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PubMed1998

Recovery of locomotion after ventral and ventrolateral spinal lesions in the cat. I. Deficits and adaptive mechanisms.

The recovery of treadmill locomotion of eight adult cats, subjected to chronic ventral and ventrolateral spinal lesions at low thoracic levels (T11 or T13), preserving at least one dorsolateral funiculus and the dorsal columns, was documented daily using electromyographic (EMG) and kinematic methods. The data show that all cats eventually recovered quadrupedal voluntary locomotion despite extensive damage to important pathways (such as the reticulospinal and the vestibulospinal) as verified by injection of wheat germ agglutinin-horseradish peroxidase (WGA-HRP) caudal to the site of lesion. Initially (in the early period after the spinal lesion), all the cats suffered from pronounced locomotor and postural deficits, and they could not support their hindquarters or walk with their hindlimbs. Gradually, during the recovery period, they regained quadrupedal walking, although their locomotion was wobbly and inconsistent, and they suffered from poor lateral stability. EMG and kinematic data analyses showed a tendency for an increase in the variability of the step cycle duration but no major changes in the step cycle structure or in the intralimb coupling of the joints. However, the homolateral fore- and hindlimb coupling was highly perturbed in cats with the largest lesions. Although the general alternating pattern of extensor and flexors was maintained, there were various changes in the duration and amplitude of the EMG bursts as well as a lack of amplitude modulation during walking uphill or downhill on the treadmill. In cats with larger lesions, the forelimbs also seem to take a greater propulsive role than usual as revealed by a consistent increase of the activity of the triceps. In cats with smaller lesions, these deficits were transient, but, for the most extensively lesioned cats, they were pronounced and lasted long term postlesion even after reaching a more or less stable locomotor behavior (plateau period). It is concluded that recovery of quadrupedal locomotion is possible even after a massive lesion to ventral and ventrolateral quadrants, severing the vestibulospinal pathway and causing severe, although incomplete, damage to the reticulospinal tract. The quick recovery in the less lesioned cats can be attributed to remaining pathways normally implicated in locomotor function. However, in the most extensively lesioned cats, the long period of recovery and the pronounced deficits during the plateau period may indicate that the compensation, attributed to remaining reticulospinal pathways, is not sufficient and that other pathways in the dorsolateral funiculi, such as the corticospinal, can sustain and adapt, up to a certain extent, the voluntary quadrupedal walking.

Animals↗
PubMed1976

Locomotion and posture of the Malayan siamang and implications for hominoid evolution.

Wild, adult siamang were observed for over 800 h in lowland dipterocarp forest in the Krau Game Reserve, Pahang, West Malaysia. Siamang use four patterns of locomotion: brachiation, climbing, bipedalism and leaping. The pattern of locomotion used by the siamang varies with the size of arboreal supports and with major behavioral activity. Travel is primarily by brachiation along large boughs. Locomotion during feeding is primarily climbing among small branches. In feeding, siamang use suspensory postures among small supports and seated postures on large supports. Comparison of siamang locomotion and posture with that of other apes suggest that quadramanous climbing during feeding is the basic hominoid locomotor adaptation.

Animals↗
PubMed1986

Locomotion and feeding postures of spider and howling monkeys: field study and evolutionary interpretation.

Field observations demonstrate clear differences in locomotion and feeding postures between spider monkeys (Ateles) and howling monkeys (Alouatta). When feeding, Ateles employs sitting postures approximately half the time, and a variety of suspensory postures using the tail the other half. Ateles moves quadrupedally during 52% of locomotion, by tail-arm suspension 25%, and various mixed support-suspensory modes the remainder. Tail-arm suspension is practiced more rapidly on thinner supports, and on more negatively inclined supports than is quadrupedal movement. Howlers do not locomote by tail-arm suspension: movement is almost entirely quadrupedal and is slower than that of spider monkeys. The positional behavior of spider monkeys fits closely recent views of major adaptive changes in hominoid evolution emphasizing brachiation and speed during travel. Howler locomotion and also tissue composition appear related to diet and digestive mechanisms.

Alouatta↗
PubMed1989

Biomechanics and allometric scaling in primate locomotion and morphology.

