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Extra-pair paternity and egg dumping in birds: life history, parental care and the risk of retaliation.

Molecular techniques have revealed striking variation among bird species in the rates of extra-pair paternity (EPP) and intraspecific brood parasitism (IBP). In terms of the proportion of broods affected, rates of EPP and IBP vary across species from 0-95% and 0-50%, respectively. Despite a plethora of hypotheses and several careful comparative analyses, few robust correlates of this interspecific variation have been identified. One explanation for this shortfall is that most comparative studies have tended to focus on contemporary ecological factors and ignored fundamental differences in reproductive biology that evolved millions of years ago. We show that, for both EPP and IBP, over 50% of interspecific variation is due to differences among taxonomic families and orders. Therefore, we test hypotheses that predict interspecific variation in the rate of alternative reproductive strategies should be associated with differences in life history and the form of parental care. Our analyses largely support these predictions, with high rates of reproductive cheating being associated with 'fast' life histories. High EPP rates are associated with high rates of adult mortality and reduced paternal care. High IBP rates are associated with high-fecundity rates. These patterns remain intact whether we use species as independent data points or evolutionary contrasts based on either molecular or morphological phylogenies. These results are interpreted as supporting the idea that alternative reproductive strategies are most common in taxa in which the risks of retaliation are low. We suggest a hierarchical explanation for interspecific variation in the incidence of alternative reproductive strategies. Variation between major avian lineages in the EPP and IBP rates are determined by fundamental differences in life history and parental care that evolved many millions of years ago. Variation between populations or individuals of the same species, however, are more likely to be determined by differences in contemporary ecological and genetic factors.

Animals↗

Size breeds success: multiple paternity, multivariate selection and male semelparity in a small marsupial, Antechinus stuartii.

Mating in the marsupial genus Antechinus is a synchronous annual event that is characterized by monoestry in females and abrupt postmating mortality in males. Male semelparity (multiple copulations during a single breeding season per lifetime) is often assumed to occur as a consequence of the intense mating effort expended by males in the rut, but the forces selecting for this remain elusive. Here, we investigate selection in male brown antechinus, Antechinus stuartii, and test two hypotheses for the evolution of semelparity: intermale competition and sperm competition. If intermale competition drives semelparity, we predicted that males would be under strong selection for large body size. If sperm competition is important, we predicted that selection would be strongest on scrotal size, a surrogate for testes volume. Using microsatellite markers, we found that 92% of females in free-living conditions mated with multiple males, producing litters of eight that had up to four fathers. These observations confirm the potential for sperm competition. Using selection analysis, we then found paternity success in 119 males to be related most strongly to body mass and scrotal size, thus providing support for both hypotheses. Large males presumably experience increased paternity success by gaining more matings or prolonged copulations via mate guarding, while large testes may allow increased sperm investment per copulation. Increased levels of free corticosteroid hormones in males facilitate the extreme mating effort during the short period of rut, but lead to immune suppression and consequently to the phenomenon of postmating mortality.

Animals↗

Multiple paternity in a salamander with socially monogamous behaviour.

In the majority of birds and mammals, social monogamy is not congruent with genetic monogamy. No research to date has compared social and genetic monogamy in amphibians. We analysed paternity in clutches of red-backed salamanders (Plethodon cinereus), a species in which social monogamy has been demonstrated in the laboratory, and 28% of individuals in the forest are found in male-female pairs in the noncourtship season. We collected 16 clutches of eggs of P. cinereus in the southern Appalachian Mountains of Virginia and collected tail clippings from attending mothers. We genotyped embryos and adults at five microsatellite loci in order to analyse paternity of clutches. Most clutches (84.6%) had multiple sires, with two to three sires per clutch. In this study, 25% of clutches had males in addition to females attending eggs. None of the mothers of these clutches were genetically monogamous. All attending males sired some of the offspring in the clutch that they attended (between 9% and 50%) but never sired a majority in that clutch. We conclude that, at least in this population, social monogamy in P. cinereus is not concomitant with genetic monogamy.

Animals↗

Combined influence of maternal and paternal quality on sex allocation in red-capped robins.

