| 词条 | Endocrinology of parenting |
| 释义 |
Based on cross species evidence, some aspects of these mechanisms have been phylogenetically conserved from rodents to humans,[4] meaning that these mechanisms are adaptive for mammalian parenting and that the environment of evolutionary adaptedness of some parenting mechanisms may have evolved when mammals first evolved. The importance of these mechanisms are to regulate parental investment and to inform offspring about their environment, primarily those involving responsiveness and sensitivity. These are commonly mentioned in humans as important parenting characteristics that inform their infants about their environments.[5][6] Estrogen, progesterone, prolactin, and oxytocinNonhuman femalesMany nonhuman studies can be used as both potential models for humans and to show the phylogenetic conservation some endocrine signals.[1] Estrogen and progesterone released by ovaries during pregnancy make oxytocin receptors more sensitive in female rats[7] and is associated with the onset of maternal behaviors in other species as well.[3][8][9] Maestripieri found a very similar relationship, whereas estrogen and progesterone are increased during pregnancy and oxytocin was increased postpartum.[10] Estrogen, progesterone, and estradiol in pregnant mammals, in some species, but not all, correlate with maternal behavior before the birth of their offspring, with other infants, and after with their own infants.[3] However, an increase in hormones influences maternal behavior, but it is not always the cause of the onset of maternal behavior in females. Some studies on primates in which increased estrogen and progesterone have a negative or no correlation with maternal responsivity are in black tufted-ear marmosets,[11] common marmosets,[12] lowland gorillas,[13] and baboon On the other hand, in non-human primates, lactating females of multiple species, there is an alarming correlation with increased estrogen and progesterone and prolactin.[3] These species include black tufted-ear marmosets,[11] baboons,[14] rhesus macaques,[16] and gorillas.[13] Endogenous signals, like increasing estrogen compared to progesterone, increases the amount of prolactin, the “lactation hormone,” in the bloodstream, as well as exogenous cues from infants, like suckling.[17] Oxytocin, has been found to play a key factor in breastfeeding and increase similarly to prolactin, with increases in estrogen and infant cues.[18] Suckling also increases oxytocin levels in rhesus macaques,[10] and oxytocin has also been found in other non-human species to inhibit the rejection of offspring;[3] oxytocin is essential for responsive and sensitive caregiving.[19][20][3][21] Some specific examples include Francis's study on female rats which linked high amount of oxytocin receptors to increased grooming,[22] and another study by Maestripieri which linked oxytocin levels in free-ranging macaques to increased nursing and grooming.[10] However, experimental results are less conclusive. Nulliparous mice do not respond to pup calls, but when administered with oxytocin, they do.[23] A similar study conducted by Holmon and Goy tested nulliparous rhesus females, where post injection did not elicit a drastic response to infants, however there was a notable change in the adult's behavior, including increased proximity and touching.[24] Oxytocin is more often described as a hormone that facilitates bonding and not one that directly increases care.[25][24][26][10] Also, the mice were responding to pup calls and the rhesus macaque infants weren't necessarily providing cues. Similar to the explanation provided for the variation in estrogen and progesterone, rhesus macaques live in drastically different environments. Saltzman proposes that this is due to primates living socially and having a slower developmental trajectory, in which learning is more important.[3] Human femalesLike in many nonhumnan animals, human mothers go through a period of high progesterone during pregnancy that is followed by a decrease in progesterone and a subsequent increase in estrogen, prolactin and oxytocin. During pregnancy and postpartum, a high estradiol to progesterone ratio is associated with mother reported higher feelings of attachment.[27] High levels of progesterone, which is associated with pregnancy, inhibits prolactin and therefore lactation.[28] Prolactin increases during the initial stages of lactation and can be stimulated by estrogen, but not progesterone, and infant cues.[28] Research focuses on the role of prolactin for breastfeeding and less on other behaviors.[28] Prolactin increases with infant suckling, but not other forms of infant contact.[29] Oxytocin on the other hand increases with both suckling, and physical contact in human females.[28] Oxytocin in human females is associated with the level of physical affection and bonding. Feldman (2010) found that mothers who displayed “high affectionate contact” had increased oxytocin levels post interaction, but not mothers who displayed “low affectionate contact.[21]” Oxytocin is believed to provide a feedback loop, meaning that maternal-infant contact increases oxytocin and oxytocin increases maternal behavior and facilitates bonding.