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Author  |
Arnold, W.; Ruf, T.; Kuntz, R. |
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Title |
Seasonal adjustment of energy budget in a large wild mammal, the Przewalski horse (Equus ferus przewalskii) II. Energy expenditure |
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Journal Article |
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Year |
2006 |
Publication |
The Journal of experimental biology |
Abbreviated Journal |
J Exp Biol |
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Volume |
209 |
Issue |
Pt 22 |
Pages |
4566-4573 |
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Keywords |
Animals; Animals, Wild/*physiology; Body Temperature; Body Temperature Regulation; Eating; *Energy Metabolism; Female; Heart Rate; Horses/*physiology; Male; Motor Activity; Pregnancy; Reproduction; *Seasons |
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Abstract |
Many large mammals show pronounced seasonal fluctuations of metabolic rate (MR). It has been argued, based on studies in ruminants, that this variation merely results from different levels of locomotor activity (LA), and heat increment of feeding (HI). However, a recent study in red deer (Cervus elaphus) identified a previously unknown mechanism in ungulates--nocturnal hypometabolism--that contributed significantly to reduced energy expenditure, mainly during late winter. The relative contribution of these different mechanisms to seasonal adjustments of MR is still unknown, however. Therefore, in the study presented here we quantified for the first time the independent contribution of thermoregulation, LA and HI to heart rate (f(H)) as a measure of MR in a free-roaming large ungulate, the Przewalski horse or Takhi (Equus ferus przewalskii Poljakow). f(H) varied periodically throughout the year with a twofold increase from a mean of 44 beats min(-1) during December and January to a spring peak of 89 beats min(-1) at the beginning of May. LA increased from 23% per day during December and January to a mean level of 53% per day during May, and declined again thereafter. Daily mean subcutaneous body temperature (T(s)) declined continuously during winter and reached a nadir at the beginning of April (annual range was 5.8 degrees C), well after the annual low of air temperature and LA. Lower T(s) during winter contributed considerably to the reduction in f(H). In addition to thermoregulation, f(H) was affected by reproduction, LA, HI and unexplained seasonal variation, presumably reflecting to some degree changes in organ mass. The observed phase relations of seasonal changes indicate that energy expenditure was not a consequence of energy uptake but is under endogenous control, preparing the organism well in advance of seasonal energetic demands. |
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Research Institute of Wildlife Ecology, University of Veterinary Medicine, Vienna, Savoyenstrasse 1, 1160 Vienna, Austria. walter.arnold@vu-wien.ac.at |
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0022-0949 |
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PMID:17079726 |
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1782 |
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Bennett, A.T. |
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Title |
Do animals have cognitive maps? |
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Journal Article |
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Year |
1996 |
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The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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199 |
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Pt 1 |
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219-224 |
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Animals; Cognition/*physiology; Humans; Space Perception/*physiology; Visual Pathways |
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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. |
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Department of Pure Mathematics, University of Adelaide, Australia |
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0022-0949 |
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PMID:8576693 |
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Equine Behaviour @ team @ |
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2756 |
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Bobbert, M.F.; Alvarez, C.B.G.; van Weeren, P.R.; Roepstorff, L.; Weishaupt, M.A. |
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Title |
Validation of vertical ground reaction forces on individual limbs calculated from kinematics of horse locomotion |
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Journal Article |
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Year |
2007 |
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The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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210 |
Issue |
Pt 11 |
Pages |
1885-1896 |
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The purpose of this study was to determine whether individual limb forces could be calculated accurately from kinematics of trotting and walking horses. We collected kinematic data and measured vertical ground reaction forces on the individual limbs of seven Warmblood dressage horses, trotting at 3.4 m s(-1) and walking at 1.6 m s(-1) on a treadmill. First, using a segmental model, we calculated from kinematics the total ground reaction force vector and its moment arm relative to each of the hoofs. Second, for phases in which the body was supported by only two limbs, we calculated the individual reaction forces on these limbs. Third, we assumed that the distal limbs operated as linear springs, and determined their force-length relationships using calculated individual limb forces at trot. Finally, we calculated individual limb force-time histories from distal limb lengths. A good correspondence was obtained between calculated and measured individual limb forces. At trot, the average peak vertical reaction force on the forelimb was calculated to be 11.5+/-0.9 N kg(-1) and measured to be 11.7+/-0.9 N kg(-1), and for the hindlimb these values were 9.8+/-0.7 N kg(-1) and 10.0+/-0.6 N kg(-1), respectively. At walk, the average peak vertical reaction force on the forelimb was calculated to be 6.9+/-0.5 N kg(-1) and measured to be 7.1+/-0.3 N kg(-1), and for the hindlimb these values were 4.8+/-0.5 N kg(-1) and 4.7+/-0.3 N kg(-1), respectively. It was concluded that the proposed method of calculating individual limb reaction forces is sufficiently accurate to detect changes in loading reported in the literature for mild to moderate lameness at trot. |
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Institute for Fundamental and Clinical Human Movement Sciences, Vrije Universiteit, van der Boechorstraat 9, NL-1081 BT Amsterdam, The Netherlands |
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0022-0949 |
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PMID:17515415 |
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Equine Behaviour @ team @ |
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3700 |
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Author |
