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Plasticity in juvenile hormone in male burying beetles during breeding: physiological consequences of the loss of a mate.

Burying beetles, Nicrophorus orbicollis, have facultative biparental care. They bury and prepare small vertebrate carcasses that provide food for their young. Here we establish the juvenile hormone (JH) profiles of paired females, paired males and single males and investigate some of the environmental and social factors that may affect these profiles. Before larvae hatch JH profiles of paired males and females were similar. However, after larvae hatch and during brood care, JH titers of females were very high and those of single males were significantly higher than those of paired males. We tested the hypothesis that higher JH was a response to the need for increased parental care by manipulating brood size. Although JH titers of single males caring for small versus large broods were not significantly different, when comparing JH titers and larval growth (a measure of parental effort), a significant positive correlation emerged. In contrast, we found that food quality had no effect on JH levels suggesting that increased feeding by males and females after carcass discovery cannot explain the elevation of JH. The regulation of JH in male burying beetles appears thus to be dependent on the presence of a mate and on critical stimuli from young.

Adaptation, Physiological↗

Male expulsion in cooperative Mongolian gerbils (Meriones unguiculatus).

Mongolian gerbils are territorial rodents of steppes and semideserts of Mongolia and Northern China. At least under laboratory conditions, the family life is characterized by long-term harmonious and short-term aggressive periods in which some members are expelled. The purpose of this study was to clarify which features of male offspring promote their expulsion. Therefore body mass, absolute and relative weight of testes and epididymis, histology of testes and testosterone concentrations were analyzed in males derived from five families living in semi-natural enclosures with food and water ad libitum. Males expelled (EFM, n=40) had significantly higher relative testes and relative epididymis weight and higher testosterone concentrations in harmonious periods than it did non-attacked ones (IFM, n=13). A stepwise discriminant analysis has established the relative testes weight as decisive for the affiliation to EFM or IFM. Based on this parameter, 71.7% of the offspring could be classified correctly. Moreover, histological results showed that 50.0% of integrated and 29.4% of the expelled family members had incomplete spermiogenesis, indicating a reduced fertility. It is concluded, that males do compete for reproduction within the families which triggers the expulsion of the males with the highest reproductive capacity.

Aggression↗

Men's constructions of smoking in the context of women's tobacco reduction during pregnancy and postpartum.

Men's smoking is largely under-examined despite research that has consistently linked partner smoking to pregnant women's smoking and smoking relapse in the postpartum. An on-going qualitative study involving 31 couples in Canada exploring the influence of couple interactions on women's tobacco reduction provided the opportunity to examine men's smoking in the context of women's tobacco reduction or cessation during pregnancy and postpartum. Individual open-ended interviews with 20 men who smoked were conducted at 0-6 weeks following the birth of their infants and again at 16-24 weeks postpartum. Constant comparative methods were used along with social constructivist perspectives of fatherhood and gender to guide data analysis and enhance theoretical sensitivity. Four themes emerged in men's accounts of their tobacco use: (1) expressing masculinity through smoking, (2) reconciling smoking as a family man, (3) losing the freedom to smoke, and (4) resisting a smoke-less life. Men's reliance on and commitment to dominant ideals of masculinity seemed to preclude them from viewing their partner's tobacco reduction or cessation for pregnancy as an opportunity for cessation. Expectant and new fathers who smoke, however, may be optimally targeted for cessation interventions because it is a time when men experience discomfort with their smoking and when discontinuities in everyday life associated with the transition to fatherhood and presence of a new baby provide opportunities for establishing new routines. Implications for gender-sensitive smoking cessation interventions are discussed.

Adult↗

Testosterone, reproductive stage, and territorial behavior of male and female European stonechats Saxicola torquata.

We investigated territorial behavior and circulating testosterone (T) levels in a multiple-brooded population of the European stonechat, a socially monogamous passerine bird with biparental care. Between arrival at and departure from the breeding territories, we (1) quantified behavior of both sexes in response to a simulated territorial intrusion (STI) of a male conspecific and (2) measured plasma T concentrations in males and females. Male response scores to a STI and male T concentrations varied with phase, but there was no temporal association between plasma T levels and the intensity of territorial behavior. During both two sexual and two parental phases, at least half of the tested males showed aggressive responses. About 20% of the tested males responded with courtship prior to laying of the first clutch, but none of the males courted during later phases. Age had a positive overall effect on male plasma T. Females also reacted to the STI of a male, but their responses did not vary with breeding phase. Female plasma T varied with phase, being elevated during production of the first but not of the second clutch. As with males, female responses to the STI were not correlated with T levels. Responses of pair partners were positively correlated with each other. We conclude that modulation of male territorial aggression with breeding phase is not regulated by changes of plasma T concentrations. In light of other studies showing reduced male aggression by pharmacological inhibition of cellular actions of T, we propose that T is permissive for male territorial aggression, but does not mediate short-term changes associated with breeding phase. The function of the high female plasma T concentrations during formation of first clutches could be related to the production of eggs with high concentrations of androgens.

