Central mechanisms underlying fish swimming

Although the basic swimming rhythm is created by central pattern generators (CPGs) located in each spinal segment, command signals from the brain should be indispensable for the activation of CPGs to initiate swimming. We hypothesized that the nucleus of medial longitudinal fascicles (Nflm) is the midbrain locomotor region driving swimming rhythms in teleosts. To test this hypothesis, we recorded neuronal activities from Nflm neurons in swimming carp and analyzed the cytoarchitecture of the nucleus. We identified two types of Nflm neurons exhibiting electric activities closely related to swimming rhythms. Remarkably, tonic neurons that continued firing during swimming were found. The Nflm and neighboring oculomotor nucleus contain about 600 neurons in total, and among them as many as 500 were labeled retrogradely by an intraspinal tracer implantation and 400 neurons showed glutamatergic immunoreactivity. They are the most likely candidates for the descending neurons as the origin of driving signals that initiate swimming. Double-labeling experiments demonstrated direct connections of Nflm neurons to spinal neurons consisting of the CPG. These data imply that most Nflm neurons possibly exert an excitatory drive to the spinal CPGs through the descending axons with excitatory transmitter(s), probably glutamate. Furthermore, we confirmed that the caudal part of Nflm and the rostral part of the oculomotor nucleus overlap rostrocaudally by approximately 200 mum. In connection with the control of swimming by the brain, we carried out experiments to clarify the efferent system of the cerebellum of the goldfish. Cerebellar efferent fibers terminated in most brain regions except for the telencephalon. Importantly, the cerebellum projected also to the Nflm, suggesting the involvement of this brain region in the control of swimming. We have also determined that in the carp so-called eurydendroid cells are cerebellar efferent neurons.     

Eye movements evoked by electrical stimualtion of the brain in anesthetized fishes.

Several eye movements were evoked by electrical stimulation of the brain in anesthetized sunfish and goldfish. Conjugate lateral rolling movements, similar to eye movements observed when an unoperated fish is rotated about its long axis, were evoked from the acoustico-lateral area of the medulla and the eminentia granularis and an adjacent medial portion of the cerebellum. Bilateral and unilateral backward rotations, similar to the eye movements observed when unoperated fish are rotated forward about the interpupillary axis, were evoked from the medial longitudinal fasciculus and areas related to the oculomotor nerve. Bilateral forward rotations, comparable to the eye movements resulting when unoperated fish are rotated backward about the interpupillary axis, were elicited by stimulation near the trochlear nerve roots in the valvula of the cerebellum; unilateral responses resulted from stimulation near the exiting trochlear nerves. Convergence was elicited by stimulation in the midline near the oculomotor complex and the medial longitudinal fasciculus while unilateral vergence responses were triggered by stimulation in the medial longitudinal fasciculus and areas lateral to the oculomotor nucleus. Conjugate eye movements in the horizontal plane were frequently evoked but were not studied in detail.     

Internuclear ophthalmoplegia of abduction: clinical and electrophysiological data on the existence of an abduction paresis of prenuclear origin.

Three patients showed unilateral and five bilateral abduction paresis. Five had associated adduction nystagmus of the contralateral eye. Electrophysiological testing of masseter and blink reflexes indicated an ipsilateral rostral pontine or mesencephalic lesion, and excluded a lesion of the infranuclear portion of the abducens nerve. Abduction paresis was attributed to impaired inhibition of the tonic resting activity of the antagonistic medial rectus muscle. The prenuclear origin of the disorder is based on morphological and neurophysiological evidence of an ipsilateral inhibitory connection between the paramedian pontine reticular formation and the oculomotor nucleus running close to but separated from the medial longitudinal fasciculus.     

Downward gaze in monkeys: stimulation and lesion studies.

