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PubMed1999

Behavior of hindbrain neurons during the transition from rest to evoked locomotion in a newt.

Trains of electrical stimuli were delivered to the mesencephalic 'locomotor region' in the rough skin newt. The current (3-12 mcA) and the interstimulus interval (100 to 200 ms) were adjusted so that locomotion arose in approximately 10 s, or so that the train remained subthreshold for initiation of locomotion. Impulses of single neurons in the hindbrain were recorded during the transition period from rest to locomotion. Time-locked synaptic responses were bi- or unimodal with typical latencies close to 18, 23 or 28 ms, and weak irregular mode near 13 ms. Impulses that were not locked to the stimuli arose in some silent neurons, and the rate of firing of neurons with background discharge was sometimes enhanced. Composite responses consisted of both time-locked component and impulses distributed throughout the interstimulus interval. The data suggest that short-lived, wave-like propagation of the input volley ceases or is transformed into asynchronous activity after three or four translations. The latter variant could occur if the train reached the threshold for initiation of locomotion. The asynchronous activity persisted throughout interstimulus interval and could coexist with time-locked impulses. Some neurons generated only a few impulses, while others remained active from beginning to end of the train. These active neurons could either spike at a steady rate, or decrement or augment their rate of firing during the train. The time course of their activity was related to the initial rate of firing. The augmenting type of firing in a subset of neurons may arise due to the interaction of neurons with unstable, steady state and decrementing activity.

Action Potentials↗
PubMed2002

Development of posture and locomotion: an interplay of endogenously generated activities and neurotrophic actions by descending pathways.

The adult pattern of locomotion is observed at the end of the second postnatal week in the rat. The in vitro spinal cord isolated from immature rats has served as a valuable preparation to study the mechanisms underlying the development of locomotion. Although the rat is unable to walk at birth, because of an immature posture, its spinal cord networks can generate at least two kinds of motor patterns in vitro. One activity is called 'fictive locomotion' because it shares several common features with locomotion observed in vivo. This fictive locomotor pattern is rarely observed spontaneously and its release requires either pharmacological or electrical stimulation of the spinal cord. A second endogenously generated activity observed in this preparation occurs spontaneously and exhibits phase relationships between motor outputs that are quite different from the fictive locomotor pattern. Here we review some of the developmental functions this spontaneous activity may subserve. It is likely a major trigger for the maturation of lumbar networks in the fetus, at a stage when inputs from both the periphery and supraspinal structures are weak. Pathways descending from the brainstem arrive in the lumbar enlargement during the last week in utero and the first two postnatal weeks. These pathways, through the neurotransmitters they contain, especially monoamines, are essential for the expression of some neuronal properties and may regulate several ongoing developmental processes.

Afferent Pathways↗
PubMed1991

Differential effects of D1 and D2 dopamine agonists on stereotyped locomotion in rats.

The study examines the effect of selective D1 dopamine stimulation with SKF38393 (1.25-10 mg/kg), on stereotyped locomotion induced by the D2 agonist, quinpirole (0.5 mg/kg). Quinpirole induces repeated travel along a few routes in a limited portion of the environment. Co-administration of low doses of SKF38393 (1.25-2.5 mg/kg) produces the following results: the rate of route perseveration is not affected; the area explored expands to encompass the entire periphery of the open field; and, spatial distribution of locomotion is transformed from routes that cross the center under quinpirole to travel only along the edge. Under higher doses of SKF38393, locomotion ceases. These findings suggest that D1 and D2 stimulation may control the spatial organization of locomotion in oppositional rather than synergistic manner.

2,3,4,5-Tetrahydro-7,8-dihydroxy-1-phenyl-1H-3-ben↗
PubMed2001

Clonidine immediately after immobilization stress prevents long-lasting locomotion reduction in the rat.

1. Stress-induced behavioral change in the rat has been utilized as an animal model of anxiety disorder. The authors examined the effect of early intervention by noradrenergic inhibition on stress-induced long-lasting locomotion reduction. 2. Clonidine, an alpha2 agonist, was administered immediately after a single session of 8 min immobilization stress in a restraining box, followed by locomotion measurement on day 1, day 7, and day 14 after the stress session. 3. In the saline-treated control group, locomotion on day 1, day 7, and day 14 after the 8 min stress session was significantly reduced to about 80% in comparison with that before the stress. This finding confirmed the previous report that a single stressful event could lead to long-lasting behavioral changes. When clonidine was administered, locomotion reduction was not observed on any post-stress day. 4. The results suggest that early intervention by noradrenergic inhibition to stressful events may have a preventive effect on subsequent behavioral change which may be considered as an animal model of post-traumatic stress disorder.

