geher 2007 parental investment in primates

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Geher 2007 parental investment in primates nesk investment maroc15

Geher 2007 parental investment in primates

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General Life History. Possible Appearance:. Probable Appearance:. Definite Appearance:. The Human Difference:. Universality in Human Populations:. References Caring for infants is associated with increased reproductive success for male mountain gorillas , Rosenbaum, Stacy, Vigilant Linda, Kuzawa Christopher W. Extraordinary intelligence and the care of infants. Comparative population genomics in animals uncovers the determinants of genetic diversity. Parental investment: The hominid adaptation , Lancaster, Jane B.

Age at First Reproduction. Awareness of Past and Future. Beliefs About Death. Brain Size. Care of the Infirm and Elderly. Control of Paternity. Cooperative Breeding. Cultural Transmission. Duration of Infant Arousal. Food Sharing.

Group Stability. Helplessness of the Young. Infant Locomotor Development. Inter-birth Intervals. Moral Sense. Paternal Care. Personal Names. Provisioning is mating effort if females prefer to mate with successful hunters. Provisioning can also be viewed as parental effort if offspring that receive it have better survival prospects than those who do not e.

One important benefit of this paternal care is that inter-birth intervals are shorter in humans than in comparable-sized primates with only maternal care Gangestad, This quicker return to fertility after birth may be due to hunting efforts that brought more high quality food to the family or, as Gettler suggests, it may in part be due to reduction of maternal energy expenditure in mobile groups if fathers carried offspring.

Gettler suggests that we have overstated the sexual division of foraging labor in ancestral groups based on modern hunter gathering populations and he cites evidence that infant transport by fathers, as in some other primate species, was important for reducing maternal energy expenditure in mobile foraging groups. One way we might determine whether human paternal care is adaptive is by examining the underlying hormonal mechanisms.

Gangestad argues that the association of paternal care with decreased testosterone as opposed to increased testosterone with mating effort suggests that human paternal care is an adaptation. Such an argument could also be made for the more recent research on oxytocin and vasopressin. While male and female primates have gonadal differences derived from their genetic sex, there are a number of anatomical similarities that suggest males could respond to infants, both behaviorally and hormonally, as do females.

For example, male and female primates show nipple and duct development with no gross structural pre-pubertal sex differences Daly, Only with the onset of puberty do females show changes in mammary development due to ovarian and adrenal steroids. These changes are not seen in males, although in monomorphic species such as the marmosets and tamarins, both sexes lack external differences in mammary development. Interestingly, blocking fetal androgen action in genetic males, can lead to full lactation in adult male rats with orchidectomy and stimulation from pregnancy hormones Neumann and von Berswordt-Wallrabe, Thus, males are very responsive to their hormonal milieu and this in turn influences their behavior and physiology.

Primates demonstrate a long interval between conception and the stage where males can care for the young. Close association with the pregnant female may enable males to process the signals that promote the biological changes of fatherhood. A number of studies on expectant tamarin and marmoset fathers have revealed extensive physiological changes to the males prior to the birth of their infants.

Males are therefore responsive to the reproductive state of their mates Ziegler et al. Ziegler et al. Similarly, elevated estradiol, prolactin and testosterone concentrations have also been demonstrated for common marmoset males, again with concentrations peaking in the last month of gestation Zeigler et al. Storey et al. More recently computerized dolls RealCare Baby have been used to systematically assess paternal care van Anders et al. Longitudinal studies have demonstrated hormonal changes in individual men when they become fathers Gettler et al.

Further, hormonal changes are greatest in the men who provide the most paternal care Alvergne et al. Neuropeptides, primarily oxytocin and vasopressin, are now measured in plasma, saliva and urine and these hormones have been linked to responsiveness to mates and young. Intranasal sprays of these neuropeptides provide an experimental means to assess hormone-behavior interactions in humans e.

Recent studies show that variation in neuropeptide receptor alleles is reflected in variation in parental behavior Feldman et al. Table 1 contains a comparative summary table of the hormonal mechanisms of paternal behavior in primates as is currently known. Summary of relationships between hormone levels and paternal behavior in primates at different stages e. Male primates need to have flexible hormonal responses since behaviors requiring high testosterone, such as mate guarding or territorial defense behaviors, co-occur with low-testosterone paternal care Ziegler et al.

