Sex allocation within broods: the intrabrood sharing-out hypothesis

Selection is expected to cause parents to adjust the sex oftheir offspring when the environment is predictable during development,and it is expected to affect each sex differently. When severaloffspring compete for limited resources, the environmental conditionsacting on the brood are not a good predictor of the conditionsaffecting individual offspring. There is evidence for some speciesthat, regardless of any bias in brood sex ratio, the sex ofindividual offspring within a brood may be related to its positionin the hatching/birth/weight rank, in ways that might correlatewith the expected share of available resources. Here I proposethat parents may be selected to adjust offspring sex withinthe brood, provided that some depreciable environmental qualityis unequally distributed among siblings in a predictable manner.I call this the "intrabrood sharing-out" hypothesis and presenta graphical model to derive predictions about the relationshipbetween offspring sex and positions within the brood. The modelconsiders that sibling competition not only produces differencesin the mean share of resources among siblings, but it also increasesthe predictability of the share obtained by high-ranking sibsand decreases the predictability of the share for low-rankingones. Consequently, parents should be selected to deal withsuch a distribution by promoting the conditions to make it morepredictable and then adaptively adjust the sex of particularsiblings, especially in high-ranking positions within the brood,rather than to modify the sex ratio of the brood as a whole.     

Violent men have more sons: further evidence for the generalized Trivers-Willard hypothesis (gTWH)

The generalized Trivers-Willard hypothesis (gTWH) [Kanazawa, S., 2005a. Big and tall parents have more sons; further generalizations of the Trivers-Willard hypothesis. J. Theor. Biol. 235, 583-590] proposes that parents who possess any heritable trait which increases the male reproductive success at a greater rate than female reproductive success in a given environment have a higher-than-expected offspring sex ratio, and parents who possess any heritable trait which increases the female reproductive success at a greater rate than male reproductive success in a given environment have a lower-than-expected offspring sex ratio. One heritable trait which increases the reproductive success of sons significantly more than that of daughters in the ancestral environment is the tendency toward violence and aggression. I therefore predict that violent parents have a higher-than-expected offspring sex ratio (more sons). The analysis of both American samples and a British sample demonstrates that battered women, who are mated to violent men, have significantly more sons than daughters.     

Big and tall parents have more sons: further generalizations of the Trivers-Willard hypothesis

This paper proposes the generalized Trivers-Willard hypothesis (gTWH), which suggests that parents who possess any heritable trait which increases male reproductive success at a greater rate than female reproductive success in a given environment will have a higher-than-expected offspring sex ratio, and parents who possess any heritable trait which increases female reproductive success at a greater rate than male reproductive success in a given environment will have a lower-than-expected offspring sex ratio. Since body size (height and weight) is a highly heritable trait which increases male (but not female) reproductive success, the paper hypothesizes that bigger and taller parents have more sons. The analysis of both surviving children and recent pregnancies among respondents of the National Child Development Survey and the British Cohort Survey largely supports the hypothesis.     

Beautiful parents have more daughters: a further implication of the generalized Trivers-Willard hypothesis (gTWH)

The generalized Trivers-Willard hypothesis (gTWH) [Kanazawa, S., 2005. Big and tall parents have more sons: further generalizations of the Trivers-Willard hypothesis. J. Theor. Biol. 235, 583-590) proposes that parents who possess any heritable trait which increases the male reproductive success at a greater rate than female reproductive success in a given environment will have a higher-than-expected offspring sex ratio, and parents who possess any heritable trait which increases the female reproductive success at a greater rate than male reproductive success in a given environment will have a lower-than-expected offspring sex ratio. One heritable trait which increases the reproductive success of daughters much more than that of sons is physical attractiveness. I therefore predict that physically attractive parents have a lower-than-expected offspring sex ratio (more daughters). Further, if beautiful parents have more daughters and physical attractiveness is heritable, then, over evolutionary history, women should gradually become more attractive than men. The analysis of the National Longitudinal Study of Adolescent Health (Add Health) confirm both of these hypotheses. Very attractive individuals are 26% less likely to have a son, and women are significantly more physically attractive than men in the representative American sample.     

