Effect of cultural inheritance of reproductive compensation on the incidence of a sex-linked lethal disease

For a sex-linked recessive lethal disease, a model is constructed to study reproductive compensation for having at least one normal male with partial cultural inheritance. The equilibrium frequency of heterozygous females depends on the probability that a female offspring of the compensating parents will not compensate, α, the probability that a female offspring of the non-compensating parents will compensate, β, and the mutation rate, u. When α = 0, the equilibrium frequency of heterozygous females is given by √2u, whereas when α ≠ 0 it is given by 2[{β +(2α + β) (α + β)}/{α(α + β)}]u.Then, the proportion of affected males due to fresh mutations is much smaller than Haldane's estimate of $\text{1}\text{3}$ without reproductive compensation.     

A theoretical study of calcium entry in nerve terminals, with application to neurotransmitter release

In his study of a kin selection model for the evolution of workers' behavior in an incipiently social insect, Craig (1979) found that the haplodiploid mode of sex determination, combined with a female-biased sex ratio, cannot accelerate an evolutionary trend toward eusociality. This seems to contradict to Hamilton's (1964) theory. It is my intention to prove that Craig's result is due to the dependence of relative reproductive success in each sex on the sex ratio of a population. The oviposition of unfertilized eggs by workers is indispensable, in primitive stages in the evolution of eusociality, for maintaining the relative reproductive success of a female. Assuming that workers control the sex ratios, we can distinguish the following three evolutionary phases in accordance with the ratio $\text{N}{}_2\text{N}{}_{10}$ (the ratio of the number of the queen's second brood to that of the progeny laid by workers derived from the same nest). During Phase 1 in which $\text{N}{}_2\text{N}{}_{10}$ < $\text{1}\text{2}$, the reproductive successes of a male and a female are equal, and Hamilton's rule holds. In Phase 2 in which $\text{1}\text{2 <}\text{N}{}_2\text{N}{}_{10}\text{<}\text{3}\text{2}$, the queen produces females, and workers oviposit males. As $\text{N}{}_2\text{N}{}_{10}$ increases, the sex ratio in the whole population becomes female-biased and relative reproductive success of a male increases. In Phase 3 in which N₂/N₁₀ >$\text{3}\text{2}$, the queen lays some male eggs in addition to female eggs. The sociality threshold ($\text{B}\text{C}$)_crit the minimum value of the benefit/cost ratio leading to the evolution of altruism, is $\text{2}\text{3}$on Phase 1. It rises threefold as N₂/N₁₀ increases in Phase 2, and, in Phase 3, it is twice as high as that in a diploid species. The anatomical and physiological characteristics of workers must have been so developed that they are efficient in helping activities after the beginning of eusociality. The evolutionary process before the beginning of helping behavior is also discussed. The egg laying by unmated females, which stay their mother's nest, seems to have been an important preadaptation for the evolution of eusociality in Hymenoptera.     

Dominance and aggression in social groups of male and female rats

Two experiments were performed to examine aggression and dominance in domestic male and female $\text{Rattus norvegicus}$ living in small mixed-sex (3 males and 3 females) groups. Experiment 1 examined the development of aggression in females. A single female (alpha) within each of the six colonies tested showed the preponderance of attacks on male intruders placed into the home-cage when male colony residents were absent. Over 12 weeks of intruder-aggression training female alphas showed only a mild nonsignificant elevation of aggressive behavior. A comparison of aggression of male and female colony alphas tested with opponents of each sex revealed that aggression was mainly directed at like-sex opponents, and that female attack was more defensive in character than male attack regardless of opponent sex. The highest intensity of aggression occurred when male alphas confronted male intruders. Although intruders never showed offense toward male residents, 61% of intruding males showed offense in response to attack by females.Experiment 2 investigated the relationship between aggressive dominance and competitive measures of dominance within each of 10 mixed-sex colonies. Alpha stat s of male and female colony residents did not reliably predict priority of access to food or water in tests of direct resource competition with like-sex colony members. When colony males were simultaneously tested for copulation, the copulatory behavior of alpha males was significantly greater than that of other colony males. Results are discussed in relation to the role of aggression in the reproductive strategy of male and female $\text{Rattus norvegicus}$.     