Body size has a dominant influence on locomotor performance and the morphology of the locomotor apparatus. In locomotion under the influence of gravity, body mass acts as weight force and is a mechanical variable. Accordingly, the application of biomechanical principles and methods allows a functional understanding of scaling effects in locomotion. This is demonstrated here using leaping primates as an example. With increasing body size, the decreasing ratio of muscle force available for acceleration during takeoff to the body mass that has to be accelerated dictates both the movement pattern and the proportions of the hindlimbs. In an arm-swinging movement, the long, heavy arms of the large-bodied leapers are effectively used to gain additional momentum. A new perspective on decreasing size identifies the absolutely small acceleration distance and time available for propulsion as factors limiting leaping distance and extensively determining locomotor behavior and body proportions. As the mechanical constraints differ according to body size for a given mode of locomotion, a typological approach to morphology in relation to locomotor category is ruled out. Across locomotor categories, dynamic similarity (sensu Alexander) can be expected if the propulsive mechanisms as well as the selective pressures acting upon locomotion are the same.

Animals↗
PubMed2003

Exotendons for assistance of human locomotion.

BACKGROUND: Powered robotic exoskeletons for assistance of human locomotion are currently under development for military and medical applications. The energy requirements for such devices are excessive, and this has become a major obstacle for practical applications. Legged locomotion in many animals, however, is very energy efficient. We propose that poly-articular elastic mechanisms are a major contributor to the economy of locomotion in such specialized animals. Consequently, it should be possible to design unpowered assistive devices that make effective use of similar mechanisms. METHODS: A passive assistive technology is presented, based on long elastic cords attached to an exoskeleton and guided by pulleys placed at the joints. A general optimization procedure is described for finding the best geometrical arrangement of such "exotendons" for assisting a specific movement. Optimality is defined either as minimal residual joint moment or as minimal residual joint power. Four specific exotendon systems with increasing complexity are considered. Representative human gait data were used to optimize each of these four systems to achieve maximal assistance for normal walking. RESULTS: The most complex exotendon system, with twelve pulleys per limb, was able to reduce the joint moments required for normal walking by 71% and joint power by 74%. A simpler system, with only three pulleys per limb, could reduce joint moments by 46% and joint power by 47%. CONCLUSION: It is concluded that unpowered passive elastic devices can substantially reduce the muscle forces and the metabolic energy needed for walking, without requiring a change in movement. When optimally designed, such devices may allow independent locomotion in patients with large deficits in muscle function.

Ankle Joint↗
PubMed2003

Whole-body mechanics and gaits in the gray short-tailed opossum Monodelphis domestica: integrating patterns of locomotion in a semi-erect mammal.

Gaits (footfall patterns) and external mechanical energy patterns of the center of mass were quantified in a generalized, semi-erect mammal in order to address three general questions. First, do semi-erect mammals exhibit the walk/run gait transitions that have been proposed as the primitive condition for tetrapods? Second, do small, semi-erect mammals employ the energy-saving pendular and spring-based mechanics used by erect mammals? Third, how well do mechanical locomotor patterns of the center of mass correlate with gaits? Monodelphis domestica utilizes only fast walking and running trot gaits over a fivefold increase in speed, over which we could illicit constant velocity steps, although running trots were their preferred gait. In sustained level locomotion the opossums did not use other walking gaits presumed to be primitive for tetrapods. Across the full range of speeds their trotting gaits exhibited force patterns and in-phase mechanical energy fluctuations that are characteristic of spring-mass mechanics. Thus, opossums appear to prefer trotting gaits with bouncing mechanics for sustained locomotion. Integration of center-of-mass versus footfall perspectives reveals that spring-mass mechanics is associated with both walking trot and running trot gaits. Furthermore, the onset of an aerial phase was not clearly associated with either the walk/run gait transition (50% duty factor) or a change in center-of-mass mechanics. The assumption that energy-saving mechanisms are ubiquitous among mammals is tenuous because small non-cursorial mammals do not appear to use pendular-based mechanics for sustained locomotion and, although they prefer spring-based mechanics, they probably lack clear musculoskeletal spring elements that could store energy during running. Thus, it appears that simply paying for locomotion with muscular work may be the primitive condition for mammals.

Animals↗
PubMed2003

Muscle force-length dynamics during level versus incline locomotion: a comparison of in vivo performance of two guinea fowl ankle extensors.