Extreme sex ratio variation in relation to change in condition around conception

Adaptive theory predicts that mothers would be advantaged by adjusting the sex ratio of their offspring in relation to their offspring's future reproductive success. Studies investigating sex ratio variation in mammals have produced notoriously inconsistent results, although recent studies suggest more consistency if sex ratio variation is related to maternal condition at conception, potentially mediated by changes in circulating glucose level. Consequently, we hypothesized that change in condition might better predict sex ratio variation than condition per se. Here, we investigate sex ratio variation in feral horses (Equus caballus), where sex ratio variation was previously shown to be related to maternal condition at conception. We used condition measures before and after conception to measure the change in condition around conception in individual mothers. The relationship with sex ratio was substantially more extreme than previously reported: 3% of females losing condition gave birth to a son, whereas 80% of those females that were gaining condition gave birth to a son. Change in condition is more predictive of sex ratio than actual condition, supporting previous studies, and shows the most extreme variation in mammals ever reported.     

Engineers have more sons, nurses have more daughters: an evolutionary psychological extension of Baron-Cohen's extreme male brain theory of autism

In his extreme male brain theory of autism, Baron-Cohen postulates that having a typically male brain was adaptive for ancestral men and having a typically female brain was adaptive for ancestral women. He also suggests that brain types are substantially heritable. These postulates, combined with the insight from the Trivers-Willard hypothesis regarding parental ability to vary offspring sex ratio, lead to the prediction that people who have strong male brains should have more sons than daughters, and people who have strong female brains should have more daughters than sons. The analysis of the 1994 US General Social Survey data provides support for this prediction. Our results suggest potentially fruitful extensions of both Baron-Cohen's theory and the Trivers-Willard hypothesis.     

Female common lizards (Lacerta vivipara) do not adjust their sex-biased investment in relation to the adult sex ratio

Sex allocation theory predicts that facultative maternal investment in the rare sex should be favoured by natural selection when breeders experience predictable variation in adult sex ratios (ASRs). We found significant spatial and predictable interannual changes in local ASRs within a natural population of the common lizard where the mean ASR is female-biased, thus validating the key assumptions of adaptive sex ratio models. We tested for facultative maternal investment in the rare sex during and after an experimental perturbation of the ASR by creating populations with female-biased or male-biased ASR. Mothers did not adjust their clutch sex ratio during or after the ASR perturbation, but produced sons with a higher body condition in male-biased populations. However, this differential sex allocation did not result in growth or survival differences in offspring. Our results thus contradict the predictions of adaptive models and challenge the idea that facultative investment in the rare sex might be a mechanism regulating the population sex ratio.     

Experimental manipulation of maternal effort produces differential effects in sons and daughters: implications for adaptive sex ratios in the blue-footed booby

Sex allocation theory predicts that mothers in good conditionshould bias their brood sex ratio in response to the differentialbenefits obtained from increased maternal expenditure in sonsand daughters. Although there is well-documented variationof offspring sex ratios in several bird species according tomaternal condition, the assumption that maternal condition has different fitness consequences for male and for female offspringremains unclear. The blue-footed booby (Sula nebouxii) is asexually size-dimorphic seabird, with females approximately31% heavier than males. It has been reported that the sex ratiois male biased in years with poor feeding conditions, whichsuggests that either females adjust their sex ratio in accordancewith their condition or that they suffer differential brood mortality before their sex can be determined. In this studyI tested whether the condition of mothers affected their daughters'fitness more than their sons' fitness. I manipulated maternalinvestment by trimming the flight feathers and thereby handicappingfemales during the chick-rearing period. Adult females in thehandicapped group had a poorer physical condition at end ofchick growth, as measured by mass and by the residuals of masson wing length compared to control birds. Female chicks wereaffected by the handicapping experiment, showing a lower massand shorter wing length (reduced approximately 8% in both measures)than controls. However, this effect was not found in male chicks.Hatching sex ratios were also related to female body conditionat hatching. The brood sex ratio of females in poor conditionwas male biased but was female biased for females in good condition.Overall, these results suggest that the variation in the sexratio in blue-footed boobies is an adaptive response to thedisadvantage daughters face from being reared under poor conditions.     