Ten monkeys were stimulated unilaterally and bilaterally through bipolar electrodes placed stereotactically on each side of the midline under light barbiturate anaesthesia. Bilateral simultaneous stimulation elicited straight downward binocular movements from a core of tissue about 40 mm3 on each side which included the fields of Forel, zona incerta, subthalamic nucleus, oral pole of the red nucleus, fasciculus retroflexus and 'area tegmentalis'. Unilateral stimulation of the same points yielded downward eye movements in only 25 per cent of the instances. Upward deviation of the globes could be elicited by bilateral stimulation of tissue located more caudal, ventral and medial than that from which downward movements were obtained. Bilateral electrolytic lesions within the region outlined above caused significant defects in downward gaze both in saccadic and slow pursuit binocular movements. Passive bending of the head backwards, however, resulted in downward deviation of the globes (oculocephalic reflex). Optokinetic nystagmus and after-nystagmus downward were abolished. Oblique (45 degrees) optokinetic stimulation elicited a perverted response in the horizontal plane. Vestibulo-ocular reflexes elicited by bilateral warm irrigation of both ear canals with the monkey in the erect position, or by turning the animal while lying on one side, caused a strong tonic deviation upward with absence of nystagmus downward. Some of these monkeys showed additional alterations in upward gaze but they were less severe in intensity and duration than those of downward gaze. All eye deviations in the horizontal plane were consistently normal. Recovery occurred in all types of vertical binocular movements except in the rapid motions (saccades and quick phases of nystagmus) below the horizontal meridian. A unilateral lesion had no effect. The minimal damage producing downward gaze defects was about 1.7 mm in diameter, cetred in the prerubral fields, rostral and medial to the red nuclei with minimal involvement of the oral pole of these structures. The nuclei of Cajal, Darkschewitsch and interstitialis of the posterior commissure, as well as the fasciculus retroflexus and the posterior commissure, were spared by this lesion. The so-called rostral interstitial nucleus of the medial longitudinal fasciculus and the nucleus campi Foreli appear to be destroyed. These structures are known to receive an input from the paramedian pontine reticular formation and project on to the oculomotor nerve nucleus. These results demonstrate that the prerubral fields contain structures which are critical for rapid eye movements downward, and therefore an isolated downward gaze palsy is a strong indicator of a bilateral lesion of this zone. The findings in the few reported cases with this sign and available pathological analysis suggest that our conclusions from the experimental monkey apply to man as well. The concept of bilateral innervation for vertical eye movements is amply confirmed for the downward vectors...     

Bilateral sequential motor cortex stimulation and skilled task performance with non-dominant hand

Objective

To check whether bilateral sequential stimulation (BSS) of M1 with theta burst stimulation (TBS), using facilitatory protocol over non-dominant M1 followed by inhibitory one over dominant M1, can improve skilled task performance with non-dominant hand more than either of the unilateral stimulations do. Both, direct motor cortex (M1) facilitatory non-invasive brain stimulation (NIBS) and contralateral M1 inhibitory NIBS were shown to improve motor learning.

Phasic modulation of reticulospinal neurones during fictive locomotion and other types of spinal motor activity in lamprey

The intracellular activity of different types of reticulospinal neurones was studied during fictive locomotion and other types of spinal motor activity in an in vitro preparation of the lamprey brainstem-spinal cord. The examined neurones included large Müller cells of the rhombencephalic and mesencephalic reticular formation, the Mauthner cell, and neurones in the posterior rhombencephalic reticular nucleus with different sizes and conduction velocities. During bouts of fictive swimming initiated spontaneously or by stimulation of the trigeminal nerve or spinal cord, the Müller cells were depolarized and fired action potentials. Bulbar Müller cells in addition showed a phasic modulation of membrane potential with excitation in phase with ipsilateral motoneurones of the rostral spinal cord. The Mauthner cell was depolarized in phase with contralateral motoneurones. Many neurones in the posterior rhombencephalic reticular nucleus showed modulation in phase with ipsilateral motoneurones during fictive swimming. Such oscillations were observed in both fast-conducting neurones, located mainly in the medial part of the nucleus, and slower conducting cells with a more lateral distribution. All examined reticulospinal neurones showed a strong coupling also with other types of spinal motor activity, such as slow alternating bursting and synchronous bilateral ventral root bursts, but the reticulospinal activity had no correlation with respiratory activity recorded from the Xth nerve. The consequences of a phasic reticulospinal activity during locomotion are discussed.     