Adrenergic alpha-Agonists↗
PubMed1999

What functions do reflexes serve during human locomotion?

Studies on the reflex modulation of vertebrate locomotion have been conducted in many different laboratories and with many different preparations: for example, lamprey swimming, bird flight, quadrupedal walking in cats and bipedal walking in humans. Emerging concepts are that reflexes are task-, phase- and context-dependent. To function usefully in a behaviour such as locomotion wherein initial conditions change from step to step, reflexes would have to show modulation. Papers are reviewed in which the study of different reflexes have been conducted during different behaviours, with an emphasis on experiments in humans. A framework is developed in which the modulation and flexibility of reflexes are demonstrated. Alterations in cutaneous, and muscle (stretch and load receptor) reflexes between sitting, standing and walking are discussed. Studies in which both electrical, mechanical and 'natural' receptor activation have been conducted during walking are reviewed. Reflexes are shown to have important regulatory functions during human locomotion. A framework for discussion of reflex function throughout the step cycle is developed. The function of a given reflex pathway changes dynamically throughout the locomotor cycle. While all reflexes act in concert to a certain extent, generally cutaneous reflexes act to alter swing limb trajectory to avoid stumbling and falling. Stretch reflexes act to stabilize limb trajectory and assist force production during stance. Load receptor reflexes are shown to have an effect on both stance phase body weight support and step cycle timing. After neurotrauma or in disease, reflexes no longer function as during normal locomotion, but still have the potential to be clinically exploited in gait modification regimens.

Animals↗
PubMed2002

Consequences of monosodium glutamate or goldthioglucose arcuate nucleus lesions on ethanol-induced locomotion.

It has been suggested that the endogenous opioid system, especially beta-endorphins, may play an important role in the behavioral effects of ethanol. The main site of beta-endorphin synthesis in the brain is the hypothalamic arcuate nucleus (ARC). In the present study, we used the neurotoxins monosodium glutamate (MSG) or goldthioglucose (GTG) to produce a selective ARC lesion and to assess its effects on the locomotion observed after ethanol administration. The results show that MSG or GTG pre-treatment produces a blockade of the increased locomotion produced by the injection of low and moderate doses of ethanol (0.5 and 1.5 g/kg, respectively). These effects were observed in the absence of any change in blood ethanol levels. On the other hand, MSG (but not GTG) pre-treatment enhanced the locomotor depression produced by higher doses of this alcohol (2.5 g/kg). Finally, caffeine (10 mg/kg)-induced locomotion was unaffected by the aforementioned neurotoxic agents. Thus, taken together, the present results suggest that MSG and GTG administration produce a blockade of the stimulating effects of ethanol on locomotion in mice and thus provides further support for a role of the ARC in the behavioral effects observed after ethanol administration.

Animals↗
PubMed2001

Biomechanics and kinematics of limb-based locomotion in lizards: review, synthesis and prospectus.

The sprawling pattern of locomotion in lizards is kinematically intriguing and is underpinned by a distinctive pattern of appendicular morphology. The statics of the sprawling posture dictate fundamental design principles, and these place constraints on the three-dimensional kinematics of the limbs and body axis as locomotion is effected. The fore and hind limbs accommodate these constraints and dictates in fundamentally similar, but positionally different ways, resulting in different kinematic profiles for these two appendages. Recent kinematic investigations have helped to clarify earlier generalizations about lizard locomotion and have revealed that kinematic patterns are more variable than was previously supposed. Such analyses, and attendant detailed studies of the anatomy of the locomotor system, promise a new synthesis and enhanced understanding of evolutionary patterns of locomotion of lizards and adjustment to various locomotor substrata and modes of progression.

Animals↗
PubMed2002

Interactions between locomotion and ventilation in tetrapods.

Interactions between locomotion and ventilation have now been studied in several species of reptiles, birds and mammals, from a variety of perspectives. Among these perspectives are neural interactions of separate but linked central controllers; mechanical impacts of locomotion upon ventilatory pressures and flows; and the extent to which the latter may affect gas exchange and the energetics of exercise. A synchrony, i.e. 1:1 pattern of coordination, is observed in many running mammals once they achieve galloping speeds, as well as in flying bats, some flying birds and hopping marsupials. Other, non-1:1, patterns of coordination are seen in trotting and walking quadrupeds, as well as running bipedal humans and running and flying birds. There is evidence for an energetic advantage to coordination of locomotor and respiratory cycles for flying birds and running mammals. There is evidence for a mechanical constraint upon ventilation by locomotion for some reptiles (e.g. iguana), but not for others (e.g. varanids and crocodilians). In diving birds the impact of wing flapping or foot paddling on differential air sac pressures enhances gas exchange during the breath hold by improving diffusive and convective movement of air sac oxygen to parabronchi. This paper will review the current state of our knowledge of such influences of locomotion upon respiratory system function.