Age Factors↗

Comparison of the parental behavior of pair-bonded female and male prairie voles (Microtus ochrogaster).

The behavior of primiparous lactating prairie voles (Microtus ochrogaster) and their mates individually interacting with pups was continuously assessed for 45 min after a 2-h parent-litter separation on days 3-4 and 10-11 postpartum. Both sexes were highly parental after reunion with the young, and their general pattern of behavior consisted of bouts of quiescence interspersed with bursts of heightened activity. Lactating females spent more time than males in contact with pups, and more time being quiescent, most often in the kyphotic (upright crouched) nursing posture. Even in the absence of nipples upon which the pups could suckle, males also displayed kyphosis, although for shorter durations than females. Males spent more time, however, huddled over the litter in a hunched position than their mates. In accordance with their decreased quiescence, male voles licked and carried pups more and were more exploratory than females. Compared with the first week postpartum, bouts of kyphosis were shorter during the second week postpartum for both sexes, while laying prone on the pups increased. Males spent less time licking and more time carrying older pups than younger ones, and were more exploratory during the second week postpartum. Sex differences in the parental behavior of prairie voles may reflect differences in the somatosensory stimulation that females and males receive from pups. Furthermore, the display of kyphosis by male voles indicates that the sensorimotor organization of this posture in voles differs from that of lactating rats, which require suckling stimulation for its regulation.

Aging↗

Sexual dimorphism and the NMDA receptor in alloparental behavior in juvenile prairie voles (Microtus ochrogaster).

The prairie vole (Microtus ochrogaster) exhibits parental behavior in both males and females and extensive alloparenting in juveniles. The authors studied the effects on juvenile alloparenting of antagonists for the PCP, glycine, and glutamate sites on the N-methyl-D-aspartate (NMDA) receptor. In male voles, all 3 drugs had an inverted-U dose-response curve. This change could not be attributed to fear of the pup or a nonspecific impairment of cognition, level of locomotor activity, or motor coordination. The PCP site antagonist had a U-shaped dose-response curve in females, the opposite of that in males, but neither of the other drugs changed female alloparental behavior. Both male and female voles exhibit alloparental behavior, but its neurobiological underpinnings are sexually dimorphic in juveniles.

Analysis of Variance↗

Sex differences in the parental behaviour of adult virgin prairie voles: independence from gonadal hormones and vasopressin.

Sexually and parentally experienced prairie voles display robust biparental care of pups that is similar between the sexes. Little is known, however, about possible sex differences in the parental behaviours of sexually inexperienced prairie voles. Parental behaviour of adult virgin male and female prairie voles was examined in sham-operated and gonadectomized subjects treated with vehicle or oestradiol. Since arginine-vasopressin (AVP) has been suggested to stimulate parental behaviour in sexually inexperienced males, neural AVP immunoreactivity (AVP IR) was quantified. Most sham-operated and castrated males displayed high levels of parental behaviour (9/9 controls, 6/9 castrates) during a 15-min exposure to pups 4 weeks after surgery, and few behavioural differences were seen between groups. Conversely, almost all gonadally intact (8/9) and gonadectomized (8/9) females attacked pups. Implantation of a 0.1-mg pellet of oestradiol immediately after gonadectomy had little effect on males (9/9 parental), whereas most (5/9) oestradiol-treated females acted maternally. AVP-immunoreactive (AVP-ir) fibre density in the lateral septum (LS) and lateral habenula (LHb), expressed by the number of pixels that covered AVP-ir fibres during computerized optical density analysis, was greater in males than females, was non-significantly reduced in castrated males, and doubled in the LS of oestradiol-treated females. In a second experiment, males tested 8 weeks after similar manipulations remained highly parental though castrated males had almost no AVP-ir fibres in the LS and LHb. Levels of AVP IR in males treated with oestradiol were similar to those observed in intact males. A dramatic sex difference therefore exists in the parental behaviour of adult sexually naive prairie voles which cannot be explained by sex differences in gonadal hormones. Because both castrated and intact males were highly parental, even though castrates had virtually no AVP-ir in the LS or LHb, AVP does not appear to be crucial for their responsiveness toward pups.

Animals↗

Female brain size and parental care in carnivores.

Comparative studies indicate that species differences in mammalian brain size relate to body size, ecology, and life-history traits. Previous analyses failed to show intrasexual or behavioral patterns of brain size in mammals. Across the terrestrial Carnivora, I find to the contrary. Differences in female, but not male, brain size associate with a fundamental ecological and evolutionary characteristic of female behavior. Other factors equal, females that provide the sole parental care have larger brains than those of biparental or communal species. For females, more parental investment accompanies larger brains. Future comparative studies of mammalian brain size must recognize that some patterns arise independently in the two sexes.

