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Stanley, C., & Shultz, S. (2012). Mummy’s Boys: Sex Differential Maternal Offspring Relationships in Semi-feral Horses. In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: In polygynous species with high reproductive skew in males, mothers often show differential investment between sons and daughters; however consistent sex differential investment has not been found by previous studies in horses. We investigated sex differences in mother-offspring relationships in nutritionally independent sub-adult semi-feral Carneddau Welsh mountain ponies Equus ferus caballus. Mothers and their sub-adult sons had consistently closer relationships than mothers and daughters. Stronger affiliative bonds between mothers and sons were quantified by their maintenance of closer proximity, higher rates of affiliative interactions and more frequent suckling bouts. These measures of affiliation were temporally associated with higher aggression levels directed towards sub-adults by other band members, particularly stallions. We suggest that aggression may serve as the proximate mechanism driving male dispersal in feral horses and that the stronger mother-son bond signals an attempt to delay their dispersal, highlighting conflict within the band concerning dispersal timing. Since males become increasingly central within the band over time, with mature stallions requiring excellent social skills in order to both acquire and keep a band of mares, we propose that delaying colts' dispersal allows for further development of these skills in a relatively safe environment. This additional investment is expected to maximise their reproductive success. This study illustrates how social network theory can be used to quantify individuals' social experiences, contributing to a greater understanding of the evolution of group living. It also gives us further insight into the mechanisms underlying dispersal in wild and semi-wild horse populations and how conflict often arises when individual needs differ.
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Heitor, F., & Vicente, L. (2008). Maternal care and foal social relationships in a herd of Sorraia horses: Influence of maternal rank and experience. Appl. Anim. Behav. Sci., 113(1-3), 189–205.
Abstract: The influence of maternal rank and experience on patterns of maternal care and social relationships of foals were investigated in a managed herd of Sorraia horses, Equus caballus. Social interactions and spatial relationships of 13 foals (seven females and six males) born to seven mares were examined from birth to 10 months of life, within the three major periods of foal development. Conflict over suckling between dam and foal was not generally affected by rank and experience, but higher-ranking mothers allowed more suckling during late lactation than lower-ranking mothers. Foals of higher-ranking mares spent more time in proximity to the mother during socialization. Maternal rank and experience did not significantly affect maternal protectiveness, foal independence from the mother or the development of affiliative relationships between foals and group members. Foals of higher-ranking mares received lower frequencies of aggression from other horses only in the first month of life. Dominance relationships among foals depended mainly on aggressiveness and were not associated with maternal rank. The large variability in maternal behaviour, the absence of a significant association between maternal rank and body condition at parturition and the stable social environment within this herd may partly account for the reported results.
Keywords: Horse; Rank; Experience; Maternal care; Social relationships
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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
Keywords: Equus caballus; feral horse; maternal investment; play
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Grogan, E. H., & McDonnell, S. M. (2005). Mare and Foal Bonding and Problems. Clinical Techniques in Equine Practice, 4(3), 228–237.
Abstract: A number of specific behavioral responses have been identified in mares and foals as the presumed behavioral interactive sequences supporting bonding. With the exception of the severely physically compromised foal, most failures of the mare foal bond appear to result from inadequate behavior of the mare. Six distinct forms of maternal behavior problems include ambivalence of the mare toward her foal, fear of the foal, nursing only avoidance of the foal, extreme protectiveness of the foal that becomes problematic in domestic confinement, savage attack (true rejection), and stealing or adoption of an alien foal. Management of maternal behavior problem cases in which the pair cannot be salvaged include foster (or nurse mares) and hand-rearing methods. Also presented are current practical resources related to managing certain types of inadequate maternal behavior and for rearing the orphaned foal.
Keywords: Equine; bonding; behavior; mare; neonatal; foal; inadequate maternal behavior; orphan foal; nurse or foster mare
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Broad, K. D., Curley, J. P., & Keverne, E. B. (2006). Mother-infant bonding and the evolution of mammalian social relationships. Phil. Trans. Biol. Sci., 361(1476), 2199–2214.
Abstract: A wide variety of maternal, social and sexual bonding strategies have been described across mammalian species, including humans. Many of the neural and hormonal mechanisms that underpin the formation and maintenance of these bonds demonstrate a considerable degree of evolutionary conservation across a representative range of these species. However, there is also a considerable degree of diversity in both the way these mechanisms are activated and in the behavioural responses that result. In the majority of small-brained mammals (including rodents), the formation of a maternal or partner preference bond requires individual recognition by olfactory cues, activation of neural mechanisms concerned with social reward by these cues and gender-specific hormonal priming for behavioural output. With the evolutionary increase of neocortex seen in monkeys and apes, there has been a corresponding increase in the complexity of social relationships and bonding strategies together with a significant redundancy in hormonal priming for motivated behaviour. Olfactory recognition and olfactory inputs to areas of the brain concerned with social reward are downregulated and recognition is based on integration of multimodal sensory cues requiring an expanded neocortex, particularly the association cortex. This emancipation from olfactory and hormonal determinants of bonding has been succeeded by the increased importance of social learning that is necessitated by living in a complex social world and, especially in humans, a world that is dominated by cultural inheritance. © 2006 The Royal Society.
