Sex roles and sexual selection

Sexual selection has been portrayed as acting predominantly on males who compete with each other over copulatory access to females; selection was considered to be driven by females choosing between males at the pre- or postcopulatory level. However, a broader view of sexual selection is now emerging. Examining male discrimination between females and female-female competition has been beneficial in identifying factors influencing the direction and strength of sexual selection. Furthermore, consideration of processes such as sexual coercion or genetic incompatibility, which indirectly influence an individual's set of copulation partners, gamete set or their offspring success, has helped to clarify the ways in which sexual selection may operate. Moreover, there is increasing evidence that not all copulations translate directly to paternity and that paternity does not necessarily translate into successful offspring. Postcopulatory and postfertilization mechanisms that influence not only paternity share but offspring recruitment now require further consideration. The benefits to each sex of copulating with particular partners or with more than one partner remains an area of debate. More carefully designed studies which eliminate alternative possibilities or quantify the relative importance of different selective pressures will also benefit from considering that not all copulations function solely to inseminate or receive sperm. It is also now clear that not all individuals of one sex follow the same strategy. Examining the variation between individuals in reproductive behaviour, fertilization success and offspring success will be important in establishing the selective pressures and mechanisms underlying the operation of sexual selection. (c) 1998 The Association for the Study of Animal Behaviour.     

A developmental defect in Plasmodium falciparum male gametogenesis

Asexually replicating populations of Plasmodium parasites, including those from cloned lines, generate both male and female gametes to complete the malaria life cycle through the mosquito. The generation of these sexual forms begins with the induction of gametocytes from haploid asexual stage parasites in the blood of the vertebrate host. The molecular processes that govern the differentiation and development of the sexual forms are largely unknown. Here we describe a defect that affects the development of competent male gametocytes from a mutant clone of P. falciparum (Dd2). Comparison of the Dd2 clone to the predecessor clone from which it was derived (W2'82) shows that the defect is a mutation that arose during the long-term cultivation of asexual stages in vitro. Light and electron microscopic images, and indirect immunofluorescence assays with male-specific anti-alpha-tubulin II antibodies, indicate a global disruption of male development at the gametocyte level with at least a 70-90% reduction in the proportion of mature male gametocytes by the Dd2 clone relative to W2'82. A high prevalence of abnormal gametocyte forms, frequently containing multiple and unusually large vacuoles, is associated with the defect. The reduced production of mature male gametocytes may reflect a problem in processes that commit a gametocyte to male development or a progressive attrition of viable male gametocytes during maturation. The defect is genetically linked to an almost complete absence of male gamete production and of infectivity to mosquitoes. This is the first sex-specific developmental mutation identified and characterized in Plasmodium.     

Increased competitiveness of nematode sperm bearing the male X chromosome

Male offspring, which cannot reproduce independently, represent a cost of sexual reproduction. This cost is eliminated by the production of hermaphroditic offspring in the self-fertilizing nematode Caenorhabditis briggsae. However, these hermaphrodites can outcross by mating with males. Half the sperm received from males contain no sex chromosome and therefore give rise to male progeny. Mating with males should thus impose the cost of making male offspring. We found that male sperm took immediate precedence over hermaphrodite sperm, resulting in maximized outcrossing, but the appearance of male progeny was delayed after mating. This delay is caused by the male X-bearing sperm outcompeting their nullo-X counterparts. The competitive advantage of X-bearing sperm over nullo-X sperm is limited to sperm from males; it did not occur in a mutant hermaphrodite that produces both types of sperm. The chromosomal effect on sperm competitiveness in C. briggsae, which has not been observed in other species, suggests that the X chromosome has evolved a form of meiotic drive, selfishly increasing the competitiveness of sperm that bear it over those that do not. Thus, the multiple levels of sperm competitiveness found in C. briggsae maximize outcrossing after mating while delaying the cost of making male offspring.     

The cytology of the gametes and fertilization of Allomyces macrogynus

The gametes and the process of fertilization were examined by light and electron microscopy in the lower eukaryote Allomyces macrogynus. Differences in gamete morphology included the overall larger size and the presence of a larger nuclear apparatus, along with the association of a side-body complex and many more mitochondria in the female gamete. In this species of Allomyces, fertilization was initiated by contact and fusion of specialized regions of the gamete plasma membranes resulting in a binucleate fusion cell surrounded by plasma membrane contributed by both partners. Following plasmogamy, nuclear fusion was initiated by multiple nuclear membrane contacts between adjacent outer membranes. Following inner membrane fusion, small nucleoplasmic bridges were observed which presumably fused with one another and resulted in a single bridge which widened, forming the mature diploid nucleus. After karyogamy, fusion of the nuclear caps did not always occur and zygotes with and without fused caps were observed. Coalescence of the nucleoli completed the events of fertilization, forming zygote with a single nuclear apparatus (sometimes with two caps) and two flagella. These observations are discussed in relation to fertilization mechanisms and compared to fertilization in other organisms.     