Animals↗
PubMed2006

One-sided limb preference is linked to alternating-limb locomotion in anuran amphibians.

Amphibians provide a unique opportunity for identifying possible links between lateralized behaviors, locomotion, and phylogeny and for addressing the origin of lateralized behaviors of higher vertebrates. Five anuran species with different locomotive habits were tested for forelimb and hind limb preferences during 2 stereotyped behavior sequences--wiping a foreign object off their snout and righting themselves from the overturned position. The experiments were analyzed in a broader context of previous findings on anuran lateralization involving 11 anuran species that were studied within the same experimental paradigms. This analysis shows that one-sided forelimb and hind limb motor lateralization in anurans is strongly associated with alternating-limb locomotion and other unilateral limb activity. Conclusions reached for anuran amphibians may be applicable to other vertebrates possessing paired appendages-the degree of lateralization in motor response depends on the mode of locomotion used by a species.

Amphibians↗
PubMed1992

Laufband locomotion with body weight support improved walking in persons with severe spinal cord injuries.

After low transection of the spinal cord mammalian quadrupeds can be trained to walk on a driven surface indicating that coordinating neuronal circuits persist in the spinal cord segments caudal to the lesion. We trained 8 persons with incomplete spinal cord lesion on a Laufband (driven treadmill) for 1 1/2 to 7 months (5 days a week, 30-60 minutes daily) starting 5 to 20 months after injury and found significant improvement in the utilisation of the paralysed limbs during locomotion. Locomotion is described in one additional patient who had trained independently on parallel bars for several years. Five patients had complete functional paralysis in one lower limb when tested in a resting position. In EMG recordings voluntary activity (ie activity induced upon command) was absent or residual in the main flexor and extensor muscles of this limb. In contrast, during locomotion flexion and extension movements were performed and phasic EMG activity was present. In these 5 patients, and in all others reported here, skin sensibility and proprioception are preserved to different degrees in all limbs. In the course of locomotive training of 4 severely paralysed patients the initially habituating flexion reflexes could be entrained in the paralysed limbs as was the case for knee extension during stance. Subsequently, initial body weight support (BWS) of 40% could be reduced to 0%. The distance covered on the Laufband (0-104 m in the first week) increased significantly (200-410 m) in the last week of training as did speed (0-10 to 14-23 m/min). More importantly, this training subsequently allowed patients to walk on a static surface for 100 to 200 meters while voluntary activity remained absent in the paralysed limb when tested at rest. Similar progress was achieved in the 4 less severely paralysed patients. The one patient who had trained independently on parallel bars for several years is described walking on a static surface for 40 meters with the help of a walker, though he had one completely and one near completely paralysed lower limb. It appears that bipedal stepping with consequent knee extension and stabilisation can be taught after unilateral complete or near complete loss of voluntary activity, suggesting the manifestation of complex reflex motor patterns at the spinal level.

Adult↗
PubMed1999

Contrasting mechanisms of action and sensitivity to antipsychotics of phencyclidine versus amphetamine: importance of nucleus accumbens 5-HT2A sites for PCP-induced locomotion in the rat.

In the present study, the comparative mechanisms of action of phencyclidine (PCP) and amphetamine were addressed employing the parameter of locomotion in rats. PCP-induced locomotion (PLOC) was potently blocked by the selective serotonin (5-HT)2A vs. D2 antagonists, SR46349, MDL100,907, ritanserin and fananserin, which barely affected amphetamine-induced locomotion (ALOC). In contrast, the selective D2 vs. 5-HT2A antagonists, eticlopride, raclopride and amisulpride, preferentially inhibited ALOC vs. PLOC. The potency of these drugs and 12 multireceptorial antipsychotics in inhibiting PLOC vs. ALOC correlated significantly with affinities at 5-HT2A vs. D2 receptors, respectively. Amphetamine and PCP both dose dependently increased dialysate levels of dopamine (DA) and 5-HT in the nucleus accumbens, striatum and frontal cortex (FCX) of freely moving rats, but PCP was proportionally more effective than amphetamine in elevating levels of 5-HT vs. DA in the accumbens. Further, whereas microinjection of PCP into the accumbens elicited locomotion, its introduction into the striatum or FCX was ineffective. The action of intra-accumbens PCP, but not intra-accumbens amphetamine, was abolished by SR46349 and clozapine. Parachloroamphetamine, which depleted accumbens pools of 5-HT but not DA, likewise abolished PLOC without affecting ALOC. In contrast, intra-accumbens 6-hydroxydopamine (6-OHDA), which depleted DA but not 5-HT, abolished ALOC but only partially attenuated PLOC. In conclusion, PLOC involves (indirect) activation of accumbens-localized 5-HT2A receptors by 5-HT. PLOC is, correspondingly, more potently blocked than ALOC by antipsychotics displaying marked affinity at 5-HT2A receptors.