Tectal connections in Python reticulatus

The origins of the axons terminating in the mesencephalic tectum in Python reticulatus were examined by unilateral tectal injections of horseradish peroxidase. Kutrogradely labeled cells were observed bilaterally throughout the spinal cord in all subdivisions of the trigeminal system, with the exception of nucleus principalis, which showed labeled cells only on the ipsilateral side. Labeling of the reticular formation occurred bilaterally in nucleus reticular is interiormagnocellularis, nucleus reticularis lateralis, nucleus reticularis medius and the mesencephalic reticular formation. The tectum also receives bilateral projections from the dorsal tegmentaJ field, the nucleus of the lateral lemniscus and nucleus isthmi, and ipsilateral projections from nucleus profundus mesencephali. A few labeled cells were found ipsilaterally in the locus coeruleus and in nuclei vestibulares ventrolateralis and centromedialis. In the diencephalon labeled cells were observed ipsilaterally in nucleus ventrolateralis thalami, nucleus ventromedialis thalami, nucleus suprapeduncularis, and in the dorsal and ventral lateral geniculate nuclei. Bilateral labeling was observed in nucleus periventricularis hypo-thalami. Furthermore, labeling was ipsilaterally present in the ventral telen-cephalic areas. The tectum in Python reticulatus receives a wide variety of afferent connections which confirm the role of the tectum as an integration center of visual and exteroceptive information.     

Projections of the lateral terminal accessory optic nucleus of the common marmoset (Callithrix jacchus)

The connections of the lateral terminal nucleus (LTN) of the accessory optic system (AOS) of the marmoset monkey were studied with anterograde ³H-amino acid light autoradiography and horseradish peroxidase retrograde labeling techniques. Results show a first and largest LTN projection to the pretectal and AOS nuclei including the ipsilateral nucleus of the optic tract, dorsal terminal nucleus, and interstitial nucleus of the superior fasciculus (posterior fibers); smaller contralateral projections are to the olivary pretectal nucleus, dorsal terminal nucleus, and LTN. A second, mejor bundle produces moderate-to-heavy labeling in all ipsilateral, accessory oculornotor nuclei (nucleus of posterior commissure, interstitial nucleus of Cajal, nucleus of Darkschewitsch) and nucleus of Bechterew; some of the fibers are distributed above the caudal oculomotor complex within the supraoculornotor periaqueductal gray. A third projection is ipsilateral to the pontine and mesencephalic reticular formations, nucleus reticularis tegmenti pontis and basilar pontine complex (dorsolateral nucleus only), dorsal parts of the medial terminal accessory optic nucleus, ventral tegmental area of Tsai, and rostral interstitial nucleus of the medial longitudinal fasciculus. Lastly, there are two long descending bundles: (1) one travels within the medial longitudinal fasciculus to terminate in the dorsal cap (ipsilateral > > contralateral) and medial accessory olive (ipsilateral only) of the inferior olivary complex. (2) The second soon splits, sending axons within the ipsilateral and contralateral brachium conjunctivum and is distributed to the superior and medial vestibular nuclei. The present findings are in general agreement with the documented connections of LTN with brainstem oculomotor centers in other species. In addition, there are unique connections in marmoset monkey that may have developed to serve the more complex oculomotor behavior of nonhuman primates. © 1995 Wiley-Liss, Inc.     

Projections of the vestibular and cerebellar nuclei in Rana pipiens

The efferent projections and cytoarchitecture of the vestibulocerebellar region were examined to determine the nuclear boundaries and potential homologies. The anterior portion of the vestibular complex projects to the ipsilateral oculomotor and trochlear nuclei and is the major source of commissural fibers. Neurons in the rostromedial portions of the complex project to the contralateral trochlear nucleus. Large neurons in the ventrolateral portion of the complex give rise to a bilateral vestibulospinal pathway. Medium-sized neurons in the neuropil and small neurons in the central gray giving rise to bilateral projections to the spinal cord and oculomotor nuclei as well as commissural and ipsilateral cerebellar efferents. Projections from the nucleus of the cerebellum reach the contralateral spinal cord and cerebellar nucleus and there is also a bilateral projection to the ventral rhombencephalic and mesencephalic basal plates. The medial portion of the nucleus gives rise to commissural, ipsilateral mesencephalic and contralateral spinal projections. The lateral portion of the nucleus projects to the contralateral ventral mesencephalon. On the whole, the results of this investigation substantiate the division of the anuran vestibular complex in anurans into nuclei which may be homologous to the superior nucleus and nucleus of Deiters in mammals. The case for distinct descending and medial nuclei is less compelling. Further, it appears possible to divide the nucleus of the cerebellum into medial and lateral components whose connectivity is similar to that of reptiles and to a lesser extent mammals.     