Resource allocation in a social wasp: effects of breeding system and life cycle on reproductive decisions

Organisms must make important decisions on how to allocate resources to reproduction. We investigated allocation decisions in the social wasp Vespula maculifrons to understand how social insects make reproductive choices. We first determined how annual colonies apportioned resources to growth and reproduction by analysing developing brood. In contrast to expectations, colonies invested in both growth (workers) and reproduction (males) simultaneously. In addition, colonies showed evidence of producing males in pulses and reversing their reproductive choices by decreasing investment in males late in the season. This reversal is consistent with theory suggesting that colonies decrease production in males if fitness of late emerging males is low. To further investigate reproductive decisions within colonies, we determined if the male mates of multiply-mated queens varied in their reproductive success over time. Sperm use by queens did vary over time suggesting that male success may depend on sperm clumping within the female reproductive tract. Finally, we tested if colony sex ratio conformed to expectations under kin selection theory that nestmate relatedness would positively correlate with investment in new queens if workers controlled sex allocation. Surprisingly, the proportion of queens produced by colonies was negatively correlated with nestmate relatedness, suggesting that allocation may be shaped by advantages arising from increased genetic diversity resulting from multiple mating by queens. Overall, our study suggests that the reproductive decisions of colonies are flexible and may depend both on environmental cues arising from energetic needs of the colony and genetic cues arising from mating behaviours of queens.     

Optimal investment allocation in primitively eusocial bees: a balance model based on resource limitation of the queen

The classical model of colony dynamics developed by Macevicz and Oster predicts that optimal colony fitness in annual eusocial insects is achieved by a bang-bang strategy of reproduction: exclusive production of workers (ergonomic phase) followed by exclusive production of sexuals (reproductive phase). We propose an alternative model that assumes colony development in discrete broods and a limited overall investment potential of the queen. Based on the costs for producing eggs, workers, and sexuals and efficiency of individuals we predict the optimal number of workers and sexuals in the colony for each brood of the colony cycle that maximizes overall colony fitness. To link our model assumptions to the real world we chose model parameters according to field data of the halictid bee Lasioglossum malachurum. However, our model is representative of a large number of species with an annual life cycle and with discrete broods. Our model shows that the optimal partitioning of resources, i.e. the optimal workers/sexuals ratio depends on rearing cost for sexuals as well as productivity of workers but not on the queens’ total investment, egg cost, or rearing cost for workers. In complete accordance to Macevicz and Oster we predict a bang-bang reproduction strategy despite the differences in the basic assumptions. Potential deviations from this strategy and transitions from social to solitary breeding are discussed in the framework of our model. Received 31 October 2006; revised 29 March 2007; accepted 17 April 2007.     

A new model for the effect of pH on microbial growth: an extension of the Gamma hypothesis

Aims: To investigate the appropriateness of the extended Lambert–Pearson model (ELPM) to model the effect of pH (as hydrogen and hydroxyl ions) over the whole biokinetic pH range in comparison with other available models. Methods and Results: Data for the effect of pH on microbial growth were obtained from the literature or in‐house. Data were examined using several models for pH. Models were compared using the residual mean of squares. Using the ELPM, pH was modelled as hydrogen ions and hydroxyl ions; hence, the model was monotonic in each. The ELPM was able to model data more successfully than the cardinal pH model (CPM) and other models in the majority of cases. Conclusions: Examining the effect of pH as hydrogen and hydroxyl ions has the advantage that the basic form of the ELPM can be retained as each is treated as a distinct antimicrobial effect. With the ELPM, each inhibitor is described by two parameters; from these parameters, the pH_min, pH_opt and pH_max can be obtained. Furthermore, the idea of a dose response, absent from other models, becomes important. Significance and Impact of the Study: The CPM is an excellent model for certain situations – where there is a high degree of symmetry between the suboptimal pH and superoptimal pH response and where there are few data points available. The ELPM is more amenable to highly asymmetric behaviour, especially where plateaus of effect around the pH optimum are observed and where the number of data points is not restrictive.     