Amphetamine↗
PubMed2000

Red nucleus lesions impair overground locomotion in rats: a kinetic analysis.

The red nucleus is a prominent brainstem nucleus in mammals which is thought to be involved in production of skilled limb movements. The presence of the red nucleus and associated rubrospinal tract in animals that do not produce skilled limb movements, however, suggests that these structures might also be involved in control of more general limb actions, such as those occurring during locomotion. The present study investigates this question by measuring the three-dimensional ground reaction forces produced by locomoting rats with unilateral excitotoxic lesions of the red nucleus. Twenty-four to 48 h after the lesion, rats moved with an asymmetric gait during which abnormal braking and propulsive forces were produced during the dual contact time of the forelimb contralateral to the lesion and the ipsilateral hindlimb. Rats did not recover normal symmetrical locomotion within the 55-day duration of the study. The persistent asymmetry produced by red nucleus ablation provides the first unequivocal demonstration that the red nucleus plays a role in ongoing overground locomotion in the rat. Species differences in phylogeny and connectivity of the red nucleus are discussed, as well as the possibility that there is a general compensatory response to unilateral CNS injury in the rat.

Animals↗
PubMed2003

Dynamic analysis of larval locomotion in Drosophila chordotonal organ mutants.

Rhythmic movements, such as peristaltic contraction, are initiated by output from central pattern generator (CPG) networks in the CNS. These oscillatory networks elicit locomotion in the absence of external sensory or descending inputs, but CPG circuits produce more directed and behaviorally relevant movement via peripheral nervous system (PNS) input. Drosophila melanogaster larval locomotion results from patterned muscle contractions moving stereotypically along the body segments, but without PNS feedback, contraction of body segments is uncoordinated. We have dissected the role of a subset of mechanosensory neurons in the larval PNS, the chordotonal organs (chos), in providing sensory feedback to the locomotor CPG circuit with dias (Dynamic Image Analysis System) software. We analyzed mutants carrying cho mutations including atonal, a cho proneural gene, beethoven, a cho cilia class mutant, smetana and touch-insensitive larva B, two axonemal mutants, and 5D10, a weak cho mutant. All cho mutants have defects in gross path morphology compared to controls. These mutants exhibit increased frequency and duration of turning (decision-making) and reduced duration of linear locomotion. Furthermore, cho mutants affect locomotor parameters, including reduced average speed, direction change, and persistence. Dias analysis of peristaltic waves indicates that mutants exhibit reduced average speed, positive flow and negative flow, and increased stride period. Thus, cho sensilla are major proprioceptive components that underlie touch sensitivity, locomotion, and peristaltic contraction by providing sensory feedback to the locomotor CPG circuit in larvae.

Animals↗
PubMed1987

Abnormal gait sequence in locomotion after atropine treatment of catecholamine-deficient akinetic rats.

Excessive, abnormal locomotion occurs after a high dose (25-50 mg/kg) of atropine sulfate to rats already akinetic due to catecholamine deficiency from intraventricular administration of 6-hydroxydopamine. This abnormal locomotion involves an abnormal gait sequence [right (R) hindleg (H), left (L) foreleg (F), LH, RF] instead of the normal gait sequence (RH, RF, LH, LF). In such animals atropine progressively (i) decreases hindleg step size, (ii) decreases arching of the trunk, and (iii) increases foreleg step size. These factors combine to change the ratio of front/hind body support. If the body stretches too far and the hindleg step is too small, a given hindleg step supports insufficient weight to remove weight from the ipsilateral foreleg; consequently, the opposite foreleg must execute the next step, producing the abnormal gait sequence. Thus, atropine affects gait sequence indirectly; it acts on at least three variables that affect how body weight is distributed and shifted during locomotion. To maintain stability during such locomotion, gait sequence is appropriately altered.

Animals↗
PubMed1994

Kinematics of treadmill versus overground locomotion in horses.