Anatomical and physiological characteristics of vestibular neurons mediating the horizontal vestibulo-ocular reflex of the squirrel monkey

The anatomical characteristics of vestibular neurons, which are involved in controlling the horizontal vestibulo-ocular reflex, were studied by injecting horseradish peroxidase (HRP) into neurons whose response during spontaneous eye movements had been characterized in alert squirrel monkeys. Most of the vestibular neurons injected with HRP that had axons projecting to the abducens nucleus or the medial rectus subdivision of the oculomotor nucleus had discharge rates related to eye position and eye velocity. Three morphological types of cells were injected whose firing rates were related to horizontal eye movements. Two of the cell types were located in the ventral lateral vestibular nucleus and the ventral part of the medial vestibular nucleus (MV). These vestibular neurons could be activated at monosynaptic latencies following electrical stimulation of the vestibular nerve; increased their firing rate when the eye moved in the direction contralateral to the soma; had tonic firing rates that increased when the eye was held in contralateral positions; and had a pause in their firing rate during saccadic eye movements in the ipsilateral or vertical directions. Eleven of the above cells had axons that arborized exclusively on the contralateral side of the brainstem, terminating in the contralateral abducens nucleus, the dorsal paramedian pontine reticular formation, the prepositus nucleus, medial vestibular nucleus, dorsal medullary reticular formation, caudal interstitial nucleus of the medial longitudinal fasciculus, and raphé obscurus. Eight of the cells had axons that projected rostrally in the ascending tract of Deiters and arborized exclusively on the ipsilateral side of the brainstem, terminating in the ipsilateral medial rectus subdivision of the oculomotor nucleus and, in some cases, the dorsal paramedian pontine reticular formation or the caudal interstitial nucleus of the medial longitudinal fasciculus. Two MV neurons were injected that had discharge rates related to ipsilateral eye position, generated bursts of spikes during saccades in the ipsilateral direction, and paused during saccades in the contralateral direction. The axons of those cells arborized ipsilaterally, and terminated in the ipsilateral abducens nucleus, MV, prepositus nucleus, and the dorsal medullary reticular formation. The morphology of vestibular neurons that projected to the abducens nucleus whose discharge rate was not related to eye movements, or was related primarily to vertical eye movements, is also briefly presented.     

Cooperative hand movements in tetraplegic spinal cord injury patients: Preserved neural coupling

Objectives

To evaluate whether the task-specific neural coupling mechanism during the performance of cooperative hand movements is preserved in tetraplegic subjects.

Vertical one-and-a-half syndrome. Supranuclear downgaze paralysis with monocular elevation palsy

A patient with bilateral infarction in the mesodiencephalic region showed impairment of all downward rapid eye movements (including vestibulo-ocular movements) and foveal smooth pursuit (nondissociated downgaze paralysis) associated with monocular paralysis of elevation (vertical one-and-a-half syndrome). Bell's phenomenon and all types of horizontal eye movements were preserved. The lesions may have affected the efferent tracts of the rostral interstitial nucleus of the medial longitudinal fasciculus bilaterally and the premotor fibers to the contralateral superior rectus subnucleus and ipsilateral inferior oblique subnucleus, either before or after decussation in the posterior commissure.     

Effects of stimulating the reticular formation during fictive locomotion in lampreys

The effects of stimulating the reticular formation were studied during fictive locomotion in lampreys (Ichthyomyzon unicuspis). The in vitro isolated preparation of the brainstem and spinal cord was used and fictive locomotion was induced by bath application of N-methyl-

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-aspartate (NMDA; 50–100 μM). During different phases of the locomotor cycle, short trains of stimuli (10 pulses at 80–100 Hz; 10 μA) were delivered through glass-coated tungsten microelectrodes positioned within the middle rhombencephalic reticular nucleus (MRRN) and their effects were studied on ipsi- and contralateral ventral root locomotor discharges. Irrespective of the locomotor phase during which the stimulation train was delivered, a resetting effect occurred. It was characterized by a re-synchronization of the locomotor discharges with a constant latency for each ventral root on the ipsilateral side. The latency increased as the recorded root was located further caudally. This increase in latency was in the range of the phase lag observed between roots during control bouts of locomotion. These results suggest that reticulospinal neurones exert strong resetting effects on spinal locomotor networks. These effects may play a significant role with respect to changes of direction during swimming.     