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Brennan, P. A., & Kendrick, K. M. (2006). Mammalian social odours: attraction and individual recognition. Phil. Trans. Biol. Sci., 361(1476), 2061–2078.
Abstract: Mammalian social systems rely on signals passed between individuals conveying information including sex, reproductive status, individual identity, ownership, competitive ability and health status. Many of these signals take the form of complex mixtures of molecules sensed by chemosensory systems and have important influences on a variety of behaviours that are vital for reproductive success, such as parent-offspring attachment, mate choice and territorial marking. This article aims to review the nature of these chemosensory cues and the neural pathways mediating their physiological and behavioural effects. Despite the complexities of mammalian societies, there are instances where single molecules can act as classical pheromones attracting interest and approach behaviour. Chemosignals with relatively high volatility can be used to signal at a distance and are sensed by the main olfactory system. Most mammals also possess a vomeronasal system, which is specialized to detect relatively non-volatile chemosensory cues following direct contact. Single attractant molecules are sensed by highly specific receptors using a labelled line pathway. These act alongside more complex mixtures of signals that are required to signal individual identity. There are multiple sources of such individuality chemosignals, based on the highly polymorphic genes of the major histocompatibility complex (MHC) or lipocalins such as the mouse major urinary proteins. The individual profile of volatile components that make up an individual odour signature can be sensed by the main olfactory system, as the pattern of activity across an array of broadly tuned receptor types. In addition, the vomeronasal system can respond highly selectively to non-volatile peptide ligands associated with the MHC, acting at the V2r class of vomeronasal receptor.The ability to recognize individuals or their genetic relatedness plays an important role in mammalian social behaviour. Thus robust systems for olfactory learning and recognition of chemosensory individuality have evolved, often associated with major life events, such as mating, parturition or neonatal development. These forms of learning share common features, such as increased noradrenaline evoked by somatosensory stimulation, which results in neural changes at the level of the olfactory bulb. In the main olfactory bulb, these changes are likely to refine the pattern of activity in response to the learned odour, enhancing its discrimination from those of similar odours. In the accessory olfactory bulb, memory formation is hypothesized to involve a selective inhibition, which disrupts the transmission of the learned chemosignal from the mating male. Information from the main olfactory and vomeronasal systems is integrated at the level of the corticomedial amygdala, which forms the most important pathway by which social odours mediate their behavioural and physiological effects. Recent evidence suggests that this region may also play an important role in the learning and recognition of social chemosignals.
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Rudran, R. (1973). Adult male replacement in one-male troops of purple-faced langurs (Presbytis senex senex) and its effect on population structure. Folia Primatol (Basel), 19(2), 166–192. |
Bernstein, I. S., & Dobrofsky, M. (1981). Compensatory social responses of older pigtailed monkeys to maternal separation. Dev Psychobiol, 14(2), 163–168.
Abstract: Thirteen 3-5-year-old pigtailed monkeys were subjected to five 2-hr maternal separations while remaining in their normal social group. Significant changes in activity profiles were noted during separation and reunion phases. This suggests the continued social dependence of older offspring upon the matriarch. The shift in social activities reflected attempts by the juvenile and adolescent subjects to compensate for maternal absence by intensification of other affiliative social behavior and avoidance of potentially socially disruptive situation. The subjects oriented more towards kin in the absence of the matriarch, but actual time with kin decreased. Upon the return of the matriarch, the intensified some responses depressed during her absence and returned to preseparation social relationships. Play and aggressive responses declined whereas social approaches increased during maternal absences. Submissive responses declined upon the return of the matriarch, and play increased. The subjects also showed a marked, temporary increase of direct interaction, largely sniffing and grooming, with the matriarch upon her return.
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Hoff, M. P., Nadler, R. D., & Maple, T. L. (1981). Development of infant independence in a captive group of lowland gorillas. Dev Psychobiol, 14(3), 251–265.
Abstract: In March 1976, 3 lowlands gorillas (Gorilla gorilla gorilla) were born to primiparous females living with an adult male in a large compound at the field station of the Yerkes Regional Primate Research Center of Emory University. Observations of parent and infant behavior began at the birth of the infants, using several methods of data collection. This report focuses on the development of independence in these infants over the 1st 1 1/2 years of life. As expected, measures of mother-infant contact and proximity decreased with age. Several measures suggested that infant independence developed as an interactive process between mothers and infants, with primary responsibility changing over the months of study. Maternal behaviors that served to maintain mother-infant contact were found to decrease with age, with an eventual shift to infant responsibility for contact maintenance. Additionally, the adult male appeared to influence developing independence as reflected in the maternal protectiveness evoked by his behavior.
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Zhang, T. - Y., Parent, C., Weaver, I., & Meaney, M. J. (2004). Maternal programming of individual differences in defensive responses in the rat. Ann N Y Acad Sci, 1032, 85–103.
Abstract: This paper describes the results of a series of studies showing that variations in mother-pup interactions program the development of individual differences in behavioral and endocrine stress responses in the rat. These effects are associated with altered expression of genes in brain regions, such as the amygdala, hippocampus, and hypothalamus, that regulate the expression of stress responses. Studies from evolutionary biology suggest that such “maternal effects” are common and often associated with variations in the quality of the maternal environment. Together these findings suggest an epigenetic process whereby the experience of the mother alters the nature of the parent-offspring interactions and thus the phenotype of the offspring.
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