Animals↗

Brighter yellow blue tits make better parents.

Whether or not bird ornaments are a signal for direct (e.g. good parents) or indirect (e.g. good genes) benefits to prospective partners in sexual selection is controversial. Carotene coloration in Parus species is directly related to the ingestion of caterpillars, so that a brightly carotene-coloured tit may be signalling its ability to find caterpillars, a main high-quality food source for good fledgling development, and hence its parental abilities. If carotene-based plumage coloration is related to the good-parent hypothesis, we predict that yellow plumage brightness of tit fathers should be positively correlated to their investment in offspring provisioning. Here, we use cross-fostering experiments in blue tits (Parus caeruleus) to show that chick development (as measured by tarsus length) is related to yellowness of the foster father, but not to that of the genetic parents. Using these data, we were able to measure, for the first time to our knowledge, the separate contribution of genetic and environmental factors (i.e. parental plumage coloration) to chick development, and hence parental investment. Our data, which relate carotenoid coloration to models of good parents, and data from other authors, which relate ultraviolet coloration to good-genes models, stress that different kinds of coloration within an individual may provide different units of information to prospective females.

Animals↗

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.

Age Factors↗

Parentally biased favouritism: why should parents specialize in caring for different offspring?

'Parentally biased favouritism' occurs when the two parents differentially care for individual offspring or kinds of offspring. Examples in birds include brood division and differential investment by the two parents in relation to the size or sex of the offspring. This paper uses mathematical models to investigate which ideas can, in theory, explain parentally biased favouritism. One previous explanation is that the parents differ in their cost of reproduction and that the parent who consequently invests least concentrates its care on the more valuable offspring. However, a mathematical model predicts the total care given by each parent and received by each offspring, not how much each parent cares for each offspring, and hence does not explain parentally biased favouritism. Parentally biased favouritism towards particular types of offspring can be explained by a difference between the parents in the benefits of caring for a given type of offspring or in the effort incurred in providing care to a given type of offspring, but then it is extreme, with at least one of the parents providing care to only one type of offspring. Parentally biased favouritism towards particular individual offspring (brood division) can be explained by parent-offspring conflict or sexual conflict.

Animals↗

Phylogenetic reconstruction of parental-care systems in the ancestors of birds.

Due to the controversy surrounding incipient avian parental care, ancestral parental care systems were reconstructed in a phylogeny including major extant amniote lineages. Using two different resolutions for the basal avian branches, transitions between the states no care, female care, biparental care and male care were inferred for the most basal branches of the tree. Uniparental female care was inferred for the lineage to birds and crocodiles. Using a phylogeny where ratites and tinamous branch off early and an ordered character-state assumption, a transition to biparental care was inferred for the ancestor of birds. This ancestor could be any organism along the lineage leading from the crocodile-bird split up to modern birds, not necessarily the original bird. We discuss the support for alternative avian phylogenies and the homology in parental care between crocodiles and birds. We suggest that the phylogenetic pattern should be used as a starting point for a more detailed analysis of parental care systems in birds and their relatives.

Animals↗

Intrafamilial conflict and parental investment: a synthesis.

We outline and develop current theory on how inherent genetic conflicts of interest between the various family members can affect the flow of parental investment from parents to offspring, and discuss the problems for empirical testing that this generates. The parental investment pattern realized in nature reflects the simultaneous resolution of all the conflicts between the family players. This depends on the genetic mechanism, the mating system and reproductive constraints, on whether extra demand by progeny affects current or future sibs, and particularly on the behavioural mechanisms underlying demand (begging or solicitation) and supply (provision of parental investment by parents). The direction of deviation from the optimal parental investment for the parent(s) depends on the slope of what we term the 'effect of supply on demand', the mechanism that determines how changes in food supply affect begging levels. If increasing food increases begging (positive slope), less parental investment is supplied than the parental optimum and if increasing food decreases begging (negative slope), more parental investment is supplied. The magnitude of deviation depends on both the 'effect of supply on demand' and on the 'effect of demand on supply' (the mechanism determining how changes in begging affect food supply, which always has a positive slope). We conclude that it will often be impossible to deduce the extent of underlying conflict by establishing the amount of parental investment given relative to the ideal optimum for the parent. Some possible directions for future research are discussed.

Animals↗

Heritability of parental effort in a passerine bird.

The study of the evolution of parental care is central to our understanding of social systems, sexual selection, and interindividual conflict, yet we know virtually nothing about the genetic architecture of parental care traits in natural populations. In this paper, we use data from a long term field study of a passerine bird, the long-tailed tit (Aegithalos caudatus), to examine the heritability of the rate at which parents feed offspring. This measure of effort is positively related to offspring survival, is repeatable within individuals, and does not appear to be confounded by environmental effects. Using both parent-offspring regression, and an animal model approach, with a pedigree derived from ringing data, we show that our measure of effort has a significant heritable component.

Animals↗