This flexibility is most evident in species where infant care behaviors co-occur with postpartum ovulation. However, mating behavior does not alter the high rates of father—infant interactions. Male marmosets that are not living with young show a significant increase in testosterone concentrations when smelling a novel periovulatory scent while fathers with young show little response to these odors Ziegler et al.

Testosterone concentrations are low while males are caring for their offspring Nunes et al. Additionally, testosterone concentrations in black tufted-ear marmosets are lower for males that have more paternal experience and carry young more frequently than for other males Nunes et al.

Marmoset infant odors may work as signals that promote recognition of offspring and as primer by affecting paternal hormones. Experienced marmoset fathers show a significant decrease in serum testosterone concentrations within 20 minutes of contact with an isolated scent from their own infant Prudom et al. In contrast, parentally inexperienced males show no changes in testosterone concentrations in response to unfamiliar infant odors.

Fathers also showed an increase in estrogen in response to odors from their own dependent infants Ziegler et al. Of all our results, this finding is associated with the most subsequent research. Generally in North American studies, testosterone concentrations are lower in pair-bonded men and fathers than in single, childless men Gray and Campbell, No differences have been found in the testosterone concentrations of fathers and non-fathers in some of the non-North American studies reviewed in Gray and Campbell, Testosterone concentrations were lower in fathers than non-fathers in the Hazda who are highly paternal but not different in the nearby Dagota who spent little time with young children Muller et al.

Gettler et al. This decrease in testosterone was greater in men whose children were younger when they were sampled, in men who spent more time caring for their children Gettler et al. Similarly, Alvergne et al. New fathers had variable, but on average, increased testosterone concentrations after listening to taped infant cries and holding their newborn for 30 minutes Storey et al. These findings appear to contradict the general findings that lower testosterone concentrations are associated with greater paternal responsiveness.

Van Anders et al. When men could not comfort the infant surrogates, computer control bracelet was not provided their testosterone concentrations did not change. None of the men in the comfort group showed an increase in testosterone whereas at least half of the men in the other two groups showed such an increase. Similarly, Weisman et al. Taken together, this literature suggests that it is not just being a father that affects testosterone concentrations but rather the amount and effectiveness of the paternal care that men provide.

Alternatively, these increases may also be involved in helping men counter any external threats to their infants i. Prolactin is known for its role in promoting maternal behaviors in mammals Bridges et al. In the rat, prolactin regulates the onset of maternal behavior but not the maintenance, which is controlled by pup exposure Rosenblatt and Siegel, The role of prolactin in mammalian paternal behavior is more subtle and possibly more species-specific than for some other hormones.

Prolactin has been suggested as being involved in the transition from a non-paternal to a paternal state in male mammals e. Males from a number of biparental mammalian species show elevated prolactin while they are actively participating in infant care. These species include the Mongolian gerbil, California mouse and Siberian dwarf hamsters Brown et al. The naturally biparental Siberian dwarf hamster, Phodopus campbelli, shows higher mRNA for prolactin receptors in the choroid plexus of the brain than non-paternal males of Phodopus sungorus Ma et al.

Prolactin suppression, however, has variable effects in animal studies: infant care behavior was lower in some studies mixed sex group, Roberts et al. Differences in behavioral testing conditions, specifically testing in small cages, may explain why some studies find no deficits because the constant close proximity between fathers and young may over-ride the effects of hormonal suppression. Support for this possibility comes from a study of prolactin suppression in female voles: no effects on maternal behavior were seen for females tested in small cages.

In contrast, when families were housed in larger enclosures, prolactin-suppressed females stayed away longer from their pups than controls, though once they returned to the nest, their bout times were equivalent to controls Walsh et al. Similarly, Ziegler et al. These results suggest prolactin concentrations and suppression should be evaluated in more naturalistic conditions. In one study to do this, Carlson et al. We have provided details on these studies to set the context for the further discussion in primates.

Researchers have suggested that prolactin has both a preparatory role and short-term response role in primate paternal behavior. Results have differed as to when and how much prolactin changes, with some differences being related to whether males have previous paternal experience and whether they are living with young during the experiment. This prenatal increase was not seen, however, when the experienced males were not living with offspring Almond et al.

