Abstract: The behaviour of humans around horses is thought to have a substantial impact on how people are perceived in subsequent interactions and many horse trainers give detailed advice on how handlers should behave when initially approaching a loose horse. Here we report on three studies designed to explore the effect of different human approach styles on the behaviour of naïve and experienced horses. In the first study, the change in flight distance (distance at which horses started to avoid an approaching human) of twelve semi-feral Dartmoor ponies, undergoing training to allow handling, was assessed. Over the 10 handling sessions median flight distance decreased significantly (p < 0.001) from 2.38 m to 0.00 m and there was a significant positive shift in the ponies' behaviour following the appearance of the researcher (p = 0.002). In a second study the effect of a direct (vigorous, swinging a lead rope and with eye contact) versus indirect (relaxed, no rope swinging and without eye contact) approach style was assessed on six adult experienced riding horses. The mean flight distance during a direct approach style (6.87 m) was significantly greater than that which occurred during an indirect approach style (2.32 m). Direction of approach was not found to significantly affect flight distance. In a third study, the effect of the rope was removed and a similar method to the second study applied to a group of naïve, feral ponies. The effect of different components of approach style, speed of approach, handler body posture and direction of gaze, which might contribute to observed differences in behavioural responses, were then examined systematically in this population. This revealed no significant difference in mean flight distance between the two approach styles (2.28 m indirect versus 2.37 m direct approach), but ponies were significantly more likely to move off in trot (p = 0.025) and to travel further (p = 0.001) when a direct approach was used. Speed of approach was the most salient factor, with a fast approach increasing both the tendency to move off in trot (p < 0.001) and distance travelled (p < 0.001). Body posture (relaxed or tense) had no effect, while flight distance was significantly greater when the person was looking away (p = 0.045). These results suggest horses may have an important egocentric spatial barrier, which perhaps relates to personal space and triggering of the flight response. Contrary to popular belief, body posture did not appear to be very important in the contexts examined unless accompanied by extraneous aids, while the speed of approach is particularly significant. These results are of important practical relevance in reducing the risk of injury, and the effective management of horses with minimal stress.

Abstract: An adult female chimpanzee with previous training in the use of Arabic numerals 1&#8211;9 was introduced to the meaning of “zero” in the context of three different numerical tasks. The first two were cardinal tasks where the subject was required either to select numerals corresponding to the number of items presented on a computer screen (productive use of numerals) or to match sets of the appropriate size to numerals presented as samples (receptive use). The third task addressed the ordinal meaning of the same symbols where the subject was required to respond to numerals sequentially, arranging them into an ascending series. The subject mastered the recognition of the meaning of zero in all three tasks. However, details of her usage of the symbol revealed that transfer of the meaning between different kinds of tasks was incomplete, suggesting that the level of ion characteristic of human numerical ability was not attained in the chimpanzee. Over the course of acquisition leading to the high levels of accuracy eventually observed, the newly introduced zero appeared to shift along the length of a continuous numerical scale toward the lower end, while confusions with 1 remained the most frequently encountered mistakes. Such patterns of error thus suggest that Ai's understanding of the meaning of zero in relation to the rest of the number symbols was not consistent with an “absence of items versus presence of items” scheme.

Abstract: The integration of spatial maps in pigeons was investigated using a spatial analog to sensory preconditioning. The pigeons were tested in an open-field arena in which they had to locate hidden food among a 4x4 grid of gravel-filled cups. In phase 1, the pigeons were exposed to a consistent spatial relationship (vector) between landmark L (a red L-shaped block of wood), landmark T (a blue T-shaped block of wood) and the hidden food goal. In phase 2, the pigeons were then exposed to landmark T with a different spatial vector to the hidden food goal. Following phase 2, pigeons were tested with trials on which they were presented with only landmark L to examine the potential integration of the phase 1 and 2 vectors via their shared common elements. When these test trials were preceded by phase 1 and phase 2 reminder trials, pigeons searched for the goal most often at a location consistent with their integration of the L-->T phase 1 and T-->phase 2 goal vectors. This result indicates that integration of spatial vectors acquired during phases 1 and 2 allowed the pigeons to compute a novel L-->goal vector. This suggests that spatial maps may be enlarged by successively integrating additional spatial information through the linkage of common elements.

Abstract: A model for the evolution of cooperation shows that two conditions are necessary for cooperation to be stable: a hunting success rate that is low for single hunters and increases with group size, and a social mechanism limiting access to meat by non-hunters. Testing this model on TaI chimpanzees, Pan troglodytes, showed that (1) it pays for individuals to hunt in groups of three or four rather than alone or in pairs, and (2) cooperation is stable because hunters gain more at these group sizes than cheaters, owing to a meat-sharing pattern in which hunting, dominance and age, in that order, determine how much an individual gets. In addition, hunters provide cheaters (about 45% of the meat eaters) with the surplus they produce during the hunts. Thus, cooperation in Tai male chimpanzees is an evolutionarily stable strategy, and its success allows cheating to be an evolutionarily stable strategy for Tai female chimpanzees. In Gombe chimpanzees, cooperation is not stable, first, because hunting success is very high for single hunters, and second, because no social mechanism exists that limits access to meat by non-hunters. The analysis showed that some assumptions made when discussing cooperation in other social hunters might be wrong. This might downgrade our general perception of the importance of cooperation as an evolutionary cause of sociality.

Abstract: The four questions that Niko Tinbergen identified for behavioural biology ? evolution, function, development and causation ? are all important and should be studied in their own right. Recently, there has been a debate as to whether these four questions should be investigated separately or whether they should be integrated. Integration of the four questions has been attempted in novel research disciplines such as cognitive ecology, evolutionary psychology and neuroecology. Euan Macphail and I have criticised these integrative approaches, suggesting that they are fundamentally flawed as they confound function and mechanism. Investigating the function or evolutionary history of a behaviour or cognitive system is important and entirely legitimate. However, such investigations cannot provide us with answers to questions about the mechanisms underlying behaviour or cognition. At most, functional or evolutionary considerations can provide clues that may be useful for a causal analysis of the underlying mechanisms. However, these clues can be misleading and are often wrong, as is illustrated with examples from song learning and food storing in birds. After summarising the main issues in the neuroecology debate, I discuss some misunderstandings that were apparent in the responses to our critique, as well as some recent relevant data. Recent results do not support the neuroecological approach. Finally, I suggest that the way forward is a cautious and critical use of functional and evolutionary clues in the study of the mechanisms of behaviour.