Abstract: The purpose of the present study was to gain more insight into the contribution of the forelimbs and hindlimbs of the horse to energy changes during the push-off for a jump. For this purpose, we collected kinematic data at 240 Hz from 23 5-year-old Warmbloods (average mass: 595 kg) performing free jumps over a 1.15 m high fence. From these data, we calculated the changes in mechanical energy and the changes in limb length and joint angles. The force carried by the forelimbs and the amount of energy stored was estimated from the distance between elbow and hoof, assuming that this part of the leg behaved as a linear spring. During the forelimb push, the total energy first decreased by 3.2 J kg(-1) and then increased again by 4.2 J kg(-1) to the end of the forelimb push. At the end of the forelimb push, the kinetic energy due to horizontal velocity of the centre of mass was 1.6 J kg(-1) less than at the start, while the effective energy (energy contributing to jump height) was 2.3 J kg(-1) greater. It was investigated to what extent these changes could involve passive spring-like behaviour of the forelimbs. The amount of energy stored and re-utilized in the distal tendons during the forelimb push was estimated to be on average 0.4 J kg(-1) in the trailing forelimb and 0.23 J kg(-1) in the leading forelimb. This means that a considerable amount of energy was first dissipated and subsequently regenerated by muscles, with triceps brachii probably being the most important contributor. During the hindlimb push, the muscles of the leg were primarily producing energy. The total increase in energy was 2.5 J kg(-1) and the peak power output amounted to 71 W kg(-1).

Abstract: The ability to jump over an obstacle depends upon the generation of work across the joints of the propelling limb(s). The total work generated by one hind limb of a horse and the contribution to the total work by four joints of the hind limb were determined for a jump. It was hypothesized that the hip and ankle joints would have extensor moments performing positive work, while the knee would have a flexor moment and perform negative work during the jump. Ground reaction forces and sagittal plane kinematics were simultaneously recorded during each jumping trial. Joint moment, power and work were determined for the metatarsophalangeal (MP), tarsal (ankle), tibiofemoral (knee) and coxofemoral (hip) joints. The hip, knee and ankle all flexed and then extended and the MP extended and then flexed during ground contact. Consistent with our hypothesis, large extensor moments were observed at the hip and ankle joints and large flexor moments at the knee and MP joints throughout ground contact of the hind limb. Peak moments tended to occur earlier in stance in the proximal joints but peak power generation of the hind limb joints occurred at similar times except for the MP joint, with the hip and ankle peaking first followed by the MP joint. During the first portion of ground contact (approximately 40%), the net result of the joint powers was the absorption of power. During the remainder of the contact period, the hind limb generated power. This pattern of power absorption followed by power generation paralleled the power profiles of the hip, ankle and MP joints. The total work performed by one hind limb was 0.71 J kg(-1). Surprisingly, the knee produced 85% of the work (0.60 J kg(-1)) done by the hind limb, and the positive work performed by the knee occurred during the first 40% of the take-off. There is little net work generated by the other three joints over the entire take-off. Velocity of the tuber coxae (a landmark on the pelvis of the animal) was negative (downward) during the first 40% of stance, which perhaps reflects the negative work performed to decrease the potential energy during the first 40% of contact. During the final 60% of contact, the hip, ankle and MP joints generate positive work, which is reflected in the increase of the animal's potential energy.

Abstract: A horse's legs are compressed during the stance phase, storing and then returning elastic strain energy in spring-like muscle-tendon units. The arrangement of the muscle-tendon units around the lever-like joints means that as the leg shortens the muscle-tendon units are stretched. The forelimb anatomy means that the leg can be conceptually divided into two springs: the proximal spring, from the scapula to the elbow, and the distal spring, from the elbow to the foot. In this paper we report the results of a series of experiments testing the hypothesis that there is minimal scope for muscle contraction in either spring to adjust limb compliance. Firstly, we demonstrate that the distal, passive leg spring changes length by 127 mm (range 106-128 mm) at gallop and the proximal spring by 12 mm (9-15 mm). Secondly, we demonstrate that there is a linear relationship between limb force and metacarpo-phalangeal (MCP) joint angle that is minimally influenced by digital flexor muscle activation in vitro or as a function of gait in vivo. Finally, we determined the relationship between MCP joint angle and vertical ground-reaction force at trot and then predicted the forelimb peak vertical ground-reaction force during a 12 m s(-1) gallop on a treadmill. These were 12.79 N kg(-1) body mass (BM) (range 12.07-13.73 N kg(-1) BM) for the lead forelimb and 15.23 N kg(-1) BM (13.51-17.10 N kg(-1) BM) for the non-lead forelimb.

Abstract: Drawing on studies of humans, rodents, birds and arthropods, I show that 'cognitive maps' have been used to describe a wide variety of spatial concepts. There are, however, two main definitions. One, sensu Tolman, O'Keefe and Nadel, is that a cognitive map is a powerful memory of landmarks which allows novel short-cutting to occur. The other, sensu Gallistel, is that a cognitive map is any representation of space held by an animal. Other definitions with quite different meanings are also summarised. I argue that no animal has been conclusively shown to have a cognitive map, sensu Tolman, O'Keefe and Nadel, because simpler explanations of the crucial novel short-cutting results are invariably possible. Owing to the repeated inability of experimenters to eliminate these simpler explanations over at least 15 years, and the confusion caused by the numerous contradictory definitions of a cognitive map, I argue that the cognitive map is no longer a useful hypothesis for elucidating the spatial behaviour of animals and that use of the term should be avoided.

Abstract: During locomotion, mammals update their position with respect to a fixed point of reference, such as their point of departure, by processing inertial cues, proprioceptive feedback and stored motor commands generated during locomotion. This so-called path integration system (dead reckoning) allows the animal to return to its home, or to a familiar feeding place, even when external cues are absent or novel. However, without the use of external cues, the path integration process leads to rapid accumulation of errors involving both the direction and distance of the goal. Therefore, even nocturnal species such as hamsters and mice rely more on previously learned visual references than on the path integration system when the two types of information are in conflict. Recent studies investigate the extent to which path integration and familiar visual cues cooperate to optimize the navigational performance.