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Brinkmann, L., Gerken, M., Hambly, C., Speakman, J. R., & Riek, A. (2014). Saving energy during hard times: Energetic adaptations of Shetland pony mares. J. Exp. Biol., 217, 4320–4327.
Abstract: Recent results suggest that wild Northern herbivores reduce their metabolism during times of low ambient temperatures and food shortage in order to reduce their energetic needs. It is however not known if domesticated animals are also able to reduce their energy expenditure. We exposed ten Shetland pony mares to different environmental conditions (summer and winter) and to two food quantities (60 and 100% of maintenance energy requirement, respectively) during low winter temperatures to examine energetic and behavioural responses. In summer ponies showed a considerably higher field metabolic rate (FMR) (63.4±15.0 MJ d-1) compared to restrictively fed and control animals in winter (24.6±7.8 MJ d-1 and 15.0±1.1 MJ d-1, respectively). During summer conditions locomotor activity, resting heart rates and total water turnover were considerably elevated (P<0.001) compared to winter. Restrictively fed animals (N=5) compensated for the decreased energy supply by reducing their FMR by 26% compared to control animals (N=5). Furthermore, resting heart rate, body mass and body condition score were lower (29.2±2.7 beats min-1; 140±22 kg; 3.0±1.0 points) than in control animals (36.8±41 beats min-1; 165 ±31 kg; 4.4±0.7 points; P<0.05). While the observed behaviour did not change, nocturnal hypothermia was elevated. We conclude that ponies acclimatize to different climatic conditions by changing their metabolic rate, behaviour and some physiological parameters. When exposed to energy challenges, ponies, like wild herbivores, exhibited hypometabolism and nocturnal hypothermia.
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Roth, L. S. V., Balkenius, A., & Kelber, A. (2007). Colour perception in a dichromat. Journal of Experimental Biology, 210(16), 2795–2800.
Abstract: Most mammals have dichromatic colour vision based on two different types of cones: a short-wavelength-sensitive cone and a long-wavelength-sensitive cone. Comparing the signal from two cone types gives rise to a one-dimensional chromatic space when brightness is excluded. The so-called `neutral point' refers to the wavelength that the animal cannot distinguish from achromatic light such as white or grey because it stimulates both cone types equally. The question is: how do dichromats perceive their chromatic space? Do they experience a continuous scale of colours or does the neutral point divide their chromatic space into two colour categories, i.e. into colours of either short or long wavelengths?We trained horses to different colour combinations in a two-choice behavioural experiment and tested their responses to the training and test colours. The horses chose colours according to their similarity/relationship to rewarded and unrewarded training colours. There was no evidence for a categorical boundary at the neutral point or elsewhere.This study suggests that dichromats perceive their chromatic space as a continuous scale of colours, treating the colour at the neutral point as any other colour they can distinguish.
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Kuntz, R., Kubalek, C., Ruf, T., Tataruch, F., & Arnold, W. (2006). Seasonal adjustment of energy budget in a large wild mammal, the Przewalski horse (Equus ferus przewalskii) I. Energy intake. J Exp Biol, 209(22), 4557–4565.
Abstract: 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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Bobbert, M. F., Alvarez, C. B. G., van Weeren, P. R., Roepstorff, L., & Weishaupt, M. A. (2007). Validation of vertical ground reaction forces on individual limbs calculated from kinematics of horse locomotion. J Exp Biol, 210(Pt 11), 1885–1896.
Abstract: 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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Witte, T. H., Knill, K., & Wilson, A. M. (2004). Determination of peak vertical ground reaction force from duty factor in the horse (Equus caballus). J Exp Biol, 207(Pt 21), 3639–3648.
Abstract: Measurement of peak vertical ground reaction force (GRFz) from multiple limbs simultaneously during high-speed, over-ground locomotion would enhance our understanding of the locomotor mechanics of cursorial animals. Here, we evaluate the accuracy of predicting peak GRFz from duty factor (the proportion of the stride for which the limb is in contact with the ground). Foot-mounted uniaxial accelerometers, combined with UHF FM telemetry, are shown to be practical and accurate for the field measurement of stride timing variables, including duty factor. Direct comparison with the force plate produces a mean error of 2.3 ms and 3.5 ms for the timing of foot on and foot off, respectively, across all gaits. Predictions of peak GRFz from duty factor show mean errors (with positive values indicating an overestimate) of 0.8+/-0.04 N kg(-1) (13%; N=42; mean +/- S.E.M.) at walk, -0.3+/-0.06 N kg(-1) (3%; N=75) at trot, -2.3+/-0.27 N kg(-1) (16%; N=18) for the non-lead limb at canter and +2.1+/-0.7 N kg(-1) (19%; N=9) for the lead limb at canter. The substantial over- and underestimate seen at canter, in the lead and non-lead limbs, respectively, is attributed to the different functions performed by the two limbs in the asymmetrical gaits. The difference in load experienced by the lead and non-lead limbs decreased with increasing speed.
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McGuigan, M. P., & Wilson, A. M. (2003). The effect of gait and digital flexor muscle activation on limb compliance in the forelimb of the horse Equus caballus. J Exp Biol, 206(Pt 8), 1325–1336.
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.
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Dutto, D. J., Hoyt, D. F., Clayton, H. M., Cogger, E. A., & Wickler, S. J. (2004). Moments and power generated by the horse (Equus caballus) hind limb during jumping. J Exp Biol, 207(Pt 4), 667–674.
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.
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Etienne, A. S., Maurer, R., & Seguinot, V. (1996). Path integration in mammals and its interaction with visual landmarks. J Exp Biol, 199(Pt 1), 201–209.
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.
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Gallistel, C. R., & Cramer, A. E. (1996). Computations on metric maps in mammals: getting oriented and choosing a multi-destination route. J Exp Biol, 199(Pt 1), 211–217.
Abstract: 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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Bennett, A. T. (1996). Do animals have cognitive maps? J Exp Biol, 199(Pt 1), 219–224.
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.
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