Prenatal prolactin concentrations were higher in experienced fathers than non-fathers in two species where males begin carrying infants right after birth but in this study, there was no post-natal increase Schradin et al. Mota et al. Taken together, these studies suggest that prolactin increases at or before the time males start carrying infants and that this preparatory phase may be affected by previous experience or whether males are living in family units with young.

Such a possibility is consistent with the meerkat study Carlson et al. In an attempt to determine a causal relationship of prolactin to paternal care behaviors, Ziegler et al. Each male went through each treatment. There was little effect on paternal behavior with elevated prolactin in the experienced fathers, except as noted above. Elevated prolactin postpartum, when males are actively caring for infants, may work to prevent excessive weight loss during their added energetic demands Ziegler et al.

Males lose a significant amount of weight while they are caring for infants. Prolactin implants prevented males from losing weight during the first three weeks after infant birth while lowering prolactin with cabergoline increased the amount of weight males lost. It would be interesting to conduct such a study on a natural population as it is likely that males with significant weight loss would show a decrease in paternal care. While all marmosets and tamarins gain experience with infants through caring for their younger siblings, as they are cooperative breeders, it is the experience of being a father that provides for the major physical, endocrine and behavioral changes that appear to be important in ensuring the successful development of the infant.

As in male tamarins and marmosets, human fathers also have been reported to exhibit increased prolactin concentrations before the birth Storey et al. Fathers with higher prolactin concentrations were more responsive to infant cries Fleming et al.

Prolactin concentrations are higher in fathers than non-fathers in a sample of highly paternal men in the Philippines Gettler et al. Experienced fathers of newborns show a significantly greater increase in prolactin after hearing infant cries or holding their babies than first-time fathers Fleming et al. In contrast, women showed a significant prolactin increase in response to baby cries and a birth video even before their first babies were born Delahunty et al.

Thus, there appears to be a sex difference such that, for males, learning and exposure to social cues may be more important in the development of prolactin responsiveness than for females. Prolactin reactivity short-term changes following a social interaction in human fathers is also related to the interaction patterns in father-infant contact. Two studies found that prolactin concentrations significantly declined after fathers interacted for min with their toddlers Gettler et al.

Greater prolactin declines were associated with greater paternal responsiveness: prolactin declines were greater in men who engaged in more childcare and whose wives judged them to be better fathers Gettler et al. As with the animal studies, these results suggest that prolactin is involved in the timing and possibly the motivation to initiate bouts of parental care, but it may be less involved in the actual bouts.

It would be better to test men before they re-connect with their infants and examine whether there is relationship between pre-contact anticipation? An earlier general view of glucocorticoids is that increases are always associated with negative consequences in social and parental interactions. In keeping with this more nuanced view, elevated glucorticoids are associated with greater maternal responsiveness in humans Fleming et al.

One of the complexities of glucorticoid-behavior research is illustrated by the contradictory findings on the same population of meerkats: natural elevations in cortisol were associated with higher pup-feeding rates Carlson et al. In keeping with the models mentioned above, glucocorticoids effects on parental behavior reported in meerkats may differ with concentrations i.

Concentrations may also vary with how quickly the increase occurs: the change in cortisol from an injection may be perceived as a sudden emergency e. We will try to evaluate whether such a nuanced view is also appropriate for understanding the effects of glucocorticoids in primate fathers.

Glucorticoids in male primates appear to function in two main ways related to reproduction: they may facilitate paternal behavior in new fathers Nunes et al. Nunes et al. This result suggests that carrying infants is not extremely stressful but suggests that other stressors may impact both glucocorticoid levels and carrying frequency. Fathers raising their first litters, had higher cortisol levels than fathers raising their second consecutive litters, even without differences in carrying behavior, suggesting, as we do later for human males, that glucocorticoids may help new fathers focus on the appropriate behavioral requirements of the new role.

