Subtle manipulation of egg sex ratio in birds

Mothers are predicted to overproduce male or female eggs when the relative fitness gains from one sex are higher and outweigh the costs of manipulation. However, in birds such biases are often difficult to distinguish from differential embryo or chick mortality. Using a molecular technique to identify the sex of early embryos, we aim to determine the effect of maternal nutrition on zebra finch (Taeniopygia guttata) egg sex ratios after 2 days of incubation, which is as close to conception as is currently possible. We found no overall bias in the sex ratio of eggs laid and sex did not differ with relative laying order under any diet regime. However, mothers on a low-quality diet did produce a female bias in small clutches and a slight male bias in large clutches. On a high-quality diet, mothers produced a male bias in small clutches and a female bias in large clutches. Those on a standard diet produced a roughly even sex ratio, irrespective of clutch size. These observed biases in egg sex are partly in line with predictions that, in this species, daughters suffer disproportionately from poor rearing conditions. Thus, when relatively malnourished, mothers should only rear daughters in small broods and vice versa. Sex-ratio patterns in this species therefore appear to be subtle.     

Hormonal control of sex ratio

It has been hypothesized that the maternal gonadotrophin level at the time of conception is causally related to the sex of the resultant human zygote, high levels of hormone being associated with the production of female offspring. In this note, evidence for and against this hypothesis is reviewed. There seems so much evidence in its favour that one might conclude that it contains at least a kernel of truth. It is suggested here that of the components of gonadotrophin, the active one in this hypothesized sex-selective process is luteinizing hormone, rather than follicle-stimulating hormone. The hypothesis is nevertheless unable to accommodate several well-established sets of data. Accordingly it is suggested that other hormones, oestrogen and testosterone, have sex-selective properties too, high levels being associated with male offspring. This elaboration of the hypothesis, if it were true, would explain most, if not all, of the epidemiological data on the human secondary sex ratio. In particular it would explain Guerrero's data which have hitherto resisted explanation of any kind. No suggestion is made about possible mechanisms underlying these hypothesized sex-selective properties. But it seems that sex-selective abortion is not the only one. It is hoped that other workers may be stimulated not only to test the hypothesis as outlined here, but--if it survives this testing--to suggest such mechanisms.     

Opposing effects of mortality factors on progeny operational sex ratio may thwart adaptive manipulation of primary sex ratio

Despite extensive research on mechanisms generating biases in sex ratios, the capacity of natural enemies to shift or further skew operational sex ratios following sex allocation and parental care remains largely unstudied in natural populations. Male cocoons of the sawfly Neodiprion abietis (Hymenoptera: Diprionidae) are consistently smaller than those of females, with very little overlap, and thus, we were able to use cocoon size to sex cocoons. We studied three consecutive cohorts of N. abietis in six forest stands to detect cocoon volume‐associated biases in the attack of predators, pathogens, and parasitoids and examine how the combined effect of natural enemies shapes the realized operational sex ratio. Neodiprion abietis mortality during the cocoon stage was sex‐biased, being 1.6 times greater for males than females. Greater net mortality in males occurred because male‐biased mortality caused by a pteromalid parasitic wasp and a baculovirus was greater and more skewed than female‐biased mortality caused by ichneumonid parasitic wasps. Variation in the susceptibility of each sex to each family of parasitoids was associated with differences in size and life histories of male and female hosts. A simulation based on the data indicated that shifts in the nature of differential mortality have different effects on the sex ratio and fitness of survivors. Because previous work has indicated that reduced host plant foliage quality induces female‐biased mortality in this species, bottom‐up and top‐down factors acting on populations can affect operational sex ratios in similar or opposite ways. Shifts in ecological conditions therefore have the potential to alter progeny fitness and produce extreme sex ratio skews, even in the absence of unbalanced sex allocation. This would limit the capacity of females to anticipate the operational sex ratio and reliably predict the reproductive success of each gender at sex allocation.     

Parental care and adaptive brood sex ratio manipulation in birds

Under many circumstances, it might be adaptive for parents to bias the investment in offspring in relation to sex. Recently developed molecular techniques that allow sex determination of newly hatched offspring have caused a surge in studies of avian sex allocation. Whether females bias the primary brood sex ratio in relation to factors such as environmental and parental quality is debated. Progress is hampered because the mechanisms for primary sex ratio manipulation are unknown. Moreover, publication bias against non-significant results may distort our view of adaptive sex ratio manipulation. Despite this, there is recent experimental evidence for adaptive brood sex ratio manipulation in birds. Parental care is a particularly likely candidate to affect the brood sex ratio because it can have strong direct effects on the fitness of both parents and their offspring. We investigate and make predictions of factors that can be important for adaptive brood sex ratio manipulation under different patterns of parental care. We encourage correlational studies based on sufficiently large datasets to ensure high statistical power, studies identifying and experimentally altering factors with sex-differential fitness effects that may cause brood sex ratio skew, and studies that experimentally manipulate brood sex ratio and investigate fitness effects.     