Sex allocation theory predicts females will adaptively manipulate sex ratios to maximize their progeny's reproductive value. Recently, the generality of biased sex allocation in birds has been questioned by meta-analytic reviews, which demonstrate that many previously reported significant results may simply reflect sampling error. Here, we utilize a robust sample size and powerful statistical approach to determine whether parental quality is correlated with biased sex allocation in red-capped robins. Indices of maternal quality (including interactive effects of age and condition) were strongly related to sex allocation. These relationships were in the predicted directions, with larger effect sizes than those of previous studies in this field. There were also paternal correlates, involving age and the source of paternity. We propose that biased sex allocation occurs in this species, and is maintained by differing production costs of each sex and genetic benefits to females of producing sons when fertilized by high-quality males.

Animals↗

Red cell esterase D in studies of paternity cases in the United Kingdom.

The recently discovered genetic polymorphism of human red cell esterase D has been applied to 156 cases of disputed paternity, along with other well-established systems. The results indicate that esterase D is a useful additional genetic marker for use in paternity testing.

ABO Blood-Group System↗

Paternity cases.

Through analysis of 350 expertises in suits, where the man was accused of paternity, was found that these examinees often lived in a sterile marriage. When examined they often try to influence negatively the result of spermiologic examination. Sterility was ascertained here in only about 10% of cases. In the material consisting of 98 expertises in suits for denial of paternity it was found that about one half of these men were either divorced or in the course of divorce proceedings and about one third of them stated the infidelity of the wife. In more than one third of these cases sterility was proved. The court expert must often express his opinion as to the fertility of the man two or more years previously. His conclusion is especially difficult, where the spermiologic findings repeatedly vary within the limits of severe oligozoospermia with less than 5 million spermatozoids per 1 ml and where the motility is poor.

Adult↗

Effects of paternal cigarette smoking on testicular function, sperm fertilizing capacity, embryonic development, and blastocyst capacity for implantation in rats.

We evaluated the effects of paternal smoking on testicular function, sperm fertilizing capacity, embryonic development, and blastocyst capacity for implantation. Rats of group A were exposed to cigarette smoke for 10 weeks. Rats of group B were exposed to the smoke of incense sticks for 10 weeks. Rats of group C served as a control group. Rats of group D were exposed to cigarette smoke for 7 weeks only. Experimental period was 10 weeks in all groups. At the end of the experimental period serum testosterone responses to human chorionic gonadotropin stimulation, androgen-binding protein activity in testicular cytosols, epididymal sperm motility, and oocyte fertilization rate, oocyte cleavage rate, and blastocyst development rate after in vitro fertilization (IVF) trials were significantly smaller in group A compared with groups B and C. In contrast, fertilization rate, cleavage rate, and blastocyst development rate after intracytoplasmic sperm injection (ICSI) procedures were not significantly different among groups A, B, C, and D. Both after IVF trials and ICSI techniques, the proportion of the alive offspring to the number of transferred oocytes was significantly smaller in group A than in groups B and C. Cigarette smoke-exposure results in a secretory deficiency of Leydig and Sertoli cells leading to an impaired epididymal sperm maturation process and diminished capacity of spermatozoa to penetrate oocytes. In addition paternal cigarette smoke exposure affects the embryonic ability for implantation.

Animals↗

The secondary sex ratio in the United States 1969-71: association with race, parental ages, birth order, paternal education and legitimacy.

The simultaneous effects of several variables on the secondary sex ratio have been examined using data from over 5,000,000 births which occurred in the United States during 1969-71. The previously described negative association of birth order and sex ratio has been confirmed. For legitimate and illegitimate births combined, maternal age and paternal age are unimportant factors once account is made of birth order. This pattern obtains for both blacks and whites. The sex ratio for legitimate and illegitimate births is equivalent but the fact of legitimacy or illegitimacy may affect the association of the ratio with birth order and the parental ages. Paternal education is not significantly related to the ratio, but the highest probability of a male birth is found among fathers with intermediate levels of attainment. Even though the association between sex ratio and order of birth is highly significant in the statistical sense, the proportion of male births changes less than 2% over the extreme values of birth order. Furthermore, it is emphasized that the association accounts for only a very minor (less than 10%) proportion of the total variation in the ratio.

Birth Order↗

Statistical methods for detecting a moderate paternal age effect on incidence of disorder when a maternal one is present.

A statistical method is developed for detecting a moderate effect on the incidence of a disorder caused by advancing age of one parent when it is known that advancing age of the other parent is of great aetiological importance. The method is a conditional test procedure given the parental ages, therefore no assumptions about the parental age distributions have to be made. The method is applied on Danish material on Down's syndrome, for which a paternal age effect is demonstrated. Methods used in some well-known previous investigations have been discussed. Several of them, e.g. the classical one of Penrose (1933), could hardly detect any paternal age effect in Down's syndrome on the available data, because these methods are heavily affected by certain fertility patterns not recognized previously.