For a terrestrial animal to move in the complex natural environment, the limb muscles must modulate force and work performance to meet changing mechanical requirements; however, it is not clear whether this is accomplished via a collective shift in function by all limb muscles, or a division of labor among limb muscles. Do muscles differ in their ability to modulate force-length contractile function to meet the mechanical demands of different locomotor tasks? We explore this question by examining the in vivo force-length performance of the guinea fowl Numida meleagris lateral gastrocnemius (LG) and digital flexor-IV (DF-IV), during level and incline locomotion. During level locomotion, the LG and DF-IV exhibit differing muscle fascicle strain patterns: the LG shortens by 10-15% while developing force, whereas the DF-IV undergoes a stretch-shorten cycle with large strain amplitudes and small net strains of 1-8%. Furthermore, the DF-IV operates at higher muscle stresses (92-130 kPa, compared to 23-39 kPa for LG) and possesses a longer tendon, which allows the DF-IV tendon to recover greater elastic energy than the LG tendon. During incline locomotion, these muscles contribute only one-third of the energy expected for their mass, with the DF-IV exhibiting high stride-to-stride variability in work output. While the stretch-shorten cycle of the DF-IV muscle may allow more economic force production, it also leads to large changes in work output with small changes in the relative timing of force and strain. Thus, while the primary determinants of LG work are net strain and mean force, the primary determinant of DF-IV work is the phase relationship between force and strain. Our results suggest that, in addition to influencing a muscle's mechanical performance during steady level locomotion, morphology also affects its capacity and mechanism for altering work output for different locomotor tasks.

Analysis of Variance↗
PubMed1985

Metabolic indicators of fibre recruitment in mammalian muscles during locomotion.

Fast-twitch-oxidative-glycolytic (FOG), fast-twitch-glycolytic (FG) and slow-twitch-oxidative (SO) fibres are distributed within and among physiological extensor muscles in mammals in predictable patterns. Deep muscles and the deep portions of extensor muscles are primarily composed of SO and FOG fibres, and the more peripheral portions of the muscles have higher concentrations of FG fibres. During terrestrial locomotion, the fibres are recruited in this same general order from postural standing through high speed running to jumping (i.e. during standing deep SO fibres are active and during locomotion there is a progressive peripheral recruitment of fibres from SO to FOG to FG). Several metabolic indicators may be used to map these fibre recruitment patterns, including glycogen loss in fibres, metabolic enzyme changes during training at different speeds, and distribution of blood flow within and among the muscles. Concerning the latter, during standing in rats blood flows in the hindlimb muscles are directly proportional to the SO fibre populations in the muscles. However, during locomotion the elevations in blood flow over pre-exercise are a function of the populations of FOG fibres in the muscles. Blood flows in the peripheral white portions (FG fibres) of extensor muscles are not significantly elevated until the rats run at high speeds, when the FG fibres presumably are recruited. During swimming, when flexor muscles are relatively more active than extensor muscles (as compared with terrestrial locomotion), blood flows in the flexors are correspondingly higher. Thus, there exists a clear 'biological economy' in the matching of blood flow to the specific fibres that are active within and among muscles during exercise.

Animals↗
PubMed1985

Force development during sustained locomotion: a determinant of gait, speed and metabolic power.

This paper develops three simple ideas about force development during sustained locomotion which provide some insights into the mechanisms that determine why animals change gait, how fast they can run, and how much metabolic energy they consume. The first idea is that the alternate stretch-shorten pattern of activity of the muscles involved in locomotion allows muscle-tendon units to function as springs, affecting the amount of force a given cross-sectional area of muscle develops, and the metabolic requirements of the muscles for force development. Animals select speeds and stride frequencies which optimize the performance of these springs. The second idea is that muscle stress (force/cross-sectional area) determines when animals change gait, how fast they run and their peak accelerations and decelerations. It is proposed that terrestrial birds and mammals develop similar muscle stresses under equivalent conditions (i.e. preferred speed within a gait) and that animals change gaits in order to reduce peak stresses as they increase speed. Finally, evidence is presented to support the idea that it is the time course of force development during locomotion, rather than the mechanical work that the muscles perform, that determines the metabolic cost of locomotion.

Animals↗
PubMed1990

Effect of variation in form on the cost of terrestrial locomotion.