Glucorcorticoid levels of male cotton-top tamarins increase at about the same time as female levels increase during pregnancy Ziegler et al. Though the mechanism of transmission within pairs is unknown, it presumably involves changes in arousal leading to behavioral changes to which the mate responds. Male cortisol and corticosterone levels peaked between weeks 13—16 of the week pregnancy, approximately 1—2 weeks after female levels increased Ziegler et al. Corticosterone concentrations were higher than cortisol concentrations during the mid-gestation peak in expectant male cotton-top tamarins, results that contrast with the general view that cortisol is the more important glucocorticoid in primates Ziegler at al.

The mid-pregnancy rise of glucocorticoids in females may stimulate a glucocorticoid response in male tamarins as in humans where cortisol concentrations of expectant human parents are correlated Storey et al. Despite being less consistent in their glucorticoid increases, less experienced males interacted with their mates more than experienced males Ziegler et al. It may be that behavioral coordination is more important for the transition to fatherhood in less experienced pairs, whereas hormonal coordination becomes more important as parental experience increases as in the human prolactin study mentioned previously, Delahunty et al.

Both male and female tamarins also experience an increase in glucocorticoids just before postpartum ovulation. At this time, male glucocorticoid levels are positively correlated with their rising testosterone levels but these hormonal changes are not associated with any decrease in paternal care. Other studies have also found hormonal synchrony between dyad partners, involving glucocorticoids Storey et al.

Cortisol concentrations are highest just prior to birth in both mothers Fleming et al. The cortisol concentrations of expectant fathers were correlated with the cortisol concentrations of their pregnant partners Storey et al. Although higher cortisol is associated with greater maternal responsiveness Fleming et al.

Similar complexities exist for men: fathers who expressed concern after hearing baby cries show less of a cortisol decrease than men who did not express concern and partners of men with higher cortisol concentrations reported that they did more domestic work after their babies were born Storey et al. In contrast, fathers whose partners reported that they found parenting more difficult that they expected had higher cortisol concentrations than other fathers Storey et al.

Further, men who spent more time with their toddlers before testing had a greater decrease in cortisol over the test period than men who had spent less time Storey et al. Elevated concentrations of cortisol are thus associated with both positive increased sensitivity and negative stress aspects of human parental care. Most reports on oxytocin in non-human primates have focused on pair-bonding behaviors rather than on paternal care.

However in one study, intranasal oxytocin increased the tolerance of marmoset fathers when transferring food to weanling infants Saito and Nakamura, Most of the work with oxytocin and vasopressin on non-human primates concerned how these hormones affect receptors in particular brain regions e. Common marmosets and several other species of New World primates, as well as the tree shrew Tupaia belangeri have a variant of the oxytocin molecule, Pro 8 OT that is unusual, as oxytocin is a conserved molecule with no variation in other nonhuman primates Lee et al.

However, several of the species with this variant are not bi-parental or monogamous, questioning whether this difference in structure influences its function. The most exciting recent developments in the biological basis of human paternal behavior involve oxytocin see Feldman, this issue. Oxytocin concentrations are higher in fathers than non-fathers Mascaro et al. Unlike some other hormones implicated in human paternal behavior, there are no sex differences in parental oxytocin concentrations in blood and saliva.

Thus, in some cases values are not analysed separately for men and women e. Infants of parents whose oxytocin concentrations increased the most were higher in affect synchrony than low-oxytocin, low affect-synchrony parents Feldman et al. Two current developments in oxytocin research have generated a great deal of attention: the effects of intranasal oxytocin and the discoveries that polymorphism in oxytocin receptor nucleotides is associated with differences in parental behavior and social responsiveness.

Several studies have now demonstrated that intranasal oxytocin increases paternal responsiveness Naber et al. The decrease was significantly less in men with early parental separation than in men from intact families. These results suggest that there is an early developmental component to this interaction between hormones that affects the degree to which oxytocin effectively reduces later stress.

Much current research has focused on single nucleotide polymorphisms SNPs , which reflect variation in alleles coding for oxytocin receptors. Other oxytocin receptor SNPs have also been implicated in parental responsiveness: Marsh et al. It may be that the study of parental behavior, like recent research on autism e.

It appears that vasopressin is involved in a wider range of male reproductive activities than is oxytocin. A particular variant in the vasopressin AVPR1a receptor allele was associated with higher relationship discord in men than other variants Walum et al.