Denmark↗

Suicide on the paternal and maternal sides of depressed patients with a lifetime history of attempted suicide.

The computational model of Slater, based on the analysis of ancestral secondary cases on the paternal and maternal sides of the subjects, was applied to depressed patients with a lifetime history of attempted suicide, either violent or nonviolent, in order to investigate possible modes of transmission of suicidal behavior. Among 549 patients, 15 had 2 or more ascendant first- and second-degree relatives who committed suicide. The results of the distribution of these cases were compatible with polygenic inheritance of suicidal behavior in depressed patients with a history of attempted suicide. In patients using violent methods, a significantly greater loading of ancestral secondary cases of suicide was observed on the maternal side and, in the nonviolent attempter group, on the paternal side.

Adult↗

Medical and social paternalism. Regulation of and attitudes towards compulsory psychiatric care.

In Sweden, recommendations and reforms in psychiatric care have increasingly stressed respect for patient autonomy and justice with less emphasis on medical and social paternalism. This is the official policy. But what are the attitudes of the people involved in or affected by compulsory psychiatric care? To answer this question, the attitudes of committed and voluntarily admitted patients, their relatives, psychiatric staff, health and welfare personnel of primary care and a sample of the general public were studied in 2 Swedish counties. Strong support for medical and social paternalism was reported, and according to most of the people asked, doctors, not legal authorities, should decide about commitment. These attitudes are discordant with the recent legislative changes in Sweden.

Adolescent↗

Maternal and paternal age at delivery, birth order, and risk of childhood onset type 1 diabetes: population based cohort study.

OBJECTIVE: To estimate the associations of maternal and paternal age at delivery and of birth order with the risk of childhood onset type 1 diabetes. DESIGN: Cohort study by record linkage of the medical birth registry and the national childhood diabetes registry in Norway. SETTING: Norway. SUBJECTS: All live births in Norway between 1974 and 1998 (1.4 million people) were followed for a maximum of 15 years, contributing 8.2 million person years of observation during 1989-98. 1824 cases of type 1 diabetes diagnosed between 1989 and 1998 were identified. MAIN OUTCOME MEASURES: Incidence of type 1 diabetes. RESULTS: There was no association between maternal age at delivery and type 1 diabetes among firstborn children, but among fourth born children there was a 43.2% increase in incidence of diabetes for each five year increase in maternal age (95% confidence interval 6.4% to 92.6%). Each increase in birth order was associated with a 17.9% reduction in incidence (3.2% to 30.4%) when maternal age was 20-24 years, but the association was weaker when maternal age was 30 years or more. Paternal age was not associated with type 1 diabetes after maternal age was adjusted for. CONCLUSIONS: Intrauterine factors and early life environment may influence the risk of type 1 diabetes. The relation of maternal age and birth order to risk of type 1 diabetes is complex.

Adult↗

Measuring paternal discrepancy and its public health consequences.

Paternal discrepancy (PD) occurs when a child is identified as being biologically fathered by someone other than the man who believes he is the father. This paper examines published evidence on levels of PD and its public health consequences. Rates vary between studies from 0.8% to 30% (median 3.7%, n = 17). Using information from genetic and behavioural studies, the article identifies those who conceive younger, live in deprivation, are in long term relationships (rather than marriages), or in certain cultural groups are at higher risk. Public health consequences of PD being exposed include family break up and violence. However, leaving PD undiagnosed means cases having incorrect information on their genetics and fathers continuing to suspect that children may not be theirs. Increasing paternity testing and use of DNA techniques in clinical and judicial procedures means more cases of PD will be identified. Given developing roles for individual's genetics in decisions made by health services, private services (for example, insurance), and even in personal lifestyle decisions, the dearth of intelligence on how and when PD should be exposed urgently needs addressing.

Child↗

Rational non-interventional paternalism: why doctors ought to make judgments of what is best for their patients.

This paper argues that doctors ought to make all things considered value judgments about what is best for their patients. It illustrates some of the shortcomings of the model of doctor as 'fact-provider'. The 'fact-provider' model fails to take account of the fact that practising medicine necessarily involves making value judgments; that medical practice is a moral practice and requires that doctors reflect on what ought to be done, and that patients can make choices which fail to express their autonomy and which are based on mistaken judgments of value. If doctors are properly to respect patient autonomy and to function as moral agents, they must make evaluations of what their patients ought to do, all things considered. This paper argues for 'rational, non-interventional paternalism'. This is a practice in which doctors form conceptions of what is best for their patients and argue rationally with them. It differs from old-style paternalism in that it is not committed to doing what is best.