Mating frequency, genetic structure, and sex ratio in the intermorphic female producing ant species Myrmecina nipponica

1. Myrmecina nipponica has two types of colonies: a queen colony type, in which the reproductive females are queens and new colonies are made by independent founding, and an intermorphic female colony type, in which reproductive females belong to a wingless intermediate morphology between queen and worker, and where colonies multiply through colonial budding. 2. The mating frequencies of reproductive females in both types indicate monoandry. The relatedness among nestmates in both types was almost 0.75, however relatedness between mother and daughter in intermorphic female colonies was slightly higher than that of queen colonies. 3. The sex ratio (corrected investment female ratio) was 0.70 at the population level, suggesting that the sex ratio is controlled by workers in this species, however the ratio differed greatly between the two types of colonies. Queen colonies (n = 37) had a female‐biased sex ratio of 0.77 while intermorphic female colonies (n = 33) had a ratio of 0.56. 4. Each reproductive intermorphic female was accompanied by an average of 2.9 workers (including virgin intermorphic females) in the colonial budding, and when the investment to those workers was added to the female investment, the sex ratio reached 0.81. 5. The frequency distribution of sex ratio was bimodal, with many colonies producing exclusively males or females, however mean estimated relatedness within colonies was almost 0.75. These data are inconsistent with the genetic variation hypothesis, which is one of the predominant hypotheses accounting for the between‐colony variation in sex ratio.     

Exposure of embryos to an aromatization inhibitor increases copulatory behaviour of male quail

This experiment examined the possibility that endogenous embryonic androgen contributes to sexual differentiation of behaviour in male or female quail ($\text{Coturnix}$$\text{coturnix}$$\text{japonica}$), and that it does so via aromatization (conversion to oestrogen). Eggs were injected on day 9 of incubation with oil or ATD (an aromatization inhibitor). As adults, males and females were exposed to short days, injected with testosterone propionate, tested for male-typical behaviour, then injected with oestradiol benzoate and tested for female-typical receptivity. ATD increased the level of male-typical copulatory behaviour in males. Male-typical behaviour in females was not affected, nor was female-typical behaviour in either sex. Thus normal male quail are actually slightly demasculinized by their own androgen during embryonic development, and this process is mediated by aromatization.     

COLONY SEX RATIOS,CONFLICT BETWEEN QUEENS AND WORKERS,AND APPARENT QUEEN CONTROL IN THE ANT PHEIDOLE DESERTORUM

Sex-ratio conflict between queens and workers was explored in a study of colony sex ratios, relatedness, and population investment in the ant Pheidole desertorum. Colony reproductive broods consist of only females, only males, or have a sex ratio that is extremely male biased. Colonies producing females (female specialists) and colonies producing males (male specialists) occur at near equal frequency in the population. Most colonies apparently specialize in producing one reproductive sex throughout their life. Allozyme analyses show that relatedness does not differ within male-specialist and female-specialist colonies and they do not appear to differ in available resources. In the population, workers are nearly three times more closely related to females than males; however, the investment sex ratio is near equal (1.01, female/male), which is consistent with queen control. Selection should be strong on workers to increase investment in reproductive females, so why do workers in male-specialist colonies produce only (or nearly only) males? One hypothesis is that queens in male-specialist colonies prevent the occurrence of reproductive females, perhaps by producing worker-biased female eggs. An earlier simulation study of genetic evolution of sex ratios in social Hymenoptera (Pamilo 1982b) predicts that such mechanisms can result in the evolution of bimodal colony sex ratios and queen control. Results on P. desertorum are generally consistent with that study; however, information is not currently available to test some of the model's predictions and assumptions.     

Suppression of reproductive maturation in male-stimulated virgin female microtus by a female urinary chemosignal

Urine from female $\text{Microtus ochrogaster}$ possesses a chemosignal that suppresses reproductive maturation in other females. Uterine enlargement in virgin females stimulated by a male was suppressed by subsequent association with another female or by application of female urine on the nose. Females so suppressed are not able to achieve estrus. Urine from virgin sibling and non-sibling females and from pregnant females possesses the suppressing effect.     