Sex-specific growth rates in zebra finch nestlings: a possible mechanism for sex ratio adjustment

Wild and captive zebra finches (Taenopygia guttata), like severalother species, produce a male-biased sex ratio at fledging whenfood is scarce. This is due to primary sex-ratio adjustmentand female-biased nestling mortality. Given that young femalesfledging at low body masses have been shown to have low fecundityas adults, lower returns to parents from producing female offspringin conditions of restricted food has been raised as a functionalexplanation (Trivers and Willard's hypothesis of adaptive sexualinvestment; 1973). However, an alternative, mechanistic hypothesisis that under restricted conditions female chicks are more costlyto produce. In consequence, lower returns to parents under theseconditions would happen earlier in the life of female offspringrather than later. To test this hypothesis, I hand-reared chickson a food gradient. In the absence of parent-offspring and sib-sibinteractions, final body mass and growth rates for females werelower in conditions of restricted food. For males, final bodymass and growth rates did not differ with food condition. Lowfemale growth rates in food-restricted conditions might be onepotential mechanism causing female-biased mortality in birds.More importantly, this result is the strongest evidence yetof female offspring experiencing higher marginal fitness benefitsfrom additional food than males and it has implications forprimary and secondary sex-ratio adjustment. Also, as this mechanismhas been shown in the absence of parent-offspring interactions,significant questions can now be raised as to how parental andoffspring behavior interact in their effects on secondary sex-ratioadjustment.     

Seed sex ratio in dioecious plants depends on relative dispersal of pollen and seeds: an example using a chessboard simulation model

Two principles are important for the optimal sex ratio strategy of plants. (1) Sib mating. Because seed dispersal is restricted, sib mating may occur which selects for a female bias in the seed sex ratio. (2) Local resource competition (LRC). If a plant produces pollen its nuclear genes are dispersed in two steps: first through the pollen and then, if the pollen is successful in fertilizing an ovule on another plant, through the seed. If the plant produces an ovule, its genes are dispersed only through the seed. By making pollen instead of ovules the offspring of a single plant is then spread out over a wider area. This reduces the chance that genetically related individuals are close together and need to compete for the same resource. The effect is the strongest if pollen is dispersed over a much wider area than seeds. Less LRC for paternally vs. maternally derived offspring selects for a male bias in sex allocation. We study the above‐mentioned opposite effects in dioecious plants (with separate male and female individuals), with maternal control over the sex ratio (fraction males) in the seeds. In a two‐dimensional spatial model female‐biased sex ratios are found when both pollen and seed dispersal are severely restricted. If pollen disperses over a wider area than seeds, which is probably the common situation in plants, the seed sex ratio becomes male‐biased. If pollen and seeds are both dispersed over a wide area, the sex ratio approaches 0.5. Our results do not change if the offspring of brother–sister matings are less fit because of inbreeding depression.     

A model of neurovascular coupling and the BOLD response: PART I

The mechanisms with which neurons communicate with the vasculature to increase blood flow, termed neurovascular coupling is still unclear primarily due to the complex interactions between many parameters and the difficulty in accessing, monitoring and measuring them in the highly heterogeneous brain. Hence a solid theoretical framework based on existing experimental knowledge is necessary to study the relation between neural activity, the associated vasoactive factors released and their effects on the vasculature. Such a framework should also be related to experimental data so that it can be validated against repetitive experiments and generate verifiable hypothesis. We have developed a mathematical model which describes a signaling mechanism of neurovascular coupling with a model of pyramidal neuron and its corresponding fMRI BOLD response. In the first part of two papers we describe the integration of the neurovascular coupling unit extended to include a complex neuron model, which includes the important Na/K ATPase pump, with a model that provides a BOLD signal taking its input from the cerebral blood flow and the metabolic rate of oxygen consumption. We show that this produces a viable signal in terms of initial dip, positive and negative BOLD signals.