Bobbert, M.F.; Santamaria, S. |
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Title |
Contribution of the forelimbs and hindlimbs of the horse to mechanical energy changes in jumping |
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Journal Article |
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Year |
2005 |
Publication |
The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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208 |
Issue |
2 |
Pages |
249-260 |
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Animals; Biomechanics; Forelimb/*physiology; Hindlimb/*physiology; Horses/*physiology; Locomotion/*physiology; Muscle, Skeletal/*physiology; Time Factors |
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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). |
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Institute for Fundamental and Clinical Human Movement Sciences, Vrije Universiteit, van der Boechorstraat 9, NL-1081 BT Amsterdam, The Netherlands. MFBobbert@fbw.vu.nl |
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0022-0949 |
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PMID:15634844 |
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1895 |
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Author |
Dutto, D.J.; Hoyt, D.F.; Clayton, H.M.; Cogger, E.A.; Wickler, S.J. |
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Title |
Moments and power generated by the horse (Equus caballus) hind limb during jumping |
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Journal Article |
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Year |
2004 |
Publication |
The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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207 |
Issue |
Pt 4 |
Pages |
667-674 |
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Animals; Biomechanics; Hindlimb/*physiology; Horses/*physiology; Locomotion/*physiology |
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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. |
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Department of Kinesiology and Health Promotion, California State Polytechnic University, Pomona, CA 91768, USA. ddutto@csupomona.edu |
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0022-0949 |
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PMID:14718509 |
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Equine Behaviour @ team @ |
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3654 |
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Etienne, A.S.; Maurer, R.; Seguinot, V. |
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Title |
Path integration in mammals and its interaction with visual landmarks |
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Journal Article |
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Year |
1996 |
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The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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199 |
Issue |
Pt 1 |
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201-209 |
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Animals; Cognition/physiology; Cricetinae; Gerbillinae; Humans; Locomotion/*physiology; Mammals/*physiology; Mesocricetus; Mice; Proprioception/physiology; Rats; Visual Pathways/*physiology; Visual Perception/*physiology |
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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. |
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Laboratoire d'Ethologie, FPSE, Universite de Geneve, Carouge, Switzerland |
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0022-0949 |
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PMID:8576691 |
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Equine Behaviour @ team @ |
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2758 |
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Gallistel, C.R.; Cramer, A.E. |
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Computations on metric maps in mammals: getting oriented and choosing a multi-destination route |
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Journal Article |
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1996 |
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The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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199 |
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Pt 1 |
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211-217 |
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Animals; Brain/physiology; Cercopithecus aethiops; Cognition/*physiology; Humans; Mammals/*physiology; Movement; Orientation/*physiology; Rats; Space Perception; Visual Pathways/*physiology |
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The capacity to construct a cognitive map is hypothesized to rest on two foundations: (1) dead reckoning (path integration); (2) the perception of the direction and distance of terrain features relative to the animal. A map may be constructed by combining these two sources of positional information, with the result that the positions of all terrain features are represented in the coordinate framework used for dead reckoning. When animals need to become reoriented in a mapped space, results from rats and human toddlers indicate that they focus exclusively on the shape of the perceived environment, ignoring non-geometric features such as surface colors. As a result, in a rectangular space, they are misoriented half the time even when the two ends of the space differ strikingly in their appearance. In searching for a hidden object after becoming reoriented, both kinds of subjects search on the basis of the object's mapped position in the space rather than on the basis of its relationship to a goal sign (e.g. a distinctive container or nearby marker), even though they have demonstrably noted the relationship between the goal and the goal sign. When choosing a multidestination foraging route, vervet monkeys look at least three destinations ahead, even though they are only capable of keeping a maximum of six destinations in mind at once. |
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Department of Psychology, University of California, Los Angeles 90095, USA |
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0022-0949 |
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PMID:8576692 |
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Equine Behaviour @ team @ |
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2757 |
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Kuntz, R.; Kubalek, C.; Ruf, T.; Tataruch, F.; Arnold, W. |
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Seasonal adjustment of energy budget in a large wild mammal, the Przewalski horse (Equus ferus przewalskii) I. Energy intake |
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Journal Article |
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2006 |
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The Journal of experimental biology |
Abbreviated Journal |
J Exp Biol |
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209 |
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22 |
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4557-4565 |