蜜蜂性别决定与性比调控机理研究

叙述了 4个主要蜜蜂性别决定机理的假说 :即性位点假说、基因平衡假说、蜜蜂性别决定综合假说和性基因数量决定假说。然后就蜜蜂性比由蜂王操纵 ,或是由工蜂操纵进行了论述 ,并对蜜蜂性比调控机理研究提出了一些建议     

H-Y antigen in the study of sex determination and control of sex ratio

It has been proposed, on the basis of widespread phylogenetic conservation, that H-Y antigen is the inducer of primary sex, causing the undifferentiated XY gonad to become a testis in male heterogametic species such as the human and bovine. That proposition has withstood extensive testing in vivo and in vitro. Freemartin gonads are H-Y⁺, for example, and masculinization of the freemartin gonad has been attributed to soluble H-Y, borne and transmitted in the serum of the bull twin, and bound in ovarian receptors of the female. We have applied monoclonal H-Y antibodies to the identification of gender in embryos of the bovine. Our preliminary results imply presence of H-Y in bovine embryos of the morula and blastocyst stages recovered at about 6–12 days of gestation. Assignment of H-Y phenotype -- positive in males and negative in females -- allows selective implantation of male and female during embryo transfer. Thus in an early study, we correctly identified gender in 6 of 7 calves born healthy at term, after transfer of 8 blastocysts.     

Evolutionary conflict over the control of offspring sex ratio

There are many theories which predict how animals should control the sex ratio of their offspring. In diploids, however, such control is rarely seen. Two explanations have been suggested for this. One is that parents are simply unable to control the sex ratio of their offspring. The other is that sperm actively oppose such control. This paper examines the possibilities and consequences of parent-gamete conflict over the sex ratio. Such conflict may occur between any of the parties concerned--sperm, ova, fathers, mothers, offspring. It is concluded that gametes are indeed almost always opposed to any parental manipulation of the sex ratio. However, it is probable that the rarity of adaptive parental control of progeny sex ratio in diploids is because parents are physiologically incapable of altering the sex ratio.     

Towards a genetic theory for the evolution of the sex ratio

A genetical model is formulated in which the sex ratio in broods and the relative size of broods are determined by the genotype at an autosomal locus. The results also apply to the case in which the sex-ratio locus is sex linked and expressed in the homogametic sex and to the case in which the locus is expressed in the diploid sex of a haplodiploid organism. Fisher (1930) argued that the sex ratio evolves under natural selection to a value such that parental expenditure is equalized between the sexes. Shaw and Mohler (1953) and MacArthur (1965) proposed that the sex ratio evolves to increase a certain expression for fitness. The sex ratio suggested by Fisher (1930) is in fact identical to the sex ratio specified by these maximization principles. Further, in our model, the Fisherian sex ratio corresponds exactly to the sex ratio at certain equilibria that are approached whenever they exist.     

Towards a genetic theory for the evolution of the sex ratio II. Haplodiploid and diploid models with sibling and parental control of the brood sex ratio and brood size

Population genetic models involving sister, brother, and father control of the brood sex ratio and brood size in both the haplodiploid and diploid cases are constructed and analyzed. The results are interpreted in light of the verbal theories which predict the evolution of the sex ratio to a value which is proportional to the ratio of relatedness of the controlling members of the family to males and to females produced in the brood. In our models, the sex ratio in a certain class of polymorphic equilibria evolves to equal investment in males and females in those cases where the controlling members of the family are symmetrically related to males and females as predicted by the verbal theory. However, the sex ratio in the case of sister control in haploidiploids does not evolve to 1:3, but rather to a value proportional to the ratio of the regression coefficients of additive genotypes. Even so, the predicted sex ratio, which is proportional to 1:3, is in fact an “ESS” in the sense that fixation of a genotype specifying that sex ratio is resistant to the initial increase of all other genotypes.     

类胡萝卜素生物合成途径及其控制与遗传操作

类胡萝卜素在真菌和植物细胞胞液／内质网上是由乙酰CoA经甲羟戊酸途径合成的,在细菌与植物质体中由磷酸甘油醛与丙酮酸经1-脱氧木酮糖-5-磷酸途径合成。形成的异戊烯基焦磷酸经多次缩合生成第一个类胡萝卜素八氢番茄红素,再经脱氢、环化、羟基化、环氧化等转变为其它类胡萝卜素。类胡萝卜素生物合成中涉及的酶都是膜结合的或整合入膜中的。类胡萝卜素合成是通过底物可利用性与环化分支方式进行控制的。白色体到叶绿体的转变以及花与果实成熟时类胡萝卜素合成增加是在基因转录水平调节的。进行类胡萝卜素合成酶基因的转化,可增加转化体类胡萝卜素的积累。     