The mass-specific minimum cost of terrestrial locomotion (Cmin) decreases with an increase in body mass. This generalization spans nearly eight orders of magnitude in body mass and includes two phyla. The general relationship between metabolic cost and mass is striking. However, a significant amount of unexplained interspecific variation in Cmin exists at any given body mass. To determine how variation in morphology and physiology affects metabolic energy cost, we measured the oxygen consumption of three comparably sized insects running on a miniature treadmill; the American cockroach Periplaneta americana, the caterpillar hunting beetle Calosoma affine and the Australian field cricket Teleogryllus commodus. Steady-state oxygen consumption (VO2ss) increased linearly with speed. Cmin was similar for crickets and cockroaches (8.0 and 8.5 ml O2 g-1km-1, respectively), but was substantially lower for beetles (4.6 ml O2 g-1km-1). The predicted value of Cmin for all three insects was within the 95% confidence intervals of the Cmin versus body mass function. However, the 95% confidence intervals extend approximately 2.5-fold above and 40% below the regression line, making the variation at any given body mass nearly sixfold. Normalizing for the rate of muscle force production by determining the metabolic cost per stride failed to account for the interspecific variation in the cost of locomotion observed in the three insects. Ground contact costs (i.e. VO2ss multiplied by leg contact time during a stride) in insects were similar to those measured in mammals (1.5-3.1 J kg-1) and were independent of speed, but did not explain the interspecific variation in the cost of locomotion. Muscles of the caterpillar hunting beetle may have a greater mechanical advantage than muscles of the Australian field cricket and American cockroach. Variation in musculo-skeletal arrangement, apart from variation in body mass, could translate into significant differences in the minimum cost of terrestrial locomotion.

Animals↗
PubMed1990

The work and energetic cost of locomotion. II. Partitioning the cost of internal and external work within a species.

Previous studies have shown that large animals have systematically lower mass-specific costs of locomotion than do smaller animals, in spite of there being no demonstrable difference between them in the mass-specific mechanical work of locomotion. Larger animals are somehow much more efficient at converting metabolic energy to mechanical work. The present study analyzes how this decoupling of work and cost might occur. The experimental design employs limb-loaded and back-loaded dogs and allows the energetic cost of locomotion to be partitioned between that used to move the center of mass (external work) and that used to move the limbs relative to the center of mass (internal work). These costs were measured in three dogs moving at four speeds. Increases in the cost of external work with speed parallel increases in the amount of external work based on data from previous studies. However, increases in the cost of internal work with speed are much less (less than 50%) than the increase in internal work itself over the speeds examined. Furthermore, the cost of internal work increases linearly with speed, whereas internal work itself increases as a power function of speed. It is suggested that this decoupling results from an increase with speed in the extent to which the internal work of locomotion is powered by non-metabolic means, such as elastic strain energy and transfer of energy within and between body segments.

Animals↗
PubMed1992

Scaling of muscle fibres and locomotion.

To reconcile the scaling of the mechanics and energetics of locomotion to recent data on the scaling of the mechanics of muscle fibres, I have extended the theory of Taylor and colleagues that the energetic cost of locomotion is determined by the cost of generating force by the fibres. By assuming (1) that the cost of generating force in a fibre is proportional to V(max) (maximum velocity of shortening) and (2) that, at physiologically equivalent speeds, animals of different body sizes recruit the same fibre types, this extension quantitatively predicts the scaling of the energetics of locomotion, as well as other observations, from the scaling of V(max) of the muscle fibres. First, the energetic cost of locomotion at physiologically equivalent speeds scales with Mb-0.16, where Mb is body mass, as does V(max) of a given fibre type. However, the energetic cost at absolute speeds (cost of transport) scales with Mb-0.30, because small animals must compress their recruitment order into a narrower speed range and, hence, recruit faster muscle fibre types at a given running speed. Thus, it costs more for small animals to move 1 kg of their body mass 1 km not only because a given muscle fibre type from a small animal costs more to generate force than from a large one, but also because small animals recruit faster fibre types at a given absolute running speed. In addition, this analysis provides evidence that V(max) scales similarly to 1/tc (where tc is foot contact time) and muscle shortening velocity (V), in agreement with recent models. Finally, this extension predicts that, at physiologically equivalent speeds, the weight-specific energetic cost per step is independent of body size, as has been found empirically.

Animals↗
PubMed1996

Function of the intercostal muscles in trotting dogs: ventilation or locomotion?