In contrast, for women, oxytocin was more closely related to these factors than vasopressin: certain variants in oxytocin receptors genes Walum et al. Apter-Levi et al. Further, Gray found that the fathers of younger children had higher vasopressin concentrations than fathers of older children. Weisman et al. One focus in the animal research on paternal behavior is the role of the distribution of vasopressin receptors AVPR1a in the brain.

The focus in the human literature has been on vasopressin, oxytocin and to a lesser extent, testosterone in relation to brain areas associated with emotion, reward and decision-making. First-time and experienced marmoset fathers have a higher density of dendritic spines on pyramidal vasopressin neurons in prefrontal cortex compared to non-fathers Kozorovitskiy et al.

These results provide evidence for structural reorganization in the prefrontral cortex and a parallel enhancement in the abundance of vasopressin V1a receptors that indicate plasticity in the brain with parenting experience. Most of the other neuropeptide studies on biparental non-human primates have been mainly focused on pair bonding.

Few studies have examined the brain for receptor mapping, gene expression and structural plasticity in relationship to paternal care. The distribution of vasopressin producing cells, their projections and AVP receptors and oxytocin binding sites have been mapped in the common marmoset opening the door for understanding the role of neuropeptides in social affiliations Wang et al. Marmosets show differences in hypothalamic and pituitary hormones with paternal experience Woller et al.

Parentally experienced male marmosets without resident dependent infants show significantly higher concentrations of prolactin in their hypothalamus and reduced concentrations of dopamine compared to non-fathers.

Only the experienced fathers had elevated oxytocin concentrations. This study suggests that differences in secretion of prolactin and oxytocin may be related to paternal experience in male marmosets. Unlike the other studies revealing experiential changes, the changes here for the experienced males were not a direct priming effect from infant sensory stimuli since none of the males were caring for dependent infants.

These results lead to speculation that male brains undergo long-term changes after parental experience, as occurs in females. In biparental species, neuroplasticity and altered brain hormones apparently occurs in both sexes. Mascaro et al. First, hormonal changes may facilitate the development of paternal responsiveness by enhancing the emotional or reward brain circuitry and by increasing the salience of sensory cues from social companions.

Finally, hormonal changes may be involved in differential brain activation that diverts males from the mate acquisition component of reproductive effort towards the provision of paternal care. We will evaluate these possibilities in light of a number of the recent brain imaging studies.

Fathers showed greater brain activation to videos of infants averaged between their own and a matched unfamiliar infant than to a doll in several cortical areas including the orbitofrontal cortex, the superior, middle and inferior frontal gyri as well as in the caudate nucleus Kuo et al. The inferior frontal gyrus was also more active in fathers viewing a video of their own infant compared to the unfamiliar infant, as were the supramarginal and middle temporal gyri.

Fathers with higher testosterone concentrations after the infant interaction showed a significantly higher activation in caudate to their own compared to the other infant. Though low levels of testosterone are generally more closely linked to paternal responsiveness, the authors suggest that hormones may enhance the rewarding properties of infant stimuli. It may also be the case that men with higher testosterone focus more attention on their own babies than on infants in general, compared to possibly more paternal men with lower testosterone.

Fathers showed greater activation than non-fathers in areas associated with processing emotion in faces caudal middle frontal gyrus , reward processing orbital frontal cortex and mentalizing temporo parietal junction. Testosterone was lower and oxytocin concentrations were higher in fathers than non-fathers. Compared to fathers, non-fathers showed greater activation to sexual stimuli in areas associated with reward and motivation to approach nucleus accumbens and dorsal caudate.

They found that activity in the anterior insula, an area involved in empathy, was positively related to the number of CAG repeats and negatively correlated with having a restricting negative interaction, suggesting that the most effective fathers had intermediate activity in the anterior insula.

Taken together, these studies provide support for all three hypotheses in Mascaro et al. Changes in testosterone concentrations and individual differences in androgen receptors are implicated in these changes. Despite the lack of difference in plasma concentrations of oxytocin and vasopressin in mothers and fathers, there are differences in activated brain areas that may support the sex differences in parent-infant behavioral interactions. Atzil et al. Parents showed a high level of intra-couple synchrony in several brain areas involved in empathy and social cognition.

