ÐÏࡱá > þÿ Q S þÿÿÿ P ÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿÿì¥Á À ð¿ N bjbj,(,( .R NB NB ]9 ½ ÿÿ ÿÿ ÿÿ · R R 8 8 8 8 8 ÿÿÿÿ L L L 8 L s% r ´ ´ ´ ´ ´ ´ ´ ´ Æ$ È$ È$ È$ È$ È$ È$ , å( ² + T ô$ 9 8 ´ ´ ´ ´ ´ ô$ à! 8 8 ´ ´ -% à! à! à! ´ 8 ´ 8 ´ Æ$ à! ´ Æ$ à! à! : D$ , $ ÿÿÿÿ ptkcÇÍ ÿÿÿÿ ´ , p$ ²$ C% 0 s% $ ë+ à! ë+ $ à! 8 $ ´ ´ ´ ô$ ô$ à! ´ ´ ´ s% ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ÿÿÿÿ ë+ ´ ´ ´ ´ ´ ´ ´ ´ ´ h þ : Prolactin hormone and paternal care in primates
Prolactin a peptide neurohormone secreted by the anterior pituitary gland and regulated by the hypothalamus. Even though it has been reported to have around 85 different biological functions it is best known for its effect on lactation and maternal behavior (Bole- Feysot C., Goffin V., Edery M., Binart N. & Kelly P. A (1998)). For this reason it is generally considered a female hormone, but it is also present in the male gender, where it naturally does not have any affect on lactation (Physiology of Domestic animals). Despite the absence of lactation in the male, studies have shown that prolactin might actually have an influence on paternal care for the infants in various species. Paternal care will in this essay be considerd as a suite of behaviors performed by a mature male (the putative/social father of the immature young), which would not be perfomed in the absence of the young (Fernandez-Duque, E., Valeggia, C. R. & Mendoza, S. P. (2009)). The intention of this essay is to explore more deeply the effect prolactin has on the expression of paternal care, focusing on our own animal order, the primates.
The influence prolactin has on paternal care have been intensely studied in birds and fish, and suggest that there is a significant relationship between prolactin plasma concentrations and paternal care in these species. This has lead scientists to believe that prolactin is important in the expression of paternal care. (Schradin, C. & Anzenberger, G. (1999)), (Ziegler, T. E., Prudom S. L., Refetoff Zahed, S., Parlow, A. F. & Wegner, F. (2009)). Fewer studies have been carried out concerning primates but as more exploration of the domain is conducted there appear to be a connection between prolactin and paternal behaviour in several primate species as well (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)). It has not been proved that prolactin and paternal care has a causal relationship (Fernandez-Duque, E., Valeggia, C. R. & Mendoza, S. P. (2009)) but a number of studies show that the prolactin plasma level is significantly higher in fathers than in non-fathers. Additionally, it is know that horomones can have a direct effect on behavior when they implement their action on the Central Nervous System (Schradin, C. & Anzenberger, G. (1999)). Even though prolactin is a hormone built up by approximately 200 amino acids, and there is some debate to whether or not it can cross the blood-brain barrier, the coroid plexus does have prolactin receptors and is believed to play a considerable role in the movement of prolactin across the barrier and into the brain (Schradin, C. & Anzenberger, G. (1999)). These factors may suggest that prolactin has a designated role in paternal behaviour. Indeed, it has been established that in some primate species the level of circulating prolactin is different in males that are fathers and males who are not fathers (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)). Several investigations show that prolactin levels are increased pre-partum in fathers, as well as in the pregnant female, for instance in the cotton-top tamarine (Saguinus oedipus) and the common mormoset (Callithrix jacchus) (Ziegler, T. E., Prudom S. L., Refetoff Zahed, S., Parlow, A. F. & Wegner, F. (2009)), (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)). Another common observation is that experienced fathers, usually having fathered three or more offspring, have a higher level of prolactin than inexperienced fathers and that they also show a higher amount of parternal care and react in a more positive way to the infant (Ziegler, T. E., Prudom S. L., Refetoff Zahed, S., Parlow, A. F. & Wegner, F. (2009)), (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)). It has been discovered that when the father holds for the first time or he hears the cry of his first child, the prolactin concentration would decrease compared to the pre-partum period, whereas it increases if the father is not a first-time father. These observations has lead to the belief that some particular hormonal changes happens when the male becomes a father for the first time (Schradin, C., Reeder D. M., Mendoza, S. P. & Anzenberger, G. (2003)), (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)). Also, according to (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)), there seems to be a base-line level of prolactin which implement a trait-like characteristic of the individual (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)). when he has his first child. These observations coincide with care-giving being one of the main factors for infant survival, and in that way ensuring the transfer of the fathers genes to the new generations (Schradin, C. & Anzenberger, G. (1999)). Prolactin level in human fathers are higher before birth than after, and demonstrate a quite stable hormone concentration for the first six months of the infants life (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)), whereas in Cottontop tamarins the prolactin level in the father increase around two weeks prepartum and reach a maximum level two weeks after birth (Schradin, C. & Anzenberger, G. (1999)), (Almond, R. E. A., Ziegler, T. E & Snowdon, C. T. (2008)).