Workers' sons rescue genetic diversity at the sex locus in an invasive honey bee population

The hallmark of eusociality is the division of labour between reproductive (queen) and nonreproductive (worker) females. Yet in many eusocial insects, workers retain the ability to produce haploid male offspring from unfertilized eggs. The reproductive potential of workers has well‐documented consequences for the structure and function of insect colonies, but its implications at the population level are less often considered. We show that worker reproduction in honey bees can have an important role in maintaining genetic diversity at the sex locus in invasive populations. The honey bee sex locus is homozygous‐lethal, and, all else being equal, a higher allele number in the population lead to higher mean brood survival. In an invasive population of the honey bee Apis cerana in Australia, workers contribute significantly to male production: 38% of male‐producing colonies are queenless, and these contribute one‐third of all males at mating congregations. Using a model, we show that such male production by queenless workers will increase the number of sex alleles retained in nascent invasive populations following founder events, relative to a scenario in which only queens reproduce. We conclude that by rescuing sex locus diversity that would otherwise be lost, workers' sons help honey bee populations to minimize the negative effects of inbreeding after founder events and so contribute to their success as invaders.     

REPRODUCTIVE SKEW AND SPLIT SEX RATIOS IN SOCIAL HYMENOPTERA

Abstract I present a model demonstrating that, in social Hymenoptera, split sex allocation can influence the evolution of reproductive partitioning (skew). In a facultatively polygynous population (with one to several queens per colony), workers vary in their relative relatedness to females (relatedness asymmetry). Split sex‐ratio theory predicts that workers in monogynous (single‐queen) colonies should concentrate on female production, as their relatedness asymmetry is relatively high, whereas workers in the polygynous colonies should concentrate on male production, as their relatedness asymmetry is relatively low. By contrast, queens in all colonies value males more highly per capita than they value females, because the worker‐controlled population sex ratio is too female‐biased from the queens' standpoint. Consider a polygynous colony in a facultatively polygynous population of perennial, social Hymenoptera with split sex ratios. A mutant queen achieving reproductive monopoly would gain from increasing her share of offspring but, because the workers would assess her colony as monogynous, would lose from the workers rearing a greater proportion of less‐valuable females from the colony's brood. This sets an upper limit on skew. Therefore, in social Hymenoptera, skew evolution is potentially affected by queen‐worker conflict over sex allocation.     

The development of lactate and malate dehydrogenases in the C57BL-6 mouse kidney

The development of lactate dehydrogenase (LDH; EC 1.1.1.27) and malate dehydrogenase (MDH; EC 1.1.1.37) was measured in the kidney of male and female $\text{C57BL}\text{6}$ mice from ages prenatal 16 days to 80 days. Maximum reactions rates of the enzymes were measured in vitro by following the reduction of the nicotinamide-adenine dinucleotide spectrophotometrically.Analysis of variance showed no significant sex difference for LDH and MDH. There was a significant sex difference for the ratio LDH:MDH and a significant age difference for LDH, MDH, and the ratio LDH:MDH. In the male and female, LDH activity increased from prenatal 16 days to 30 days. Malate dehydrogenase activity reached adult values at 22 days in the male and at 30 days in the female. The ratio LDH:MDH in the male decreased from prenatal 16 days to 3 days, after which the ratio continued to decline to 20 days at a less rapid rate. This general pattern was also found in the female followed by a further decline in the ratio at 50 days.The development of LDH and MDH in the $\text{C57BL}\text{6}$ mouse is tissue specific and probably parallels the development of the tissue's function. In the case of the kidney, LDH and MDH development may reflect maturation of mitochondrial function and the kidney's ability to concentrate urine.     

Sex ratio determination by queens and workers in the ant Pheidole desertorum

Because workers in colonies of eusocial Hymenoptera are more closely related to sisters than to brothers, theory predicts workers should bias investment in reproductive broods to favour reproductive females over males. However, conflict between queens and workers is predicted. Queens are equally related to daughters and sons, and should act to prevent workers from biasing investment. Previous study of the ant Pheidole desertorum showed that workers are nearly three times more closely related to reproductive females than males; however, the investment sex ratio is very near equal, consistent with substantial queen control of workers. Near-equal investment is produced by an equal frequency of colonies whose reproductive broods consist of only females (female specialists) and colonies whose reproductive broods consist of only males or whose sex ratios are extremely male biased (male specialists). Because natural selection should act on P. desertorum workers to bias investment in favour of reproductive females, why do workers in male-specialist colonies rear only (or mostly) males? We tested the hypothesis that queens prevent workers from rearing reproductive females by experimentally providing workers with immature reproductive broods of both sexes. Workers reared available reproductive females, while failing to rear available males. Worker preference for rearing reproductive females is consistent with queens preventing their occurrence in colonies of male specialists. These results provide evidence that queens and workers will act in opposition to determine the sex ratio, a fundamental prediction of queen-worker conflict theory. Copyright 2000 The Association for the Study of Animal Behaviour.     

