic study together with a multivariate selection analysis to infer how the genetic architecture and selection interact. We find abundant genetic variance and covariance in elements of the sex comb. However, there was little evidence for directional selection in either arena. Significant nonlinear selection was detected prior to copulation when males were mated to non-virgin females, and post-copulation during sperm offence (again with males mated to non-virgins). Thus contrary to our predictions, the evolution of the D. simulans sex comb is limited neither by genetic constraints nor by antagonistic selection between pre- and post-copulatory arenas, but nonlinear selection on the multivariate phenotype may prevent sex combs from evolving to reach some fitness maximising optima. This article is protected by copyright. All rights reserved.This work was supported by a BBSRC fellowship to A.W, Royal Society Fellowship to J.H., a NERC grant and a Leverhulme Research Fellowship to D.J.H. and a Leverhulme Early Career Fellowship to C.M.H. We thank two anonymous reviewers for comments on the manuscript

Rival male chemical cues evoke changes in male pre- and post-copulatory investment in a flour beetle.

This is the final version of the article. Available from Oxford University Press via the DOI in this record.Males can gather information on the risk and intensity of sperm competition from their social environment. Recent studies have implicated chemosensory cues, for instance cuticular hydrocarbons (CHCs) in insects, as a key source of this information. Here, using the broad-horned flour beetle (Gnatocerus cornutus), we investigated the importance of contact-derived rival male CHCs in informing male perception of sperm competition risk and intensity. We experimentally perfumed virgin females with male CHCs via direct intersexual contact and measured male pre- and post-copulatory investment in response to this manipulation. Using chemical analysis, we verified that this treatment engendered changes to perfumed female CHC profiles, but did not make perfumed females "smell" mated. Despite this, males responded to these chemical changes. Males increased courtship effort under low levels of perceived competition (from 1-3 rivals), but significantly decreased courtship effort as perceived competition rose (from 3-5 rivals). Furthermore, our measurement of ejaculate investment showed that males allocated significantly more sperm to perfumed females than to control females. Together, these results suggest that changes in female chemical profile elicited by contact with rival males do not provide males with information on female mating status, but rather inform males of the presence of rivals within the population and thus provide a means for males to indirectly assess the risk of sperm competition.S.M.L.  was funded by a Natural Environment Research Council (NERC) studentship, J.H.S. was funded by NERC, a Royal Society Fellowship, and a Royal Society Equipment Grant (UF120087), and C.M.H. by a Leverhulme Early Career Fellowship (ECF/2010/0067)

Polyandry and fitness in female horned flour beetles, Gnatocerus cornutus

This is the author accepted manuscript. The final version is available from the publisher via the DOI in this record.Although polyandry is common, it is often unclear why females mate with multiple males, because although polyandry may provide females with direct or indirect fitness benefits, it can also be costly. Our understanding of polyandry is also restricted by the relative paucity of studies that disentangle the fitness effects of mating more than once with a single male and mating with multiple males. Here we investigated potential benefits and costs of polyandry in the horned beetle, Gnatocerus cornutus, while controlling for the number of matings. We found that female life span was independent of mating frequency, indicating that mating itself is not very costly. However, females that mated more than once laid more eggs and had greater lifetime reproductive success than singly mated females. Because the magnitude of these effects was similar in monandrous and polyandrous females, this improved fertility was due to multiple mating itself, rather than mating with multiple males. However, although polyandrous females produced more attractive sons, these males tended to have smaller mandibles and so may fare less well in male-male competition. The se results indicate that polyandry is relatively cost free, at least in the laboratory, and has direct and indirect benefits to female fitness. However, because the attractive sons produced by polyandrous females may fight less well, the indirect benefits of polyandry will depend on the intensity of male-male competition and how free females are to exert mate choice. Where competition between males is intense, polyandry benefits via son attractiveness may be reduced and perhaps even carry costs to female fitness.This study was supported by a Grant-in-Aid for Scientific Research (KAKENHI 25840157) from Japanese Ministry of Education, Science, Sports and Culture. We thank the Editor and referees for helpful comments which greatly improved the manuscript