Although there is some variation among these species, it has been clearly demonstrated that the presence of a pregnant mate, and later the infants presence, could be associated with elevated levels of prolactin in the male (Almond, R. E. A., Ziegler, T. E & Snowdon, C. T. (2008)). The Titi monkey (Callicebus cupreus), Common marmoset and Goeldis monkey (Callimico goeldii) all have different commencement of the increased prolactin levels. In some the level was always high, whereas in others the level was raised in the period before the offspring was born. However, in all three cases, the level of prolactin was the highest 1-3 weeks after the birth of the offspring (Schradin, C., Reeder D. M., Mendoza, S. P. & Anzenberger, G. (2003)). These studies demonstrate that prolactin production is not only caused by infant stimuli and might mostly be important for the onset of paternal care and not so much for maintainig it (Schradin, C. & Anzenberger, G. (1999)). In female rats it has been discovered that prolactin is necessary for the mother's caring for the young but that steroid hormones are essential in order to induce the effect prolactin has on the maternal behaviour (Schradin, C. & Anzenberger, G. (1999)). Moreover, this seems to also be the case with onset of paternal care and it is possible that steroids induce the expression of prolactin receptors (Schradin, C. & Anzenberger, G. (1999)). Prolactin and testosterone usually have a negative relationship toward one another; when one decrease the other increases and vice versa. This relationship is reduced both during elevated and decreased prolactin levels. Estradiol also has a negativ relationship with prolactin, but it, unlike the relationship with testosterone, are mentained with both decreased and increased prolactin levels; decreased prolactin levels resulted in an elevated estradiol level and vice versa. (Ziegler, T. E., Prudom S. L., Refetoff Zahed, S., Parlow, A. F. & Wegner, F. (2009)). Paternal care in primates is clearly varied among species (Fernandez-Duque, E., Valeggia, C. R. & Mendoza, S. P. (2009)) but in human and some monkey species, such as tamarins, marmosets, titi monkey and owl monkey there is a considerable level of paternal care (Fernandez-Duque, E., Valeggia, C. R. & Mendoza, S. P. (2009)) (Schradin, C. & Anzenberger, G. (1999)). It is direct and lasts for a considerable amount of time and is observable at different circumstances (Fernandez-Duque, E., Valeggia, C. R. & Mendoza, S. P. (2009)). In the marmosets and tamarins it is also obligate (Schradin, C. & Anzenberger, G. (1999)). In different species of the New world monkeys, the onset of the paternal behavoir occured at variable times in the infants life in different species of the new world monkeys . In two species , the father started caring for the infant within the first day, while in one the start for paternal care was 3 weeks after the birth of the offspring (Schradin, C., Reeder D. M., Mendoza, S. P. & Anzenberger, G. (2003)). The levels of prolactin were most similar in those monkeys having a similar way of interacting with the infant. In the monkey species where the primary care is given by the father, the level of prolactin were the highest (Schradin, C., Reeder D. M., Mendoza, S. P. & Anzenberger, G. (2003)). In human fathers, the males that were most influenced by signals given by the infant presented higher proalctin levels than the more passive fathers. The prolactin hormone in the father was associated with his promting and reinforcing of the exploratory behaviour and play of the child (Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010)). In all of the New world monkeys studied in this case, the