The effect of sex on adaptation to high temperature in heterozygous and homozygous yeast

Most explanations for the evolutionary maintenance of sex depend on the assumption that sex produces variation by recombining parental haplotypes in the offspring. Therefore, meiosis is expected to be useful only in heterozygotes. We tested this assumption by competing sexual strains of yeast against constitutive asexuals in a hot (37 degrees C) culture for 500 generations, in either heterozygous or homozygous genetic backgrounds. We found that there was an initial cost of sex for all the sexual strains, which was indicated by a sharp increase in the proportion of asexuals after the induction of sex. The cost was larger in the heterozygotes than in the homozygotes, probably because of recombinational load. However, in two of the three heterozygote backgrounds, after the initial success of the asexuals, the remaining sexuals eventually drove them out of the population. These two heterozygotes also suffered the largest initial cost of sex. In the other heterozygote and in the three homozygote backgrounds it appeared to be a matter of chance whether sexuals or asexuals won. The average relative fitness increased in all the strains, but the increase was largest in the two strains that showed both the clearest advantage and the largest cost of sex. We conclude that these results are consistent with the traditional view that sex has a short-term cost but a long-term benefit.     

Membrane differentiations at sites specialized for cell fusion

Fusion of plasma membranes between Chlamydomonas reinhardtii gametes has been studied by freeze-fracture electron microscopy of unfixed cells. The putative site of cell fusion developes during gametic differentiation and is recognized in thin sections of unmated gametes as a plaque of dense material subjacent to a sector of the anterior plasma membrane (Goodenough, U.W., and R.L. Weiss. 1975.J. Cell Biol. 67:623-637). The overlying membrane proves to be readily recognized in replicas of unmated gametes as a circular region roughly 500 nm in diameter which is relatively free of "regular" plasma membrane particles on both the P and E fracture faces. The morphology of this region is different for mating-type plus (mt+) and mt- gametes: the few particles present in the center of the mt+ region are distributed asymmetrically and restricted to the P face, while the few particles present in the center of the mt- region are distributed symmetrically in the E face. Each gamete type can be activated for cell fusion by presenting to it isolated flagella of opposite mt. The activated mt+ gamete generates large expanses of particle-cleared membrane as it forms a long fertilization tubule from the mating structure region. In the activated mt- gamete, the E face of the mating structure region is transformed into a central dome of densely clustered particles surrounded by a particle-cleared zone. When mt+ and mt- gametes are mixed together, flagellar agglutination triggeeeds to fuse with an activated mt- region. The fusion lip is seen to develop within the particle-dense central dome. We conclude that these mt- particles play an active role in membrane fusion.     

A direct role for arrestins in desensitization of the luteinizing hormone/choriogonadotropin receptor in porcine ovarian follicular membranes

A heterologous bovine in vitro fertilization (IVF) system was used to study the functional competence of scimitar-horned oryx spermatozoa after cryopreservation. Four sperm-freezing methods were compared after dilution of ejaculates from six oryx with an equine semen extender: 1) dry ice, 2) dry shipper one-step, 3) dry shipper two-step, and 4) liquid nitrogen vapor. Post-thaw sperm motility, longevity, and acrosomal status were assessed and zona pellucida penetration, fertilization, and embryo cleavage were evaluated after coincubation of thawed oryx spermatozoa with in vitro-matured domestic cow oocytes. Sperm motility index (SMI) decreased (p < 0.05) over a 6-h period, but a high percentage (>/= 65%) of spermatozoa contained intact acrosomes in all treatments. Despite differences in sperm motility among methods, oocyte penetration, fertilization, and embryo cleavage did not differ (p >/= 0.05). However, cleavage success was < 50% across all treatments. There were positive correlations (p < 0.05; r = 0.81-0.97) between sample SMI at 3 and 6 h and fertilization, penetration, and cleavage, but no correlations (p >/= 0.05) between SMI at 0 or 1 h and IVF success. This study demonstrates that compatible heterologous gamete interaction allows thorough assessment of post-thaw sperm function in an endangered antelope. Scimitar-horned oryx spermatozoa appear relatively tolerant of varied cryopreservation methods, and preserved samples exhibit adequate post-thaw function to warrant use for assisted reproduction.     