Karyotypic normalcy and quasi-normalcy of developmentally totipotent mouse teratocarcinoma cells

Malignant mouse teratocarcinoma cells are, in some cases, able to undergo normal, complete differentiation after injection into blastocysts. Thus far, only three lines—of unrelated origin—have been found (all in this laboratory) to be developmentally totipotent in blastocyst tests. The karyotypes of these lines, and their somatic- and germ-cell derivatives, were investigated by G-banding methods, as a possible clue to their developmental superiority. The first, OTT 6050 (129 strain), is an embryo-derived induced tumor maintained as an ascites transplant line. Its stem cells (from embryoid body “cores”) have 40 chromosomes in the modal class, which comprises two subclasses: one all normal and one with a metacentric chromosome (isochromosome-8). However, mosaic animals from injected blastocysts have only the normal subclass in their teratocarcinomaderived cells; all are of $\text{X}\text{Y}$ male sex chromosome type. Presence of the Y chromosome was verified after transmission through the germ line of two fertile mosaic males, in their F₁ male progeny. The second teratocarcinoma line, 72484-395 (LT strain), is a spontaneous ovarian solid tumor maintained by subcutaneous transplantation. Karyotypes of cells from the tumor, and also of teratocarcinoma-derived cells in mosaic animals, were normal and of $\text{X}\text{X}$ female sex chromosome type. Karyotypes of the F₁ progeny, from tumor-strain germ cells of a fertile mosaic female, were also normal. The third line, NG 2 (129 strain), is a mutant clonal in vitro line deficient in hypoxanthine phosphoribosyltransferase. It originated from an embryo-derived experimental tumor (OTT 5568) that was established in culture (PSA1 line); the culture was then mutagenized and selected for 6-thioguanine resistance. The NG 2 line proved to be quasi-normal, with only two karyotypic anomalies: trisomy of chromosome 6 and $\text{X}\text{O}$ female sex chromosome constitution. Thus, developmental totipotency in all three lines, including one maintained in vitro, is accompanied by karyotypic normalcy or near-normalcy. Other culture lines reported to be aneuploid have not yet given evidence of totipotency. Karyotypic normalcy may therefore have predictive value useful in choosing teratocarcinoma lines with relatively high developmental prospects. This is of importance in identifying those mutant lines that would be promising candidates for introduction, via blastocyst injection, of specific mutant genes into mice.     

Male demographic constraints to extreme sex ratio in the twospotted spider mite

Summary Twospotted spider mites, Tetranychus urticae Koch are arrhenotokous. As a result of this genetic structure, primary and secondary sex ratios diverge from the 1:1 female:male ratio commonly found in diploid systems. Ratios vary, but 3:1 is the most common. The influence of life history parameters on spider mite sex ratio is unclear, although maternal genetic effects, resource quality and maternal age are known to play a role. An area that has not been studied is the relationship between male reproductive capabilities and spider mite sex ratio. A prior study on male reproduction in spider mites suggests that males have far too high a reproductive capacity to explain sex ratio patterns, but that study was not conducted under realistic mating conditions. Thus, this study was conducted to determine if there is a link between male reproduction and spider mite sex ratio. This was done by exposing males to various exposure regimes of females and recording the number of copulations, inseminations and daughters fathered by each male. Results include the following: i) males are most virile when one day old; ii) virgin males become nearly completely devoid of sperm (or other copulatory products) after about 15 matings and then take about four days to recover; and iii) the optimal number of matings per day (defined as that which produces the greatest number of daughters in the least number of inseminations) is four females per day. The principle conclusion is that although males have a high potential reproductive output, insemination quality is only high for the first four or five matings each day. Therefore, to ensure full inseminations of all daughters by sons, females are constrained to producing primary sex ratios of about 5:1 or less.     