families were built up by not only the mating pair, but also by adult and subadult offspring, called helpers. The level of circulating prolactin were not as high in the helpers even though they took part in the care of the young (Schradin, C., Reeder D. M., Mendoza, S. P. & Anzenberger, G. (2003)). There was a significant difference in the level of prolactin in general between the father of the infant and their adult sons that served as helpers. Also the same male showed differences from the time he himself was a helper until the time he was the father. In the second case the level of prolactin was higher. In one species (C. Jacchus) fathers had higher levels of prolactin during times without preambulatory infants than their adult sons did (Schradin, C., Reeder D. M., Mendoza, S. P. & Anzenberger, G. (2003)). Cotton-top tamarines also show increased prolactin levels during midpoint of their mate´s pregnancy, but this conserned only experienced fathers and not first time fathers. The age of the male showed importance only when compared with the adult sons. When the two groups were considered alone, age had no significant importance. A young father showed higher levels of prolactin than an older male who only surved as a helper (Schradin, C., Reeder D. M., Mendoza, S. P. & Anzenberger, G. (2003)). Another interesting fact concerning the cotton-top tamarines is that the timing of the prolactin peak as averaged by month, correlates exactly with the number of infants surviving from the previous birth (Almond, R. E. A., Ziegler, T. E & Snowdon, C. T. (2008)). Studies conserning the Common marmoset has given some very interesting results. In an investigation in which 15 experienced fathers where chosen and studied during three pregnancy periods, one with normal prolactin levels, one with decreased and the last one with elevated prolactin levels. The decreased prolactin gave a decrease in the fathers responsiveness toward the young, as one could expect, but the elevated dose also gave the same result. Another important discovery that this study showed was prolactins effect on weight loss. Normally fathers of the common marmoset loses a lot of weight during infant caring, because of the extra energy recuired for carrying, feeding etc. With the decreased treatment the faters lost even more weight than normal, but higher levels resulted in less or even no amont of weight loss (Ziegler, T. E., Prudom S. L., Refetoff Zahed, S., Parlow, A. F. & Wegner, F. (2009)). Untill now there is no absolute proff that prolactin and paternal care have a direct relationship, but much reasearch are suggesting that they do. One major fact pointing towards this is that prolactin plasma level is significantly higher in fathers than in non-fathers. The cause of this increase in prolactin levels conserning fathers have both been suggested to be caused by the precens of infants and an experienced mother, but neither has b e e n s u c c e s f u l l y c o n f i r m e d . ( T h e r e a r e m a n y i n t e r s p e c i a l d i f f r e n c e s a m o n g t h e p r i m a t o r d e r c o n s e r n i n g t h e s u p p o s e d r e l a t i o n s h i p o f p a t e r n a l c a r e a n d p r o l a c t i n , b o t h c o n c e r n i n g t h e t i m i n g o f i n c r e a s e d p r o l a c t i n l e v e l s a n d i t s e f f e c t . O n e t h i n g t h a t s o f a r e h a s b e e n c o m m o n i n a l l s t u d i e d s p e c i e s i s t h e h i g h e r l e v e l o f p r o l a c t i n i n e x p e r i e n c e d f a t h e r s c o m p a r e d t o n e w o n e . ( E v e n t h o u g h t p r o l a c t i n h a v e n o t b e e n p r o v e n t o h a v e a d i r e c t r e l a t i o n s h i p w i t h p a t e r