Why do asthmatic children need referral to an emergency room?

This review assesses whether human follicular fluid (hFF) is able to modify human sperm function in vitro. Addition of hFF has been found to stimulate the motility of washed human spermatozoa, to increase the percentage of hyperactivated spermatozoa, to induce the acrosome reaction, to attract spermatozoa to the site of fertilization and to facilitate penetration of the oocyte by spermatozoa. It is possible that hFF could provide a favourable environment around the oocyte for fertilization by spermatozoa. Inclusion of hFF in gamete transfer medium may also improve the success rate of assisted reproduction technology. Purification of individual components in hFF which modify different aspects of sperm function awaits further investigation.     

Differentiating between parthenogenetic and "positive development" embryos in turkeys by molecular sexing

In mated or inseminated turkeys, 5 to 15% of eggs set for incubation show only rudimentary development. Most of these embryos die during the first 24 to 48 h of incubation and contain only unorganized sheets of tissue. This abnormal development is termed "positive development" (PD). Turkey eggs also show incidence of parthenogenesis and the resulting progeny is believed to be always male. As both types of embryos are morphologically similar at the early stage of incubation, it has been speculated that PD embryos may in fact be parthenogens. By identifying the sex at the blastodermal stage with the help of DNA markers, we have differentiated between the PD embryos and parthenogens. Parthenogenetic embryos were obtained from eggs laid by uninseminated or virgin Beltsville Small White (BSW) hens, and the PD embryos were obtained from eggs of inseminated Nicholas and British United Turkeys of America (BUTA) hens. DNA was extracted from blastoderms of parthenogenetic and PD embryos. Turkey W-chromosome specific DNA probe and primers were used to detect females in all samples by Southern blot and polymerase chain reaction (PCR), respectively. No female was detected among the 35 parthenogens examined, whereas there were 3 females among the 11 PD embryos. The presence of both males and females among PD embryos suggests that they are products of fertilization, and that at least these 3 female embryos, if not all the 11 PD embryos, are not of parthenogenetic origin. It is concluded, therefore, that PD embryos result from errors in fertilization or from early embryonic mortality following successful fertilization, and that they are unlikely to be of parthenogenetic origin.     

Effects of sex guilt and sexual stimulation on the recall of word associations.

Elicited free associations to a mixed list of double-entendre sexual and asexual words from 84 male undergraduates under conditions of sexual and no sexual stimulation (control). Ability to recall the associations was then studied in relation to individual differences in sex guilt. Recall was tested immediately after associations were elicited. Results indicate a significant interaction between stimulation and guilt. High-sex-guilt Ss showed better recall for associations elicited under the control condition. Conversely, low-sex-guilt Ss showed better recall under the sexual stimulation condition. Results also suggest that the sexuality-asexuality of stimulus words might be a significant factor in the recall errors of high- and low-sex-guilt Ss. (18 ref.) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Manipulation of the fertility of marsupials for conservation of endangered species and control of over-abundant populations

Marsupials present a dichotomy in population management; the numbers of many Australian marsupial species have declined due to loss of habitat, competition from introduced herbivores and predation by introduced carnivores, but other species have become locally overabundant in Australia or are introduced pests in New Zealand. The manipulation of reproduction offers the means to increase or decrease productivity; however, considerable fundamental research is required before reproductive technologies can be applied to marsupials. Marsupials differ from eutherian mammals in several aspects of their reproduction including sex differentiation, gamete function and endocrinology, as well as in the relative lengths of gestation and lactation. Although these differences present unique problems in the application of reproductive technologies to marsupials, they also present unique opportunities for marsupial-specific fertility control. This paper summarises the assisted breeding technologies currently being applied to marsupials including superovulation, artificial insemination, in vitro fertilization and gene banking; unique marsupial targets for contraceptive intervention including gamete production, sperm capacitation, gamete surface antigens and embryonic development; and some options for the delivery of contraceptive vaccines to marsupial populations.     