Choice Behavior↗

Paternal exposure to agricultural pesticides and cause specific fetal death.

AIMS: To study the association between fetal death and paternal agricultural occupation in areas and time periods with different levels of use of agricultural pesticides. METHODS: A total of 1 473 146 stillbirths and births occurring in Spain between 1995 and 1999 were analysed. RESULTS: The offspring of agricultural workers had the highest risk of fetal death from congenital anomalies in the southern and eastern area (where pesticide use is greatest) and the lowest risk in the rest of Spain. In both areas the offspring of agricultural workers had a similar excess risk of fetal death from the remaining causes of death. The relative risk of fetal death from congenital anomalies in infants conceived between April and September (the months of greater use of pesticides) in the southern and eastern area was 0.90 in manual workers and 1.62 in agricultural workers, compared to non-manual workers; in individuals who were conceived during the rest of the year, the relative risk was 0.87 and 0.85, respectively. In both periods the offspring of agricultural workers had an excess risk of fetal death from the remaining causes of death. CONCLUSIONS: Paternal agricultural work in the areas where pesticides are massively used increases the risk of fetal death from congenital anomalies. The risk is also increased for fetuses conceived during the time periods of maximum use of pesticides The higher risk of fetal death from the remaining causes of death in the offspring of agricultural workers seems unrelated to pesticide exposure.

Abnormalities, Drug-Induced↗

Occupational paternal exposure to benzene and risk of spontaneous abortion.

OBJECTIVE: The purpose of this study was to evaluate the risk of spontaneous abortion among the wives of male workers occupationally exposed to benzene. METHODS: The wives of 823 men working in two chemical plants at the time of the study were asked to complete a questionnaire describing their pregnancies. The analysis of the 1739 pregnancies that ended in a spontaneous abortion or a birth is presented. The firms' payroll records provided all workers' employment history, including dates. Benzene exposure, graded at two levels (< 5, > or = 5 ppm), was determined for every job, so that benzene exposure for each worker's entire professional life (at these companies) could be assessed. This information was linked to the dates of the pregnancies reported in the questionnaires to enable the exposure status of each pregnancy to be defined (1270 non-exposed and 274 exposed). The frequency of spontaneous abortion, defined as the number of spontaneous abortions divided by the total of spontaneous abortions and births was evaluated. RESULTS: When adjusted for tobacco consumption, mother's age and pregnancy order, the odds ratio of the association between paternal exposure to approximately 5 ppm of benzene and the risk of spontaneous abortion was close to and statistically not different from unity (OR = 1.1; 95% CI (0.7-1.8). CONCLUSION: In this study paternal exposure to benzene did not increase the risk of spontaneous abortion.

Abortion, Spontaneous↗

A Prisoner's Dilemma model of the evolution of paternal care.

The heavy energetic demands of gestation, lactation and rearing of offspring mean that studies of paternal care in primates usually focus on female reproductive effort. Here it is shown that both male and female reproductive effort must be considered in order to understand how paternal care evolved. This is done using the Prisoner's Dilemma, best known as a model of reciprocal altruism. It is found that the relative cost of reproduction for males and females is crucially important in determining co-operative and competitive strategies. In particular, when male reproductive costs are less than female reproductive costs, males co-operate with females even when females do not reciprocate. This surprising behaviour, termed non-reciprocal altruism, is comparable with male investment in a female and her offspring.

Altruism↗

Human red cell galactose-1-phosphate uridylyltransferase (EC 2.7.7.12). Electrophoretically determined polymorphism in Denmark and its use in paternity cases.

The electrophoretically detectable phenotypes of human red cell galactose-1-phosphate uridylyltransferase (GALT) were determined in 2,074 unrelated Danes. The gene frequencies were: GALT1 = 0.9233 and GALT2 = 0.0767. The segregation of phenotypes in 765 mother-child pairs was consistent with autosomal codominant inheritance. One apparent mother-child incompatibility with respect to phenotypes was observed which, however, appeared to be due to the segregation of a silent gene. The results of an investigation of 248 paternity cases are reported, and the application of the GALT polymorphism to paternity cases is discussed.

Child↗