n a l c a r e , i t i s v e r y l i k e l y t h a t i t w i l l h a v e a i n d i r e c t e f f e c t , s i n c e i t g e n e r a l l y h a v e m o r e t h a n 8 5 d i f f e r e n t b i o l o b i c a l f u n c t i o n s a n d h a v e a c l o s e r e l a t i o n s h i p w i t h o t h e r s t e r o i d a l h o r m o n e s a n d h a s b e e n s h o w n t o r e d u c e w e i g h t l o s s . ( T h e r e i s c e r t a i n l y a r e l a t i o n s h i p b e t w e e n p a t e r n a l c a r e a n d p r o l a c t i n l e v e l s in primates, but in what form more research is needed to answer that question. Especially more species specific studies, since there are many interspecies differennces. References: Gordon, I., Zagoory-Sharon, O., Leckman, J. F. & Feldman, R. (2010) Prolactin, Oxytocin, and the development of paternal behavior across the first six months of fatherhood. Hormones and behavior 58 (2010) 513-518 Fernandez-Duque, E., Valeggia, C. R. & Mendoza, S. P. (2009) The Biology of Paternal Care in Human and Nonhuman Primates. The Annual Review of Arthropology 2009 38:115-30 Schradin, C. & Anzenberger, G. (1999) Prolactin, the Hormone of Paternity. News in physiological sciences volume 14, December 1999 Schradin, C., Reeder D. M., Mendoza, S. P. & Anzenberger, G. (2003) Prolactin and Paternal Care: Comparison of Three Species of Monogamous New Worl Monkeys (Callicebus cupreus, Callithrix jacchus and Callimico goeldii) Jornal of Comparative Psychology. 2003, Volume 117 No. 2 166-175 Ziegler, T. E., Prudom S. L., Refetoff Zahed, S., Parlow, A. F. & Wegner, F. (2009) Prolactin´s Meditative Role in Male Parenting in Parentally experienced Marmosets (Callithrix jacchus)Hormones and Behavior. Volume 56, issue 4, October 2009, 436-443 Almond, R. E. A., Ziegler, T. E & Snowdon, C. T. (2008) Changes in Prolactin and Glucocorticoid Levels in Cotton-top Tamarin Fathers During their Mate´s Pregnancy: The Effect of Infants and Paternal Experience. American Jornal of Primatology. 70:1-6 2008 Bole- Feysot C., Goffi 0 > @ A o ¹ ø ù Þ ^ 6 { Î 6 u á ¯ Á  _ o Æ Ç × Ø Ù øìÚʺªìÚìÊÚìÊìÊÚÊÚÊÚÊìÊÊxÊìÊìÊìÊìÊÚdÊ 'h Oi B*CJ OJ PJ QJ ^J aJ phÿ !h Oi 6CJ OJ PJ QJ ^J aJ $h Oi 6CJ OJ PJ QJ ]^J aJ h Oi CJ OJ QJ ^J aJ h Oi B*CJ OJ QJ aJ ph h Oi B* CJ OJ QJ aJ ph h Oi CJ OJ PJ QJ ^J aJ "h Oi CJ OJ QJ ^J aJ mH sH h Oi CJ OJ QJ aJ h Oi OJ QJ ' 0 1 ô * Ì$ b+ ]5 «> ¸> ? 8@ »@ ×A ÒB ÑC äL \M ]M _M `M bM cM eM ú ø ø ø ø ø ø ò ò è ø ã ã ã ã ã ã ã ã ø Ü ø Ü ø Ü dð ¤ & F : Ä^ Ä ÄÄ $a$ Ù 3 y ½ ê ë 0 * Ò Ó Õ Ö ð + < N v ³ ÷ ± Ù °! Ø! l" " " t$ Ì$ % D% Ò% 8& & à& ' 9' ;' ( Ö( µ) û) + + , Ç, . Ð. ôäôäÐäôоÐäôоªôÐôÐÐооооÐôооооÐô¾ô¾ôäô¾ô¾ h Oi CJ OJ PJ QJ ^J aJ *h Oi 6B*CJ OJ PJ QJ ^J aJ ph 'h Oi B*CJ OJ PJ QJ ^J aJ phÿ "h Oi CJ OJ QJ ^J aJ mH sH 'h Oi B*CJ OJ PJ QJ ^J aJ ph h Oi CJ OJ PJ QJ ^J aJ h Oi CJ OJ QJ aJ 4Ð. z0 À0 Á0 È1 É1 ]5 «> û> e? {? @ (@ @ ¡@ XA A A A A ¶A ×A ~B §B ¥C ÃC çC L L ¾L äL (M EM \M ]M ^M `M aM cM dM fM gM iM jM ÒM ôM õM N N N ôâÎô¾ôâôâ«â«â«â«â«â«âôôôôôâ«â hWj j h Oi Uh Oi Uh Oi 6CJ OJ QJ aJ %h Oi 6CJ OJ QJ ^J aJ mH sH h Oi B*CJ OJ QJ aJ phÿ 'h Oi B*CJ OJ PJ QJ ^J aJ phÿ "h Oi CJ OJ QJ ^J aJ mH sH h Oi CJ OJ QJ aJ 1n V., Edery M., Binart N. & Kelly P. A (1998) Prolactin (PRL) and its receptor: actions, signal transduction pathways and phenotypes observed in PRL receptor knockout mice. Endocrine Reviews, June 1 1998 volume 19 no. 3 225-268 Physiology of Domestic animals, Sjaastad, O. V, Hove, K. & Sand, O. Scandinavian Veterinary Press 2003, first edition.
Physiology, Prolactin hormone and paternal care in primates Supervisor English program István Tóth
Zara Broström, Linnea Carlsson and Tone Hansen 20.11.12. Page 1/3
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