Glucose participates in sperm-oocyte fusion in the mouse

Glucose metabolism is necessary for successful fertilization in the mouse, but the precise step of this process that requires glucose has not been determined. In this study we have investigated the involvement of glucose in the gamete binding and fusion process. The relationship between the metabolism of this carbohydrate in the oocyte and this particular step of fertilization has also been studied. A binding and fusion assay was performed by inseminating zona-free oocytes with capacitated sperm. Sperm binding and the fusion reaction were assessed by counting the sperm bound to the oolemma and the sperm heads decondensed in the ooplasm, respectively. To determine the importance of glucose metabolism, nonmetabolized glucose analogues (L-glucose, 2-deoxyglucose, 3-O-methylglucose) were used instead of glucose. The binding/fusion assay was also performed in the presence of glucose uptake inhibitors (cytochalasin B, phloretin, phlorizin). We have obtained evidence that glucose was specifically required for sperm penetration into zona-free oocytes (gamete fusion) but not for sperm binding to the oolemma. The metabolism of this carbohydrate appears to be essential, since nonmetabolized glucose analogues were unable to support this process. Although glucose metabolism was detectable in the oocyte, it did not appear to be related to the gamete fusion function. We hypothesize that the male gamete, which utilizes glucose as an energy substrate for various functions, may have a specific need to metabolize this hexose during fusion and incorporation within the oocyte.     

Sex differences in learned helplessness: IV. An experimental and naturalistic study of failure generalization and its mediators.

Tested the hypothesis that attributions for failure can mediate the generalization of failure effects across situations: When perceived causal factors remain present in otherwise novel situations, failure effects should transfer; when perceived causal factors are removed, failure effects should be attenuated. Specifically, it was predicted that sex differences in attributions would result in differential transfer to novel situations, with boys showing greater recovery of success expectancies when the evaluator changes, but girls showing greater recovery of success when the ability areas change. Two studies are reported: one a field study (40 female and 40 male 5th graders) examining changes in expectancy of academic success over the school year, and the other a laboratory analog (171 female and 143 male 4th–6th graders) examining directly the effects of evaluator and task change. Results provide strong support for the hypothesis and suggest an explanation for sex differences in long-term academic achievement. (14 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Individual and situational markers of condom use and sex with nonprimary partners among alcoholic inpatients: Findings from the ATRISK Study.

Alcoholics in treatment have demonstrated increased rates of HIV risk behavior and infection. This study explored individual and situational variables associated with HIV risk behaviors such as condom nonuse and sex with nonprimary partners reported during structured interviews of 802 male and female alcoholic inpatients. Logistic regression analyses indicated that person variables, sexual history variables, and situation variables were independently associated with sexual risk behavior in the most recent episode. Results of paired t tests revealed that participants who had engaged in sex with both primary and nonprimary partners during the previous 6 months reported significantly more frequent alcohol and condom use in situations involving sex with nonprimary partners. Prevention efforts need to target alcoholic inpatients as a group and, within this group, to focus on both high-risk individuals and high-risk situations. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Reflections on the beginning of a new human individual

Although there is very little doubt that when a child is born a new actual person can be identified, there is continuous debate as to the moment in embryological development when that same person begins its existence. Based on today's knowledge of human fertilization and the early stages of embryo development, this position paper examines three theses that deal with the establishment of personhood. The first thesis stipulates that a human individual exists prior to syngamy. Although the sperm has penetrated the plasmatic membrane of the oocyte, the genetic information contained in both gametes remain separated in the male and female pronucleus; thus, the oocyte contains the sum of two identities responsible for creating a new individual. This paper will argue that a human individual has not yet formed. The second thesis recognizes that with syngamy, a unicellular structure (zygote) is established, endowed with genetic individuality and with the potential to become a person maintaining that same genetic framework throughout its lifetime. The third thesis argues that although genetic individuality is established with syngamy, the ontological individuality is only reached once genetic expression and cellular specialization are achieved and twinning is no longer possible (15 days after fertilization).     

New technology and new challenges for assisted reproduction

As assisted reproduction technology advances, more types of procedures are becoming available, bringing more success at solving many types of infertility. In vitro fertilization has become simpler and less invasive, with success rates as high as 30% per cycle. Intracytoplasmic sperm injection has solved many types of male infertility. This article explains in vitro fertilization technology and discusses such ethical issues as embryo ownership, multiple births, and embryo genetic testing.     