Identification of the midbrain locomotor region and its relation to descending locomotor pathways in the Atlantic stingray,Dasyatis sabina

The midbrain locomotor region (MLR) in the Atlantic stingray,Dasyatis sabina, was identified and characterized. Stimulation (50–100 μA, 60 Hz) of the midbrain in decerebrated, paralyzed animals was used to elicit locomotion monitored as alternating activity in nerves innervating an antagonist pair of elevator and depressor muscles. Effective sites for evoking locomotion in the midbrain included parts of several nuclei: the caudal portion of the interstitial nucleus of the medial longitudinal fasciculus and the caudomedial parts of the cuneiform and subcuneiform nuclei. This region did not include the red nucleus, any parts of the optic tectum or the medial or lateral mesencephalic nuclei. Electrical stimulation in the MLR evokes locomotion in either the ipsi- or contralateral pectoral fin, wheres stimulation in the medullary reticular formation evokes locomotion only in the contralateral fin. Lesion experiments were performed to identify the location of descending pathways from the midbrain to the medullary reticular formation. To abolish locomotion evoked by electrical stimulation in the MLR, the medial reticular formation in the rostral medulla had to be lesioned bilaterally, or the ipsilateral medial medullary reticular formation and fibers projecting from the MLR to the contralateral midbrain had to be disrupted. Injections of HRP into the magnocellular/gigantocellular reticular formation confirmed that this area received bilateral projections from the MLR. The MLR of the Atlantic stingray appears to be similar to the lateral component of the mammalian MLR and to the MLR in other non-mammalian vertebrates.     

Uncrossed retinal projection to the medial terminal nucleus of the accessory optic system in the hereditary unilaterally microphthalmic rat

The uncrossed optic projection to the medial terminal nucleus of the accessory optic system was studied in adult rats with hereditary unilateral microphthalmia. The vestigial eye of the mutant completely lacks the optic nerve. After injection of HRP into the remaining eye the ipsilateral labeled fibers of the accessory optic system via the superior fasciculus distinctly covered the three terminal nuclei. No ipsilateral inferior fasciculus of the accessory optic system was labeled. Aberrant expansion of the uncrossed superior fasciculus fibers was observed in the medial terminal nucleus.     

Efferent projections of the deep mesencephalic nucleus (pars medialis) in the rat

The projections of the medial part of the deep mesencephalic nucleus (DMN) were traced by autoradiography and retrograde horseradish peroxidase (HRP) techniques. No ascending projections were observed from the medial part of the DMN; however, two groups of descending fibers were observed. One group crossed the midline and coursed to the caudal part of the red nucleus. At this point, these fibers divided into two distinct bundles. One bundle of fibers passed caudally to terminate in the contralateral pontine reticular nucleus, superior olive-trapezoid body complex, gigantocellular nucleus, and upper cervical spinal cord. The other bundle entered the medial longitudinal fasciculus (MLF) and coursed through the pons and medulla without termination, to enter the cervical spinal cord where terminations were noted in the dorsal horn. The other group of fibers from the medial DMN descended through the ipsilateral pons and medulla, projecting to the pontine reticular nucleus, superior olive-trapezoid body complex, gigantocellular nucleus, and upper cervical spinal cord. These findings indicate that the medial part of the DMN is likely to be involved in complex sensorimotor events via reticulobulbar and reticulospinal connections.     

The one-and-a-half syndrome--a unilateral disorder of the pontine tegmentum: a study of 20 cases and review of the literature

The one-and-a-half syndrome is a clinical disorder of extraocular movements characterized by a conjugate horizontal gaze palsy in one direction plus an internuclear ophthalmoplegia in the other. The syndrome is usually due to a single unilateral lesion of the paramedian pontine reticular formation or the abducens nucleus on one side (causing the conjugate gaze palsy), with interruption of internuclear fibers of the ipsilateral medial longitudinal fasciculus after it has crossed the midline from its site of origin in the contralateral abducens nucleus (causing failure of adduction of the ipsilateral eye). Twenty cases are reported; 14 had multiple sclerosis.     

