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.

Abstract: Gene expression affects social behavior only through changes in the excitabilities of neural circuits that govern the release of the relevant motor programs. In turn, social behavior affects gene expression only through patterns of sensory stimulation that produce significant activation of relevant portions of the nervous system. In crayfish, social interactions between pairs of animals lead to changes in behavior that mark the formation of a dominance hierarchy. Those changes in behavior result from changes in the excitability of specific neural circuits. In the new subordinate, circuits for offensive behavior become less excitable and those for defensive behavior become more excitable. Serotonin, which is implicated in mechanisms for social dominance in many animals, modulates circuits for escape and avoidance responses in crayfish. The modulatory effects of serotonin on the escape circuits have been found to change with social dominance, becoming excitatory in dominant crayfish and inhibitory in subordinates. These changes in serotonin's effects on escape affect the synaptic response to sensory input of a single cell, the lateral giant (LG) command neuron for escape. Moreover, these changes occur over a 2-week period and for the subordinate are reversible at any time following a reversal of the animal's status. The results have suggested that a persistent change in social status leads to a gradual change in the expression of serotonin receptors to a pattern that is more appropriate for the new status. To test that hypothesis, the expression patterns of crayfish serotonin receptors must be compared in dominant and subordinate animals. Two of potentially five serotonin receptors in crayfish have been cloned, sequenced, and pharmacologically characterized. Measurements of receptor expression in the whole CNS of dominant and subordinate crayfish have produced inconclusive results, probably because each receptor is widespread in the nervous system and is likely to experience opposite expression changes in different areas of the CNS. Both receptors have recently been found in identified neurons that mediate escape responses, and so the next step will be to measure their expression in these identified cells in dominant and subordinate animals.

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.

Abstract: Recent attention has been drawn to the existence of individual differences in correlated behaviour across contexts, animal `personality' (Gosling 2001, Psychological Bulletin, 127, 45-86) and behavioural syndromes (Sih et al. 2004b, Quarterly Review of Biology, 79, 241-277). The causes of these patterns of behaviour are subjects of debate. Here, we present a very simple model of how adaptively managing noisy information, combined with differences in individual state, can lead to evolutionarily stable differences in how individuals respond to environmental cues. When information is very noisy, behavioural syndromes are most likely, but as long as there is some error, some types of individuals display the same behaviour in all contexts. In extreme cases, very few individuals display flexible behaviour, and different stable behavioural types dominate the population.

Abstract: Thousands of hectares of timothy (Phleum pretense L.) grown in the Mid-Atlantic region are infected by cereal rust mite (Abacarus hysterix) that causes discoloration and curling of leaves, decreased nutritional quality, and substantial decreases in yield. A decline in production of timothy hay can lower income for hay producers and cause horse owners to search for alternative hays. Low alkaloid reed canarygrass (Phalaris arundinacea L.) hay has potential as an alternative to timothy hay because it grows well in the Mid-Atlantic region, is believed to have a similar nutrient quality to timothy, and is not as susceptible to cereal rust mite. Eleven mature, stalled Thoroughbred geldings (549 +/- 12.1 kg) that were exercised daily were used to compare voluntary DMI and apparent nutrient DM digestibility of timothy and low-alkaloid Chiefton variety reed canarygrass hay. Horses were paired by age and BW and randomly assigned to timothy or reed canarygrass hay during a 14-d period to measure voluntary DMI followed by a 4-d period to measure apparent DM digestibility. Both hays met the minimum requirements for DE, CP, Ca, P, K, Fe, and Mn, but they did not meet the minimum requirements for Cu, Zn, and Na for horses at maintenance and averaging 550 kg of BW. Timothy hay seemed to have a lower CP concentration (14.4%) compared with reed canarygrass hay (17.1%) and a more desirable Ca:P ratio at 1.6:1 compared with 0.8:1 for reed canarygrass hay. Horses fed timothy consumed more hay (P <0.001) during the voluntary DMI period compared with horses fed reed canarygrass. Greater voluntary DMI of timothy occurred on d 1, 3, and 5 (P <0.05), but DMI was similar for other days. Apparent DM digestibility was greater in horses fed timothy hay by 9.6% compared with horses fed reed canarygrass hay (P <0.05). Horses fed timothy had greater DM digestibility of ADF (P = 0.001), NDF (P = 0.001), sugar (P = 0.05), and Ca (P = 0.001) but lower apparent DM digestibility of CP (P = 0.012) and crude fat (P = 0.004). Timothy hay was superior in voluntary DMI and apparent DM digestibility compared with low-alkaloid reed canarygrass hay fed to horses.