Mammalian sex determination: from gonads to brain

In mammals, sex is determined by the Y chromosome, which encodes a testis-determining factor (TDF). This factor causes the undifferentiated embryonic gonads to develop as testes rather than ovaries. The testes subsequently produce the male sex hormones that are responsible for all male sexual characteristics. In 1990, the sex-determining gene, TDF, was identified and termed SRY in humans (Sry in mice). It encodes a protein containing a high mobility group (HMG) motif, which confers the ability to bind and to bend DNA. Genetic evidence supporting SRY as TDF came from the observation of a male phenotype in XX mice transgenic for a small genomic fragment containing Sry, and from the study of XY sex-reversed individuals who harbor de novo mutations in the SRY coding sequence. Other non-Y-linked genes involved in sex determination were subsequently found by genetic analysis of XY sex-reversed patients not explained by mutations in SRY. These genes are WT1, SF1, DAX1, and SOX9. A regulatory cascade hypothesis for mammalian sex determination, proposing that SRY represses a negative regulator of male development, was recently supported by observation of mice that expressed a DAX1 transgene and developed as XY sex-reversed females. The role of some sex-determining genes, such as DAX1 and SF1, in the development of the entire reproductive axis, a functionally integrated endocrine axis, leads to a new concept. Normal sexual development may result from the functional and developmental integration of a number of different genes that play roles in sex determination, sexual differentiation, and sexual behavior.     

The role of the paternal genome in the development of the mouse germ line

The mouse germ line originates at 6.5 days post coitum (dpc) in the proximal epiblast, apparently in response to signals from the primitive endoderm or the extraembryonic mesoderm [1,2]. Some studies have implied a significant role for imprinted genes in germ-line development [3,4]. These genes, whose expression is determined by their parental origin [5], serve complementary functions during mammalian development [6-9] and exert striking reciprocal phenotypic effects on androgenetic (AG: two paternal genomes) and parthenogenetic (GG/PG: two maternal genomes) cells [3,4,10]. This may include a fundamental effect on germ-cell development because PG but not AG cells can differentiate into viable gametes [3,4,11], suggesting that the maternal genome is obligatory for development of the mammalian germ line. Here we show unequivocally that AG cells can differentiate into germ cells, and that in chimeras with normal cells they produce functional sperm. These studies establish that the paternal and maternal genomes can individually provide both the signal and the response required for the specification of germ cells in mammals.     

Rapid evolution of sex-related genes in Chlamydomonas

Biological speciation ultimately results in prezygotic isolation-the inability of incipient species to mate with one another-but little is understood about the selection pressures and genetic changes that generate this outcome. The genus Chlamydomonas comprises numerous species of unicellular green algae, including numerous geographic isolates of the species C. reinhardtii. This diverse collection has allowed us to analyze the evolution of two sex-related genes: the mid gene of C. reinhardtii, which determines whether a gamete is mating-type plus or minus, and the fus1 gene, which dictates a cell surface glycoprotein utilized by C. reinhardtii plus gametes to recognize minus gametes. Low stringency Southern analyses failed to detect any fus1 homologs in other Chlamydomonas species and detected only one mid homolog, documenting that both genes have diverged extensively during the evolution of the lineage. The one mid homolog was found in C. incerta, the species in culture that is most closely related to C. reinhardtii. Its mid gene carries numerous nonsynonymous and synonymous codon changes compared with the C. reinhardtii mid gene. In contrast, very high sequence conservation of both the mid and fus1 sequences is found in natural isolates of C. reinhardtii, indicating that the genes are not free to drift within a species but do diverge dramatically between species. Striking divergence of sex determination and mate recognition genes also has been encountered in a number of other eukaryotic phyla, suggesting that unique, and as yet unidentified, selection pressures act on these classes of genes during the speciation process.     

Horizontal transmission of parthenogenesis-inducing microbes in Trichogramma wasps

Complete parthenogenesis (thelytoky) in species of the parasitic wasp Trichogramma is usually caused by the cytoplasmically inherited bacterium Wolbachia. This symbiont induces gamete duplication, which, in these haplodiploid organisms, results in all-female broods. Antibiotic treatment "cures' this condition, restoring normal sexual reproduction. Phylogenetic analysis of Wolbachia has shown that, in contrast with the strains in other host organisms (where the symbiont also induces different reproductive alterations), those in Trichogramma form a monophyletic group. This might be an indication of symbiont-host cocladogenesis. To test this, we performed comparative molecular phylogenetics on 20 parthenogenetic Trichogramma cultures and their Wolbachiae. We conclude that there is, in fact, little evidence for cocladogenesis. Instead, the phylogenetic distribution of the symbionts appears to result from occasional horizontal transmission, which probably takes place inside the hosts of Trichogramma parasitoids (usually lepidopteran eggs). This study therefore suggests that parthenogenesis is not only curable, it can sometimes be contagious also.