Sexual selection in gallus: Effects of morphology and dominance on female spatial behavior

Sexual selection has received a great deal of attention from field and laboratory researchers for over a century, but hard evidence of female choice of mates in mixed-sex groups remains scarce. Dominant males typically mate much more often than subordinants but evidence that females “choose” such males is elusive. In $\text{Gallus}$, which includes junglefowl and their domestic fowl descendants, females stay near and mate with dominant, territorial males. We demonstrate here that $\text{Gallus}$ females in mixed-sex flocks who have no information about the social dominance status of males orient to, approach and stay near males with larger than average combs whereas females which $\text{do}$ not have information about male dominance orient toward and stay near high ranking males. We verify that comb size correlates with male social rank. Hence, $\text{Gallus}$ females do actively respond to “high quality” males; they apparently identify such males by male physical characteristics and, if available, information concerning male-male interactions.     

Heritable determinants of left-right bias in the rat

The offspring of matings of rats having opposite or same-sided turning biases were tested for turning biases as adults and the degree of similarity to the parents' biases assessed. There were significant and equivalent tendencies for the male offspring to have the same bias as the male parent and the opposite bias as the female parent. Although, overall, female offspring were distributed randomly with respect to the parents' biases, a significant tendency for female offspring to have biases opposite those of the female parent was apparent in litters having more males than females. Based on reports indicating a relationship between the sex ratio of a litter and levels of testosterone in female fetuses, it was suggested that $\text{in utero}$ exposure to testosterone reverses the coding of a heritable female influence and induces a tendency for the offspring to have biases opposite those of the female parent. The origins of sidedness in the rat appear to involve a complex interaction between heredity and hormones.     

Evidence for Strategic Sex Allocation in the Guppy (Poecilia reticulata): Brood Sex Ratio and Size Predict Subsequent Adult Male Mating Success

Sex allocation theory predicts that females should produce more sons when the reproductive success of sons is expected to be high, whereas they should produce more daughters, not daughters when the reproductive success of sons is expected to be low. The guppy (Poecilia reticulata) is a live‐bearing fish, and female guppies are known to produce broods with biased sex ratios. In this study, we examined the relationship between brood sex ratio and reproductive success of sons and daughters, to determine whether female guppies benefit from producing broods with biased sex ratios. We found that sons in male‐biased broods had greater mating success at maturity than sons in female‐biased broods when brood sizes were larger. On the other hand, the reproductive output of daughters was not significantly affected by brood sizes and sex ratios. Our results suggest that female guppies benefit from producing large, male‐biased brood when the reproductive success of sons is expected to be high.     

Comparison in male and female rats of ACTH content and response to corticotrophin-releasing factor (CRF) of cultured adenohypophyseal cells and in vivo hypothalamic CRF content.

Although mean ACTH concentration was significantly higher in cultured adenohypophyseal cells from female than from male rats, there was no significant sex difference in the ACTH secretory response of these cells to hypothalamic extract containing CRF. Hypothalamic CRF content in the “basal” state $\text{in}$$\text{vivo}$ was slightly higher in female than $\text{in}$ male rats.     

Guppies control offspring size at birth in response to differences in population sex ratio

Females that invest adaptively in their offspring are predicted to channel more resources to the sex that will be at an advantage in the prevailing environmental conditions. Here, we report, for the first time, that female Trinidadian guppy, Poecilia reticulata, respond in reproductively distinct ways when faced with differences in operational sex ratio. We show that females assigned to a female‐biased sex ratio produce larger male offspring than females in an environment in which males predominate. Given the link between size at birth and fitness, and the marked reproductive skew in this species, larger male offspring are expected to have reproductive advantages in guppy populations with an excess of females. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 414–419.     

Evidence for an even sex allocation in haplodiploid cyclical parthenogens

Recent theoretical work has shown that haplodiploid cyclical parthenogens, such as rotifers, are expected to have an equal frequency of male‐producing and resting‐egg producing females during their sexual phase. We tested this prediction by following sexual reproduction dynamics in two laboratory populations and one field population of the rotifer Brachionus plicatilis through two growing seasons. We recorded population density, proportion of sexual females, and sex allocation (the proportion of male‐producing sexual females as a fraction of total sexual females). We found this sex allocation ratio to vary from 0.3 to 1.0 in single sampling events. However, when we computed sex allocation by using the integrated densities of both male‐producing sexual females and resting‐egg producing sexual females over time, the two laboratory populations and one of the two field growing seasons showed sex allocation ratios that did not significantly differ from the expected value of 0.5.