Modulation of physiological mirror activity with transcranial direct current stimulation over dorsal premotor cortex

Humans have a natural tendency towards symmetrical movements, which rely on a distributed cortical network that allows for complex unimanual movements. Studies on healthy humans using rTMS have shown that disruption of this network, and particularly the dorsal premotor cortex (dPMC), can result in increased physiological mirror movements. The aim of the present set of experiments was to further investigate the role of dPMC in restricting motor output to the contralateral hand and determine whether physiological mirror movements could be decreased in healthy individuals. Physiological mirror movements were assessed before and after transcranial direct current stimulation (tDCS) over right and left dPMC in three conditions: bilateral, unilateral left and unilateral right stimulation. Mirror EMG activity was assessed immediately before, 0, 10 and 20 min after tDCS. Results show that physiological mirroring increased significantly in the hand ipsilateral to cathodal stimulation during bilateral stimulation of the dPMC, 10 and 20 min after stimulation compared to baseline. There was no significant modulation of physiological mirroring in the hand ipsilateral to anodal stimulation in the bilateral condition or following unilateral anodal or unilateral cathodal stimulation. The present data further implicate the dPMC in the control of unimanual hand movements and show that physiological mirroring can be increased but not decreased with dPMC tDCS.     

Descending pathways eliciting forelimb stepping in the lateral funiculus: Experimental studies with stimulation and lesion of the cervical cord in decerebrate cats

The funicular pathways that elicit forelimb stepping were investigated with stimulation and lesion of the cervical white matter in decerebrate cats with the lower thoracic cord transected. We localized cross-sectional areas where stimulation evoked rhythmic motor-nerve discharges imitating those of stepping (fictive locomotion) in the immobilized animal, and further examined whether or not lesions made in the corresponding areas abolished actual locomotor movements. Stimulation of the C3 lateral funiculus (LF) produced fictive locomotion in the ipsilateral forelimb. The effective areas of stimulation were located separately in the dorsolateral funiculus (DLF) and in the ventrolateral funiculus (VLF), while the VLF was more effective than the DLF. Effective stimuli were pulse trains with a frequency of about 30 Hz, with a rather wide pulse duration of about 0.5 ms. Blocking axonal conduction through the lower thoracic cord by cooling reproducibly facilitated fictive locomotion in both amplitude and frequency. In the lesion experiments, forelimb locomotor movements were elicited spontaneously or by stimulation of the mesencephalic locomotor region (MLR). The locomotor movements were abolished by complete lesions of the C2-C3 LFs on both sides, but these remained when either the DLF or the VLF was intact on one side. These findings together suggested that the descending pathways for the activation of the spinal locomotor network of the single forelimb are located ipsilaterally in the DLF as well as in the VLF. Both the DLF and the VLF pathways can initiate locomotion, while the VLF pathways have a higher potential for its initiation. Lesion experiments further showed that cats can walk with both forelimbs, even though the spinal locomotor network of the single forelimb was deprived of its main descending input by unilateral lesions of the LF. However, when the unilateral lesion extended to the medial part of the LF, the bilateral walking was abolished; the limb on the lesioned side showed only rhythmic extension movements without active flexion movements, which was out of phase with the stepping movements on the intact side. This finding suggested that the medial part of the LF is important for producing flexion movements during the swing phase of stepping.     

Identification of the midbrain locomotor region and its relation to descending locomotor pathways in the Atlantic stingray, Dasyatis sabina.

The midbrain locomotor region (MLR) in the Atlantic stingray, Dasyatis sabina, was identified and characterized. Stimulation (50-100 microA, 60 Hz) of the midbrain in decerebrated, paralyzed animals was used to elicit locomotion monitored as alternating activity in nerves innervating an antagonist pair of elevator and depressor muscles. Effective sites for evoking locomotion in the midbrain included parts of several nuclei: the caudal portion of the interstitial nucleus of the medial longitudinal fasciculus and the caudomedial parts of the cuneiform and subcuneiform nuclei. This region did not include the red nucleus, any parts of the optic tectum or the medial or lateral mesencephalic nuclei. Electrical stimulation in the MLR evokes locomotion in either the ipsi- or contralateral pectoral fin, whereas stimulation in the medullary reticular formation evokes locomotion only in the contralateral fin. Lesion experiments were performed to identify the location of descending pathways from the midbrain to the medullary reticular formation. To abolish locomotion evoked by electrical stimulation in the MLR, the medial reticular formation in the rostral medulla had to be lesioned bilaterally, or the ipsilateral medial medullary reticular formation and fibers projecting from the MLR to the contralateral midbrain had to be disrupted. Injections of HRP into the magnocellular/gigantocellular reticular formation confirmed that this area received bilateral projections from the MLR. The MLR of the Atlantic stingray appears to be similar to the lateral component of the mammalian MLR and to the MLR in other non-mammalian vertebrates.