[30] In one study oxytocin also played a role on mother reported attachment to her fetus.[31] Nonhuman malesWynne-Edwards and Timonin recognize that paternal care is not primed in the same way as females simply because they do not go through pregnancy. Therefore, males do not go through the same hormonal changes as women.[32] The simplest way, through natural selection, for paternal care to evolve or be maintained is to use the same or similar pathways as females.[33] Wynne and Reburn (2001) suggest that fathers who are pair bonded and spend time with the soon to be mother may activate paternal pathways through various cues.[33] Estradiol increases just before their offspring's birth in black-tufted-ear marmosets and dwarf hamsters and possibly activates certain pathways involved in paternal behavior.[34][35] This is similar to estrogen and progesterone in pregnant females. However, the manipulation of estradiol does not increase or decrease paternal behaviors.[32] This may be similar to the finding that women who do not breastfeed or do not have vaginal births still responds to their infants.[19] Like expecting and new mothers, fathers in multiple mammals have elevated prolactin levels compared to non-fathers. These species include California mice,[36] Mongolian gerbils,[37] dwarf hamsters{{disambiguation needed|date=December 2018}},[38] meerkats,[39] marmosets,[40] and cotton-top tamarins.[41] However, the previously listed studies have different cues and are associated with different paternal behaviors, this may be due to species specific mechanisms or simply different contexts. The above species are biparental and the elevated prolactin levels in males were not exclusive to fathers. There are variable results in between males of different species as to the effects of oxytocin on paternal care. Oxytocin is unchanged in California mice before and after becoming fathers,[42] but the amount of paternal exposure to rats is associated with oxytocin and increased care.[41] However, multiple studies on biparental species show an association between paternal care and oxytocin.[42][43][33][44] Since the species in these studies are biparental, excluding rats, it is unclear as to why California mice do not have a change in oxytocin postpartum. Human malesIn human mothers, oxytocin is associated with high physical contact and affection. However, studies on fathers show that oxytocin is related to high stimulatory contact and exploratory play.[21][45] This supports three hypotheses:
Comparison of estrogen, progesterone, prolactin, and oxytocin between species and sexAcross multiple species and in some cases across sexes, there is evidence for the phylogenetic conservation of parental hormones. These include the relationships between the hormones estrogen, progesterone, prolactin, and oxytocin. In both non-human primates and humans, the increase in estrogen and progesterone during pregnancy is often followed by a decrease in progesterone and an increase in prolactin, postpartum.[11][14][13][24][28] In males across species, including humans, increased prolactin levels are associated with fatherhood.[36][37][38][39][40][41][88][89] In some studies on females across species, estrogen and progesterone prepartum is also related to oxytocin.[7][3][8][9][10] Although, the relationships between these hormones is similar across species, there is variation in the degree to how oxytocin effects behavior. For example, in some species, like rats, an increase in oxytocin greatly increases interactions with infants,[23] but an increase in oxytocin in macaques only mildly increased interactions.[24] However, the importance here is that oxytocin increased interactions in all of the relevant cited studies for females,[7][3][8][9][24] as well as the majority of studies cited for males.[42][43][33][44] In fathers across species the effects of oxytocin are more variable, however in general oxytocin is associated with increased paternal care.[42][43][33][44] In human fathers increased oxytocin is linked to increased involvement, however the type of involvement is different between fathers and mothers, where fathers focus more on stimulatory contact and exploratory play.[21][45] In human mothers oxytocin is associated with general care and affection.[21] Cortisol and prolactinNonhuman femalesContrary to the positive effects of oxytocin on maternal behavior, heightened levels of cortisol postpartum has been linked to a decrease in maternal care in nonhuman species, including the Western lowland gorilla,[46] baboons,[47] Japanese macaques,[48] and rhesus macaques.[10] However, there has been some evidence to support that the increase in cortisol levels during pregnancy results in an increased maternal care in two baboon species.[49][14] This variation possibly shows an ontogenetic difference in the role of cortisol. Human femalesCortisol in human mothers is elevated during pregnancy.[50] Human mothers with elevated cortisol during pregnancy more easily recognized, and were more attracted to, their own infant's odors postpartum, but this did not affect maternal attitudes toward their infants.[27] Behaviorally, mothers with higher levels of cortisol postpartum displayed higher levels of affectionate approach.