In addition, both fathers and mothers had significant correlations between blood concentrations of neuropeptide hormones and some critical brain areas. Fathers showed a significant positive correlation between activation of the amygdala and vasopressin concentrations whereas mothers showed a similar relationship between amygdala activation and oxytocin concentrations. The authors suggest that these sex differences reflect the more ancient and pervasive emotional role of mothering compared to the more recently evolved and more facultative, learning-based components in human paternal care.

Seifritz et al. Specifically, women responded to both infant crying and laughter with a deactivation of the anterior cingulate cortex, whereas men did not. Parents of both sexes showed greater response in the amygdala to the infant crying than laughing, whereas non-parents responded more to laughter. Abraham et al. They chose heterosexual couples in which the mothers were the distinct primary caretakers PC partnered by secondary care SC fathers.

In addition, Abraham et al. Within the homosexual group, there were no differences between the biological and adoptive fathers in behavior, oxytocin concentrations or the extent of activations in any brain areas tested.

The PC mothers and PC fathers showed equally high concentrations of behavioral synchrony with their infants and the extent of amygdala activation, with concentrations that were significantly higher than those of SC fathers. Overall for fathers, time spent in childcare was positively correlated with the extent of amygdala activation. In contrast to findings for the amygdala, activation of the superior temporal sulcus STS , part of the social-cognitive network, was equally high in PC and SC fathers and both were significantly higher than PC mothers.

Maternal oxytocin concentrations were correlated with activation in the ventral anterior cingulate cortex, a component of the emotional bonding system, whereas both groups of fathers showed oxytocin concentrations that only correlated with activity in the STS. These results suggest that at least part of what were assumed to be sex differences may actually be differences in the time and energy individuals devote to the parental role. Support for this view comes from the time allotment of same-sex and different-sex parents: parents in same-sex men or women couples and women in different-sex couples spent more time with their children than men in the different-sex couples data from the American Time Use Survey, Prickett et al.

Many researchers in this field have either looked at testosterone or some combination of oxytocin and vasopressin. They note that increases in oxytocin are involved in both the sexual system e. In contrast, testosterone concentrations change in opposite directions in these two relationship contexts.

One prediction that follows from this framework is that the social context should determine whether increased oxytocin should be associated with cause or result from? For example, increased oxytocin in a parental context should be associated with a decrease in testosterone.

This prediction was not supported in a recent study by Weisman et al. These researchers found, however, that men with the lowest baseline testosterone concentrations were initially the most paternal and had the largest testosterone increase. If increased testosterone does play a role in protective paternal tendencies, it may be that the intranasal oxytocin enhanced that response more in the lower-testosterone, highly parental men, compared to other men.

How specific are oxytocin effects? Given the spate of research on how intranasal oxytocin augments many positive and a few negative social behaviors, Churchland and Winkielman question whether oxytocin effects on behavior are specific to functional categories, or are more general, reducing anxiety in a way that facilitates social interactions e.

They suggest that we may see specific effects because of the testing situations we set up, and it would be useful to see whether there are individual patterns or similarities across behavioral categories. How important is prolactin in the onset and maintenance of primate paternal care?

There are clear effects of prolactin on energy balance in non-human primate fathers but we need more naturalistic studies to determine whether prolactin, controlled by dopamine, functions to regulate parental tendencies and motivation to approach and care for young. We do know that hormonal changes are associated with how much time fathers and offspring are together and whether the paternal care is effective.

However, we still do not know the biological basis of individual differences in the tendency for males to initiate contact with offspring as a mechanism to initiate these changes. Another major, largely unanswered, question revolves around interactions of oxytocin with other hormones, specifically testosterone and vasopressin. This interaction could occur anywhere between the receptor level and behavior, the latter occurring if multiple hormones change behavioral tendencies in incompatible ways.

A fruitful line of future research would be to compare oxytocin and testosterone responses to mating and parental stimuli in the same individuals. Further, we need to know whether peripheral concentrations of hormones can tell us much about brain mechanisms or whether plasma and cerebral spinal fluid concentrations are not related. How can we use this research to inform education and propose interventions that will improve paternal care?

