The human sex ratio from conception to birth.

We describe the trajectory of the human sex ratio from conception to birth by analyzing data from (i) 3- to 6-d-old embryos, (ii) induced abortions, (iii) chorionic villus sampling, (iv) amniocentesis, and (v) fetal deaths and live births. Our dataset is the most comprehensive and largest ever assembled to estimate the sex ratio at conception and the sex ratio trajectory and is the first, to our knowledge, to include all of these types of data. Our estimate of the sex ratio at conception is 0.5 (proportion male), which contradicts the common claim that the sex ratio at conception is male-biased. The sex ratio among abnormal embryos is male-biased, and the sex ratio among normal embryos is female-biased. These biases are associated with the abnormal/normal state of the sex chromosomes and of chromosomes 15 and 17. The sex ratio may decrease in the first week or so after conception (due to excess male mortality); it then increases for at least 10-15 wk (due to excess female mortality), levels off after ∼20 wk, and declines slowly from 28 to 35 wk (due to excess male mortality). Total female mortality during pregnancy exceeds total male mortality. The unbiased sex ratio at conception, the increase in the sex ratio during the first trimester, and total mortality during pregnancy being greater for females are fundamental insights into early human development.

Partner choice and fidelity stabilize coevolution in a Cretaceous-age defensive symbiosis.

Many insects rely on symbiotic microbes for survival, growth, or reproduction. Over evolutionary timescales, the association with intracellular symbionts is stabilized by partner fidelity through strictly vertical symbiont transmission, resulting in congruent host and symbiont phylogenies. However, little is known about how symbioses with extracellular symbionts, representing the majority of insect-associated microorganisms, evolve and remain stable despite opportunities for horizontal exchange and de novo acquisition of symbionts from the environment. Here we demonstrate that host control over symbiont transmission (partner choice) reinforces partner fidelity between solitary wasps and antibiotic-producing bacteria and thereby stabilizes this Cretaceous-age defensive mutualism. Phylogenetic analyses show that three genera of beewolf wasps (Philanthus, Trachypus, and Philanthinus) cultivate a distinct clade of Streptomyces bacteria for protection against pathogenic fungi. The symbionts were acquired from a soil-dwelling ancestor at least 68 million years ago, and vertical transmission via the brood cell and the cocoon surface resulted in host-symbiont codiversification. However, the external mode of transmission also provides opportunities for horizontal transfer, and beewolf species have indeed exchanged symbiont strains, possibly through predation or nest reuse. Experimental infection with nonnative bacteria reveals that--despite successful colonization of the antennal gland reservoirs--transmission to the cocoon is selectively blocked. Thus, partner choice can play an important role even in predominantly vertically transmitted symbioses by stabilizing the cooperative association over evolutionary timescales.

Biogeography of a defensive symbiosis.

Mutualistic microorganisms play important roles in nutrition, reproduction and defense of many insects, yet the factors contributing to their maintenance and dispersal remain unknown in most cases. Theory suggests that collaboration can be maintained by repeated interaction of the same partners (partner fidelity) or by selective discrimination against non-cooperative partners (partner choice). In the defensive mutualism between solitary beewolf wasps and their antibiotic-producing Streptomyces bacteria, partner choice by host control of vertical symbiont transmission reinforces partner fidelity and has helped to maintain this highly specific association since it originated in the late Cretaceous. However, co-phylogenetic and biogeographic analyses suggest that there has also been considerable horizontal transmission of the symbionts. While the beewolves clearly have a paleotropic or palearctic origin, with later colonization of the nearctic and neotropics via Beringia and the Aves ridge, respectively, the bacteria show only weak geographical clustering, implying global dispersal or vicariance within the confines of an otherwise apparently exclusive symbiotic relationship. We discuss several hypotheses that may explain these patterns. Future studies investigating the occurrence of beewolf symbionts in the environment could yield broadly applicable insights into the relative impact of animal-vectored and free-living dispersal on the distribution of microorganisms in nature.

Resource transfers and evolution: helpful offspring and sex allocation.