[27] Women with higher cortisol levels were also more alert and sympathetic to infant crying.[51] Fleming and colleagues discovered that there was variation in how cortisol affected mothers with only one offspring and those with more than one offspring.[27] When cortisol levels were high, women with one offspring displayed more affectionate contact, and women with multiple offspring spent more time in caretaking activities. However, the authors did not report whether or not the two groups of women had significantly different levels of cortisol. If they were different, the specific levels of cortisol may be a contributing factor in the different behaviors. Human malesCouvade syndrome, or sympathy pregnancy, is defined by Masoni and colleagues as fathers having two or more pregnancy symptoms.[52] Human fathers with couvade syndrome have higher levels of prolactin, and cortisol levels than fathers without.[88] Cross-culturally couvade syndrome is associated with how close the mother and father are just prior to birth; in the postpartum period, these fathers invested more in direct care.[53] Increased paternal care is likely associated with offspring survival. A study on Canadian fathers found that, just before the birth of their infant, the fathers had significantly higher prolactin and cortisol levels.[88] The father's prolactin and cortisol levels correlated with the mothers, although her levels were significantly higher. This provides evidence for Wynne and Reburn's hypothesis that mothers may provide cues to the father to activate paternal care pathways.[33] In a separate study, fathers with higher prolactin compared to other fathers responded more positively to infant crying.[89] These levels also positively correlated with father experience. In the same study, cortisol levels in response to infant cries were negatively correlated with father experience. Higher cortisol levels was associated with higher father alertness and responsiveness to infant crying. Increased levels of cortisol in response to infant cries was greater in inexperienced fathers.[89] Comparison of cortisol and prolactin between species and sexThe behaviors associated with elevated cortisol levels appears to be more related to whether or not the mother was pre or postpartum, however the behaviors varied across species. In multiple studies on nonhuman females across species, found that high levels of cortisol postpartum was linked to low levels of maternal care.[46][14][48][10] However, when cortisol levels were high during pregnancy there was an increase in maternal care.[49][14] This difference did not hold true for humans of females and first time fathers. Increased cortisol levels in general increased maternal care.[27][51] Increased cortisol levels in first time human fathers correlated with an increase in responsiveness to their infant's crying, but not in fathers with more than one offspring.[89] TestosteroneNonhuman malesThe most commonly associated hormone with males is testosterone. It is believed to be the “anti-parental hormone”; it inhibits the activation of paternal mechanisms.[1] In many cases testosterone levels decrease when fathers have or actively care for their infants in non-humans.[54][55][56][32] However, testosterone can be converted into estradiol, which supports paternal behavior.[32][57] Testosterone is converted into estradiol through the process of aromatization.[57] Stated previously, with the repeated presence of pups to non-parental rats, caregiving mechanisms can inhibit other mechanism, like the avoidance mechanism;[15] similar inhibitions occur in male marmosets.[58] When male marmosets hold their infants, they did not have an increased testosterone response to novel females when they otherwise would. In other words, holding the infant inhibited the mating mechanisms. This could mean that caregiving supersedes mating in some situations. For example, it may be adaptive to continue to invest in your current offspring rather than potentially create another. Human malesMultiple studies on fathers have shown that a reduction in testosterone results in increased responsiveness to infant cues[59][60][61] and that fathers in general have lower testosterone than non-fathers.[61][62][63] Testosterone in human males decreases with the number of offspring human males have.[61] However, human males with higher level testosterone had greater activation of neural mechanisms when interacting with their own infants,[64][65] this may be due to the activation of a paternal protection mechanism.[66] Human femalesNot many studies analyze the effects of testosterone and maternal behaviors in humans. However, Fleming found that lower testosterone was associated with more affectionate contact in human mothers.[27] Comparison of testosterone between species and sexesBetween species variation in behaviors associated with testosterone in biparental species was not noted. Across species father experience was negatively correlated with testosterone[58][61] and lower levels of testosterone was associate with an increase in care.[59][54][61] Testosterone is more commonly studied in males than females. 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