Carter suggests that we need to know whether humans are like the most paternal rodent species where brain mechanisms enabling paternal care are established in neonates e. It appears that humans have an extended early period where parental loss, neglect and abuse can change neural and hormonal responses, producing long-term deficits in adult parenting behaviors.

We do not know if this damage can be reversed and if so whether there is a critical period for doing so. We also do not know whether there is a physiological basis for variation in resilience after early parental deficits since there are clear individual differences in the damage resulting from these early traumas. Perhaps as we get a clearer understanding of how receptor polymorphisms and intranasal hormone applications interact with early experience to affect behavioral variation, we can move forward with more individually based treatments.

Finally, this research suggests that it is important to provide education that promotes the importance of father-child bonds starting at prenatal classes? Research on synchronous responses within parental couples also points to the important role of the parental pair bond, not only for logistical support, but also for enhancing physiological mechanisms within couples that promote effective parental care.

There appear to be several similarities in the hormonal mechanisms underlying paternal behavior in all primates studied to date, despite the differences in what studies can be done in the different species see Table 1. In most cases, lower testosterone and higher prolactin concentrations are associated with higher levels of paternal responsiveness. Moderate increases in glucocorticoids may help to synchronize hormonal changes in pair members of all species studied.

There is more neuropeptide research on humans in connection with paternal behavior, but the association of these compounds, particularly vasopressin, with pair bonding in non-human primates suggests that similar mechanisms may be involved in paternal behavior there as well.

Further, he found that virgin female rats and both intact and castrated males would act parentally if they were exposed to pups for several days, a process called sensitization Rosenblatt, Overall, it appears that the onset of primate paternal care more closely resembles the sensitization process than it does the rapid hormonally induced onset of maternal behavior in parturient females. Hormonal changes are involved in primate paternal behavior but they appear to follow, rather than precede, social interactions.

As with mammalian maternal care, hormones seem to be less involved in the maintenance of paternal behavior, although retaining low levels of testosterone may help males focus on offspring care rather than on mate acquisition. Prolactin may still play a role even after parental behavior has been established, but it may be involved in changes of motivation to approach young, effects that may be more readily observed in naturalistic settings, rather than in the lab.

Glucorticoids appear to be involved in behavioral changes that coordinate preparatory physiological and behavioral changes within pairs. There is now clear evidence in this literature that a change in the hormonal state of one individual increased via intranasal oxytocin can affect the behavior and hormonal state of the dyad partner the infant, Weisman et al.

Recent imaging studies continue to inform us about the neural substrates for hormonal action in brain areas associated with parental behavior. National Center for Biotechnology Information , U. Horm Behav. Author manuscript; available in PMC Aug 1. Anne E. Ziegler b. Toni E. Author information Copyright and License information Disclaimer. Copyright notice.

The publisher's final edited version of this article is available at Horm Behav. See other articles in PMC that cite the published article. Abstract We review recent research on the roles of hormones and social experiences on the development of paternal care in humans and non-human primates. Evolution, distribution and function of paternal care in primates 1. Overview of the hormonal basis of primate paternal care 2a.

Overview of hormonal basis of primate paternal care: Non-human primates While male and female primates have gonadal differences derived from their genetic sex, there are a number of anatomical similarities that suggest males could respond to infants, both behaviorally and hormonally, as do females. Overview of hormonal basis of primate paternal care: Humans Storey et al.

Table 1 Summary of relationships between hormone levels and paternal behavior in primates at different stages e. Open in a separate window. Testosterone 3a. Testosterone in non-human primate fathers Male primates need to have flexible hormonal responses since behaviors requiring high testosterone, such as mate guarding or territorial defense behaviors, co-occur with low-testosterone paternal care Ziegler et al.

Testosterone in human fathers Storey et al. Prolactin Prolactin is known for its role in promoting maternal behaviors in mammals Bridges et al. Prolactin in non-human primate fathers Researchers have suggested that prolactin has both a preparatory role and short-term response role in primate paternal behavior. Prolactin in human fathers As in male tamarins and marmosets, human fathers also have been reported to exhibit increased prolactin concentrations before the birth Storey et al.

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