In some vertebrates, offspring help their parents produce additional offspring. Often individuals of one sex are more likely to become "helpers at the nest". We analyze how such sex-biased offspring helping can influence sex-ratio evolution. It is essential to account for age-structure because the sex ratios of early broods influence how much help is available for later broods; previous authors have not correctly accounted for this fact. When each female produces the same sex ratio in all broods (as assumed in all previous analyses of sex-biased helping), the optimal investment strategy is biased towards the more-helpful sex. When a female has facultative control over the sex ratio in each brood and each helper of a given sex increases the resource available for offspring production by a fixed amount, the optimal strategy is to produce only the more-helpful sex in early broods and only the less-helpful sex in later broods. When there are nonlinear returns from helping, i.e., each helper increases the amount of resource available for reproduction by an amount dependent upon the number of helpers, the optimal strategy is to maximize resource accrual from helping in early broods (which may involve the production of both sexes) and then switch to the exclusive production of the less-helpful sex in later broods. The population sex ratio is biased towards the more-helpful sex regardless of whether the sex ratio is fixed or age-dependent. When fitness returns from helping exhibit environmental patchiness, females are selected to produce only males on some patches and only females on others, and the population sex ratio may be biased in either direction. We discuss our results in light of empirical information on offspring helping, and we show via meta-analysis that there is no support for the claim of Griffin et al. [Griffin, A.S., Sheldon, B.C., West, S.A., 2005. Cooperative breeders adjust offspring sex ratios to produce helpful helpers. Amer. Nat. 166, 628-632] that parents produce more of the helpful sex when that sex is rare or absent.

'Candidatus Streptomyces philanthi', an endosymbiotic streptomycete in the antennae of Philanthus digger wasps.

Symbiotic interactions with bacteria are essential for the survival and reproduction of many insects. The European beewolf (Philanthus triangulum, Hymenoptera, Crabronidae) engages in a highly specific association with bacteria of the genus Streptomyces that appears to protect beewolf offspring against infection by pathogens. Using transmission and scanning electron microscopy, the bacteria were located in the antennal glands of female wasps, where they form dense cell clusters. Using genetic methods, closely related streptomycetes were found in the antennae of 27 Philanthus species (including two subspecies of P. triangulum from distant localities). In contrast, no endosymbionts could be detected in the antennae of other genera within the subfamily Philanthinae (Aphilanthops, Clypeadon and Cerceris). On the basis of morphological, genetic and ecological data, 'Candidatus Streptomyces philanthi' is proposed. 16S rRNA gene sequence data are provided for 28 ecotypes of 'Candidatus Streptomyces philanthi' that reside in different host species and subspecies of the genus Philanthus. Primers for the selective amplification of 'Candidatus Streptomyces philanthi' and an oligonucleotide probe for specific detection by fluorescence in situ hybridization (FISH) are described.

Multiple GnRHs present in a teleost species are encoded by separate genes: analysis of the sbGnRH and cGnRH-II genes from the striped bass, Morone saxatilis.

GnRH is a neuropeptide which plays an essential role in the control of reproductive fitness for all vertebrates. Increasing evidence suggests that multiple forms of GnRH may exist in most vertebrate brains. Southern blot analysis of the three GnRHs known to be present in perciform fish, the seabream (sb)GnRH, the salmon(s) GnRH and the chicken (c) GnRH-II, demonstrates that each is present as a single gene copy in the genome of the striped bass, Morone saxatilis. In order to investigate the physiological consequences of multiple GnRHs in a single vertebrate species, we have isolated and characterized two of the GnRH genes, those for sbGnRH and cGnRH-II. Computer analysis of 3.5 kb of sequence upstream of the sbGnRH gene reveals a number of consensus DNA binding sites which implicate steroids, such as estrogen and glucocorticoids, and the steroidogenic transcription factor, SF-1, as being involved in the regulation of sbGnRH gene expression. Sequence analysis of the cGnRH-II gene reveals evidence of multiple promoters. Expression studies using (1) solution hybridization-RNAse protection mapping with several RNA probes directed at various regions of the proGnRH gene, (2) primer extension assays using two specific oligonucleotide primers, and (3) reverse transcription PCR with several oligonucleotide primers on cGnRH-II transcripts demonstrate that the cGnRH-II gene initiates transcription at numerous sites using a TATA-less promoter within the brains of sexually mature striped bass. This study is the first to characterize and compare the promoter structures of two GnRH genes present in a single vertebrate species.

Purification of gonadotropin II from a teleost fish, the hybrid striped bass, and development of a specific enzyme-linked immunosorbent assay.