A kinematic analysis was carried out to compare treadmill and overground locomotion in horses at the trot. Stride variables and limb and trunk movements of 10 Dutch Warmblood horses were measured using the CODA-3 gait analysis system. Overground recordings were made on a rubber ground surface and on an asphalt track. Treadmill recordings were taken after a controlled habituation programme and at the same velocities as measured overground. On asphalt, a shorthened stride duration and a decreased vertical displacement of the withers were found compared with those on rubber ground. On the treadmill, stance duration of the forelimbs was longer than for either overground condition. Correspondingly, the hind limbs were placed earlier than the forelimbs in making diagonal ground contact in overground locomotion, but this changed to preceding forelimbs on the treadmill. Both forelimbs and hind limbs were moved more caudally during the retraction phase on the treadmill, while no differences were found in the protraction angle. The vertical movement of the hooves as well as the withers was smaller on the treadmill than on rubber ground. Treadmill belt velocity decreased by 9% during the stance phase. This is supposed to be an important reason for the differences in biomechanics and kinematics between treadmill and overground locomotion, which must be kept in mind when data obtained during treadmill locomotion have to be extrapolated to overground conditions.

Animals↗
PubMed2000

Calcium/calmodulin-dependent protein kinase II regulates Caenorhabditis elegans locomotion in concert with a G(o)/G(q) signaling network.

Caenorhabditis elegans locomotion is a complex behavior generated by a defined set of motor neurons and interneurons. Genetic analysis shows that UNC-43, the C. elegans Ca(2+)/calmodulin protein kinase II (CaMKII), controls locomotion rate. Elevated UNC-43 activity, from a gain-of-function mutation, causes severely lethargic locomotion, presumably by inappropriate phosphorylation of targets. In a genetic screen for suppressors of this phenotype, we identified multiple alleles of four genes in a G(o)/G(q) G-protein signaling network, which has been shown to regulate synaptic activity via diacylglycerol. Mutations in goa-1, dgk-1, eat-16, or eat-11 strongly or completely suppressed unc-43(gf) lethargy, but affected other mutants with reduced locomotion only weakly. We conclude that CaMKII and G(o)/G(q) pathways act in concert to regulate synaptic activity, perhaps through a direct interaction between CaMKII and G(o).

Animals↗
PubMed2002

Central pattern generation of locomotion: a review of the evidence.

Neural networks in the spinal cord, referred to as "central pattern generators" (CPGs), are capable of producing rhythmic movements, such as swimming, walking, and hopping, even when isolated from the brain and sensory inputs. This article reviews the evidence for CPGs governing locomotion and addresses other factors, including supraspinal, sensory, and neuromodulatory influences, that interact with CPGs to shape the final motor output. Supraspinal inputs play a major role not only in initiating locomotion but also in adapting the locomotor pattern to environmental and motivational conditions. Sensory afferents involved in muscle and cutaneous reflexes have important regulatory functions in preserving balance and ensuring stable phase transitions in the locomotor cycle. Neuromodulators evoke changes in cellular and synaptic properties of CPG neurons, conferring flexibility to CPG circuits. Briefly addressed is the interaction of CPG networks to produce intersegmental coordination for locomotion. Evidence for CPGs in humans is reviewed, and although a comprehensive clinical review is not an objective, examples are provided of animal and human studies that apply knowledge of CPG mechanisms to improve locomotion. The final section deals with future directions in CPG research.

Animals↗
PubMed1988

Neural mechanisms generating locomotion studied in mammalian brain stem-spinal cord in vitro.

The neural control system for generation of locomotion is an important system for analysis of neural mechanisms underlying complex motor acts. In these studies, a novel experimental model using neonatal rat brain stem and spinal cord in vitro was developed for investigation of the locomotor system in mammals. The in vitro brain stem and spinal cord system was shown to retain functional circuitry for locomotor command generation, motor pattern generation, and sensorimotor integration. This system was exploited to investigate neurochemical mechanisms involved in neurogenesis of locomotion. Evidence was obtained for peptidergic and gamma-amino-butyric acid-mediated mechanisms in brain-stem circuits generating locomotor commands. Cholinergic, dopaminergic, and excitatory amino acid-mediated mechanisms were shown to activate spinal cord circuits for locomotor pattern generation. Endogenous N-methyl-D-aspartic acid receptors in spinal networks were found to play a central role in the generation of locomotion. The chemically induced patterns of motor activity and rhythmic membrane potential oscillations of spinal motoneurons were characteristic of those during locomotion in other mammals in vivo. The in vitro brain stem and spinal cord model provides a versatile and powerful experimental system with potentially broad application for investigation of diverse aspects of the neurobiology of mammalian motor control systems.

Animals↗