Although the intercostal muscles play an important role in lung ventilation, observations from fishes and ectothermic tetrapods suggest that their primary function may be locomotion. To provide a broader understanding of the role these muscles play in locomotion, I measured ventilatory airflow at the mouth and activity of the fourth and ninth intercostal muscles in four dogs trotting on a treadmill. During rest and thermoregulatory panting, activity of the intercostal muscles was associated with inspiratory and expiratory airflow. However, during trotting, activity of the interosseous portions of the intercostal muscles was correlated with locomotion. When ventilation and stride cycles were not synchronized, activity of the interosseous intercostal muscles stayed locked to the locomotor events and drifted in time relative to ventilation. In contrast, activity of the parasternal portion of the internal intercostal muscles was always associated with inspiratory airflow. These observations suggest that, in dogs, locomotion is the dominant function of the interosseous portions of the intercostal muscles. However, the parasternal intercostal muscles are primarily inspiratory in function.

Animals↗
PubMed1999

Dynamics of muscle function during locomotion: accommodating variable conditions.

Much of what we know about animal locomotion is derived from studies examining animals moving within a single, homogeneous environment, at a steady speed and along a flat grade. As a result, the issue of how musculoskeletal function might shift to accommodate variability within the external environment has remained relatively unexplored. One possibility is that locomotor muscles are differentially recruited depending upon the environment in which the animal is moving. A second possibility is that the same muscles are recruited, but that they are activated in a different manner so that their contractile function differs according to environment. Finally, it is also possible that, in some cases, animals may not need to alter their musculoskeletal function to move under different external conditions. In this case, however, the mechanical behavior appropriate for one environmental condition may constrain locomotor performance in another. To begin to explore the means by which animals accommodate variable conditions in their environment, we present three case studies examining how musculoskeletal systems function to allow locomotion under variable conditions: (1) eels undulating through water and across land, (2) turkeys running on level and inclined surfaces, and (3) ducks using their limbs to walk and to paddle. In all three of these examples, the mechanical behavior of some muscle(s) involved in locomotion are altered, although to different degrees and in different ways. In the running turkeys, the mechanical function of a major ankle extensor muscle shifts from contracting isometrically on a flat surface (producing little work and power), to shortening actively during contraction on an uphill gradient (increasing the amount of work and power generated). In the ducks, the major ankle extensor undergoes the same general pattern of activation and shortening in water and on land, except that the absolute levels of muscle stress and strain and work output are greater during terrestrial locomotion. In eels, a transition to land elicits changes in electromyographic duty cycles and the relative timing of muscle activation, suggesting some alteration in the functional mechanics of the underlying musculature. Only by studying muscle function in animals moving under more variable conditions can we begin to characterize the functional breadth of the vertebrate musculoskeletal system and understand more fully its evolutionary design.

Animals↗
PubMed1999

Energetics of locomotion by the Australian water rat (Hydromys chrysogaster): a comparison of swimming and running in a semi-aquatic mammal.

Semi-aquatic mammals occupy a precarious evolutionary position, having to function in both aquatic and terrestrial environments without specializing in locomotor performance in either environment. To examine possible energetic constraints on semi-aquatic mammals, we compared rates of oxygen consumption for the Australian water rat (Hydromys chrysogaster) using different locomotor behaviors: swimming and running. Aquatic locomotion was investigated as animals swam in a water flume at several speeds, whereas water rats were run on a treadmill to measure metabolic effort during terrestrial locomotion. Water rats swam at the surface using alternate pelvic paddling and locomoted on the treadmill using gaits that included walk, trot and half-bound. Water rats were able to run at twice their maximum swimming velocity. Swimming metabolic rate increased with velocity in a pattern similar to the 'humps' and 'hollows' for wave drag experienced by bodies moving at the water surface. Metabolic rate increased linearly during running. Over equivalent velocities, the metabolic rate for running was 13-40 % greater than for swimming. The minimum cost of transport for swimming (2.61 J N-1 m-1) was equivalent to values for other semi-aquatic mammals. The lowest cost for running (2.08 J N-1 m-1) was 20 % lower than for swimming. When compared with specialists at the extremes of the terrestrial-aquatic continuum, the energetic costs of locomoting either in water or on land were high for the semi-aquatic Hydromys chrysogaster. However, the relative costs for H. chrysogaster were lower than when an aquatic specialist attempts to move on land or a terrestrial specialist attempts to swim.

Animals↗
PubMed2001

Dynamics of mallard (Anas platyrynchos) gastrocnemius function during swimming versus terrestrial locomotion.