A highly purified gonadotropin II (GtH II), referred to as striped bass GtH II (stbGtH II), and its alpha and beta subunits were prepared from pituitaries of sexually mature hybrid striped bass. Pituitary glycoproteins were extracted with ethanol and intact stbGtH II purified by gel-filtration chromatography on Sephadex G-100, ion-exchange chromatography (IEC) on DE-52, and fast-performance liquid chromatography (FPLC) on Superdex 75. The presence of GtHs during the purification procedure was monitored by characteristic elution on reversed-phase high-performance liquid chromatography (rpHPLC) and in vitro steroidogenic activity. The stbGtH II alpha and beta subunits were purified from the pituitary ethanol extract by gel-filtration, IEC, and rpHPLC, and their identities assessed by sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE), rpHPLC, and N-terminal amino acid sequencing. Molecular weights of intact stbGtH II and its alpha and beta subunits, determined by SDS-PAGE, were 34.5, 14.8, and 20.4 kDa, respectively. The stbGtH II beta subunit was used to produce specific antibodies, and a competitive enzyme-linked immunosorbent assay was developed using intact stbGtH II for the standard curve. The sensitivity of the assay was 156 pg/ml (15.6 pg/well) and the intra- and interassay coefficients of variation (at 50% binding) were 7.7% (n = 16) and 8.7% (n = 10), respectively. Physiological validation of the assay was performed by measuring changes of plasma GtH II levels in mature striped bass females, after injection of GnRHa ([d-Ala6,Pro9-NEt]-mGnRH, 100 microg/kg BW). A maximum surge of GtH II in plasma was observed at 12 hr postinjection (22.5 +/- 3. 01 ng/ml), whereas GtH II levels in control fish (around 4 ng/ml) remained unchanged. Displacement curves obtained with serial dilutions of plasma and pituitaries from a number of perciform species were parallel to the standard curve, indicating that this assay can be used for GtH II measurements in a variety of fish species.

Temporal, spatial, sex-ratio and body-size heterogeneity of prey species taken by the beewolf Philanthus sanbornii (Hymenoptera: Sphecidae).

More than 3000 prey representing 108 species of bees and wasps were identified from exoskeletal remains taken from nests of the beewolf Philanthus sanbornii at a site in eastern Massachusetts over a period of five years. Quantitative reference samples totalling more than 4000 items were collected from flowers at the same site over a period of four years. These data give a uniquely detailed view of the way in which a generalist predator exploits a diverse prey community. Most species show striking year-to-year variation in relative abundance, in both the prey and reference collections, but the overall abundances of species tend to be similar in the two collections, as do their sex ratios. This shows: (i) that P. sanbornii takes virtually every bee and wasp species found at flowers during its flight season (except for the relatively small number of species too large to handle); (ii) that prey are taken at rates roughly proportional to their local abundances (with a few exceptions); and (iii) that the local bee and wasp communities have lively dynamics (at least on spatial scales equivalent to the flight ranges of P. sanbornii females). Prey species are non-randomly distributed among nests of individual females within years, and among cells within nests, in a pattern suggesting that females often return repeatedly to hunting sites at which they have had success; the pattern does not suggest that individual females develop preferences for particular prey taxa. The size-abundance distributions of female bees appear to be trimodal at both the individual and species levels, most strongly so when cleptoparasitic species are removed from the sample. Such patterns are seen weakly or not at all in the size-abundance distributions of male bees, male wasps, and female wasps. Bees and wasps of both sexes visit flowers for nectar, but only non-parasitic female bees harvest pollen; this suggests that the multimodality may be caused by aspects of pollen collection that tend to scale with size. The seven Philanthus species of eastern North America vary greatly in size, but they share a common set of relatively small prey species. As in other parts of North America, larger species of Philanthus tend to have relatively broad diets because they also take larger prey that are not available to their smaller congeners. However, long-tongued bees appear to be under-represented in the diets of most North American Philanthus outside the zebratus species group.

Some conceptual issues in the origin of eusociality.

Certain issues arising in connection with the evolutionary origins of eusociality are discussed. Previous results about when natural selection favours helping behaviour are generlised, and the differing viewpoints of both parents and offspring are considered. Particular attention is given to the evolutionary implications of different patterns of overlapping generations observed in bivoltine insects. As argued by Seger (1983), these patterns imply different conditions under which a daughter is selected to help her mother rear additional siblings in haplodiploid populations. Other factors that can alter the selective advantages of helping behaviour under haplodiploidy are also discussed, including the possibility of sex ratio manipulation and the novel result that helping behaviour may be locally favoured in populations that are spatially patchy with respect to sex-specific fitness. A new hypothesis is also presented: The fact that sisters are selected to aid their mother to parasitise other sisters may have played an important role in the origins of eusociality. A given offspring benefits from having maternally parasitised siblings because such siblings rear additional siblings (to which the given offspring is more closely related) instead of nieces and nephews. Finally, the importance of haploidiploidy in the origins of eusociality is discounted; the virtually unique biology of aculeate Hymenoptera would seem to be of much greater importance.

A test-retest reliability study on clinical measurement of masking level differences.

Measurement of binaural release from masking was studied in 30 normal and 10 hearing-impaired subjects to determine the test-retest reliability of masking level differences measured with a clinically feasible technique on generally available clinical equipment. The instrumentation/methodology was found to elicit reliable, consistent, rapid measurements for both 500-Hz and speech (spondaic) stimuli which were comparable with those generally reported in the literature as laboratory-conducted experiments.