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Large ruminants respond to changing plant phenology during winter by decreasing voluntary food intake, increasing gut passage time and utilizing body fat reserves. It is uncertain, however, how other large mammals with a non-ruminant digestive physiology cope with winter forage conditions. Therefore, we investigated seasonality of energy intake in a large herbivorous wild mammal, the Przewalski horse (Equus ferus przewalskii). Throughout all seasons we used the n-alkane method to measure daily dry matter intake (DMI), diet composition and digestion, and determined an index of gut passage time in horses living under close to natural conditions. DMI correlated positively with its content of crude protein and nitrogen-free extract. Independent of these effects, DMI further varied seasonally with a peak in autumn and a nadir in late winter. Fluctuations of DMI corresponded to the annual change in body condition, which decreased during winter while energy reserves were depleted, and increased during the fattening period. Gut passage time varied in the course of the year and was longer during winter when the diet was high in crude fibre. Nevertheless, changes in gut passage time occurred rather independently of changes in forage composition and DMI, suggesting endogenous control for timely adaption of the digestive strategy to meet predictable changes in forage quality. |
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10.1242/jeb.02535 |
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Equine Behaviour @ team @ |
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4729 |
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McGuigan, M.P.; Wilson, A.M. |
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The effect of gait and digital flexor muscle activation on limb compliance in the forelimb of the horse Equus caballus |
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Journal Article |
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2003 |
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The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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206 |
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Pt 8 |
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1325-1336 |
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Animals; Biomechanics; Forelimb/anatomy & histology/*physiology; Gait/*physiology; Horses/anatomy & histology/*physiology; Muscle Contraction/*physiology; Running |
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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. |
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Structure and Motion Laboratory, Veterinary Basic Sciences, The Royal Veterinary College, Hawkshead Lane, North Mymms, Hatfield, Hertfordshire AL9 7TA, UK. m.p.mcguigan@leeds.ac.uk |
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PMID:12624168 |
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Equine Behaviour @ team @ |
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3655 |
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Skedros, J.G.; Dayton, M.R.; Sybrowsky, C.L.; Bloebaum, R.D.; Bachus, K.N. |
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The influence of collagen fiber orientation and other histocompositional characteristics on the mechanical properties of equine cortical bone |
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Journal Article |
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2006 |
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The Journal of Experimental Biology |
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J Exp Biol |
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209 |
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Pt 15 |
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3025-3042 |
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Animals; Biomechanics; Bone and Bones/*physiology; Collagen/*physiology; Forelimb; Horses/*physiology |
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Abstract |
This study examined relative influences of predominant collagen fiber orientation (CFO), mineralization (% ash), and other microstructural characteristics on the mechanical properties of equine cortical bone. Using strain-mode-specific (S-M-S) testing (compression testing of bone habitually loaded in compression; tension testing of bone habitually loaded in tension), the relative mechanical importance of CFO and other material characteristics were examined in equine third metacarpals (MC3s). This model was chosen since it had a consistent non-uniform strain distribution estimated by finite element analysis (FEA) near mid-diaphysis of a thoroughbred horse, net tension in the dorsal/lateral cortices and net compression in the palmar/medial cortices. Bone specimens from regions habitually loaded in tension or compression were: (1) tested to failure in both axial compression and tension in order to contrast S-M-S vs non-S-M-S behavior, and (2) analyzed for CFO, % ash, porosity, fractional area of secondary osteonal bone, osteon cross-sectional area, and population densities of secondary osteons and osteocyte lacunae. Multivariate multiple regression analyses revealed that in S-M-S compression testing, CFO most strongly influenced total energy (pre-yield elastic energy plus post-yield plastic energy); in S-M-S tension testing CFO most strongly influenced post-yield energy and total energy. CFO was less important in explaining S-M-S elastic modulus, and yield and ultimate stress. Therefore, in S-M-S loading CFO appears to be important in influencing energy absorption, whereas the other characteristics have a more dominant influence in elastic modulus, pre-yield behavior and strength. These data generally support the hypothesis that differentially affecting S-M-S energy absorption may be an important consequence of regional histocompositional heterogeneity in the equine MC3. Data inconsistent with the hypothesis, including the lack of highly longitudinal collagen in the dorsal-lateral ;tension' region, paradoxical histologic organization in some locations, and lack of significantly improved S-M-S properties in some locations, might reflect the absence of a similar habitual strain distribution in all bones. An alternative strain distribution based on in vivo strain measurements, without FEA, on non-Thoroughbreds showing net compression along the dorsal-palmar axis might be more characteristic of the habitual loading of some of the bones that we examined. In turn, some inconsistencies might also reflect the complex torsion/bending loading regime that the MC3 sustains when the animal undergoes a variety of gaits and activities, which may be representative of only a portion of our animals, again reflecting the possibility that not all of the bones examined had similar habitual loading histories. |
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Utah Bone and Joint Center, 5323 S. Woodrow Street #202, Salt Lake City, UT 84107, USA. jskedros@utahboneandjoint.com |
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English |
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0022-0949 |
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PMID:16857886 |
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1868 |
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