This study investigates how the contractile function of a muscle may be modulated to accommodate changes in locomotor mode and differences in the physical environment. In vivo recordings of lateral gastrocnemius (LG) activation, force development (measured using tendon buckle transducers) and length change (measured using sonomicrometry) were obtained from mallard ducks (Anas platyrhynchos) as they swam at steady speeds in a water tank and walked or ran on land. LG force recordings were compared with combined lateral and medial gastrocnemius (MG) muscle-tendon force recordings obtained from the contralateral limb, allowing force development by the MG to be estimated relative to that of the LG. Although similar stresses were calculated to act in the LG and MG muscles during terrestrial locomotion (126 and 115 kPa, respectively), stresses were considerably greater in the LG compared with the MG during swimming (62 versus 34 kPa, respectively). During both steady swimming and terrestrial locomotion, the LG developed force while shortening over a considerable range of its length (swimming 23.6 % versus terrestrial 37.4 %). Activation of the muscle occurred near the end of passive lengthening during the recovery stroke, just prior to muscle shortening. As a result, the muscle generated broad positive work loops during both locomotor modes. LG work during swimming (4.8 J x kg(-1)) averaged 37 % of the work performed during terrestrial locomotion (13.1 J x kg(-1)), consistent with the twofold greater force and 58 % greater strain of the muscle during walking and running. Because limb cycle frequency was similar for the two locomotor modes (swimming 2.65 versus terrestrial 2.61 Hz), differences in power output (swimming 12.6 W x kg(-1 )versus terrestrial 32.4 W x kg(-1)) largely reflected difference in work per cycle. Tendon elastic energy savings was a small fraction (<5 %) of the work performed by the muscle, consistent with a fiber-tendon design of these two muscles that favors muscle work to produce limb movement with little tendon strain. These results are consistent with a higher cost of terrestrial locomotion in ducks compared with other, more cursorial birds that may operate their muscles more economically and achieve greater tendon elastic savings.

Animals↗
PubMed2001

Energetics of terrestrial locomotion of the platypus Ornithorhynchus anatinus.

The platypus Ornithorhynchus anatinus Shaw displays specializations in its limb structure for swimming that could negatively affect its terrestrial locomotion. Platypuses walked on a treadmill at speeds of 0.19-1.08 m x s(-1). Video recordings were used for gait analysis, and the metabolic rate of terrestrial locomotion was studied by measuring oxygen consumption. Platypuses used walking gaits (duty factor >0.50) with a sprawled stance. To limit any potential interference from the extensive webbing on the forefeet, platypuses walk on their knuckles. Metabolic rate increased linearly over a 2.4-fold range with increasing walking speed in a manner similar to that of terrestrial mammals, but was low as a result of the relatively low standard metabolic rate of this monotreme. The dimensionless cost of transport decreased with increasing speed to a minimum of 0.79. Compared with the cost of transport for swimming, the metabolic cost for terrestrial locomotion was 2.1 times greater. This difference suggests that the platypus may pay a price in terrestrial locomotion by being more aquatically adapted than other semi-aquatic or terrestrial mammals.

Adaptation, Biological↗
PubMed2001

Effects of serotonin, dopamine and ergometrine on locomotion in the pulmonate mollusc Helix lucorum.

The terrestrial snail Helix lucorum crawls using waves of muscular contraction (pedal waves) that spread along the sole of its foot. Crawling speed depends on wave generation frequency (step frequency) and the distance the snail moves forwards during each wave (step length). In a previous study, video recordings of a crawling snail showed that its sole length varied over a wide range and was directly correlated with mollusc speed. Speed depended on step length, which was directly related to sole length, rather than on step frequency, which remained rather constant. In the present study, the effects of dopamine, ergometrine (a blocker of dopamine receptors in molluscs) and serotonin injection on the linear relationship between sole length and locomotor speed in Helix lucorum were studied. In crawling snails, dopamine caused sole contraction, and locomotion slowed down or ceased. Ergometrine stimulated locomotion, which resembled rapid crawling with an extended sole, as observed under normal conditions. Serotonin stimulated locomotion and accelerated crawling significantly without causing changes in sole length. The acceleration of locomotion induced by serotonin injection was due to pedal wave (step) elongation. It is proposed that, during each locomotor episode, dopamine controls snail speed by regulating sole length, which determines the amplitude of contraction of the muscle cells involved in pedal waves and, as a result, step length; serotonin determines the basic step length and shifts the linear relationship between sole length and mollusc speed upwards along the axis of mollusc speed. The efficiency of the serotonergic system depends on the physiological state of the mollusc (e.g. that characteristic of summer or winter).

Animals↗