Ejaculate depletion patterns evolve in response to experimental manipulation of sex ratio in Drosophila melanogaster

We assessed the extent to which traits related to ejaculate investment have evolved in lines of Drosophila melanogaster that had an evolutionary history of maintenance at biased sex ratios. Measures of ejaculate investment were made in males that had been maintained at male-biased (MB) and female-biased (FB) adult sex ratios, in which levels of sperm competition were high and low, respectively. Theory predicts that when the risk of sperm competition is high and mating opportunities are rare (as they are for males in the MB populations), males should increase investment in their few matings. We therefore predicted that males from the MB lines would (1) exhibit increased investment in their first mating opportunities and (2) deplete their ejaculates at a faster rate when mating multiply, in comparison to FB males. To investigate these predictions we measured the single mating productivity of males from three replicates each of MB and FB lines mated to five wild-type virgin females in succession. In contrast to the first prediction, there was no evidence for differences in productivity between MB and FB line males in their first matings. The second prediction was upheld: mates of MB and FB males suffered increasingly reduced productivity with successive matings, but the decline was significantly more pronounced for MB than for FB males. There was a significant reduction in the size of the accessory glands and testes of males from the MB and FB regimes after five successive matings. However, the accessory glands, but not testes, of MB males became depleted at a significantly faster rate than those of FB males. The results show that male reproductive traits evolved in response to the level of sperm competition and suggest that the ability to maintain fertility over successive matings is associated with the rate of ejaculate, and particularly accessory gland, depletion.     

Pre-ovulation control of hatchling sex ratio in the Seychelles warbler

Females of some bird species have a high degree of control over the sex ratio of their offspring at laying. Although several mechanisms have been put forward to explain how females might control the sex of their eggs, virtually nothing is known. As females are the heterogametic sex in birds, adjustment of the clutch sex ratio could arise either by pre- or post-ovulation control mechanisms. The Seychelles warbler (Acrocephalus sechellensis) exhibits extreme adaptive egg sex ratio bias. Typically, warblers produce only single-egg clutches, but by translocating pairs to vacant habitat of very high quality, most females were induced to produce two-egg clutches. Overall, females skewed clutch sex ratios strongly towards daughters (86.6%). This bias was evident in the first egg, but critically, also in the second eggs laid a day apart, even when all absent, unhatched, or unsexed second eggs were assumed to be male. Although a bias in the first egg may arise through either pre- or post-ovulation mechanisms, the skew observed in second eggs could only arise through pre-ovulation control. Post-ovulation adjustment may also contribute to skewed hatchling sex ratios, but as sex-biased release of gametes is likely to be a more efficient process of control, pre-ovulation mechanisms may be the sole means of adjustment in this species. High fitness differentials between sons and daughters, as apparent in the Seychelles warblers, may be necessary for primary sex ratio adjustment to evolve.     

Evidence of genetic and maternal effects on secondary sex ratio in cattle

There is a paucity of estimates of genetic variation for secondary sex ratio (i.e., sex ratio at birth) in dairy cattle. The objective of this study was to estimate the direct and maternal genetic variance as well as maternal permanent environmental variance for offspring sex in dairy herds. The data consisted of 77,508 births from 61,963 dams and 2,859 sires in 1,369 Irish dairy herds across the years 2003 to 2008, inclusive. Mixed models were used to estimate all parameters. Significant genetic variation in sex ratio existed, with a heritability for secondary sex ratio estimated at 0.02; the genetic standard deviation was 0.07 percentage units. No maternal genetic effects on secondary sex ratio were identified but the proportion of phenotypic variance in secondary sex ratio attributable to maternal permanent environmental effects was similar to that attributable to the additive genetic variance (i.e., 0.02). These results, therefore, suggest that the paternal (genetic) influence on secondary sex ratio is just as large as the maternal (non-genetic) influence, both of which are biologically substantial. The results from this study will be useful in generating a sample population of divergent animals for inclusion in a controlled experiment to elucidate the physiological mechanism underpinning differences in secondary sex ratio.     

Effects of worker manipulation on the sex ratio of a Japanese ant species, Myrmecina nipponica

In order to test the effects of colony size and nutritional condition on the survivorship and sex ratio of ants, Myrmecina nipponica colonies were housed in a laboratory in colony sizes of 10 or 30 individuals and fed either daily or weekly. Under all conditions, most of the larvae successfully grew into adults, which suggests that survivorship was not significantly affected by either colony size or nutritional condition. However, the number of new queens was significantly higher in colonies that were fed daily. These results indicate that workers do not control the proportion of diploid and haploid broods by eliminating some larvae and that nutritional condition exerts a significant effect on sex ratio.