Gamma-motoneurone discharge patterns during fictive locomotion in the decerebrate cat

Triceps surae gamma-motoneurones were recorded during fictive locomotion in the paralysed high decerebrate cat. Two distinctive patterns of discharge were observed which were similar to those reported for static and dynamic gamma-motoneurones during locomotion in the same preparation, but without paralysis (Murphy, Stein & Taylor, 1984). These results suggest that movement-related afferent feedback is not essential for the generation of the basic patterns of static and dynamic gamma-motoneurone activity during locomotion. The results are discussed in relation to the generation of alpha and gamma locomotor rhythms.     

Toe flexor muscle spindle discharge and stretch modulation during locomotor activity in the decerebrate cat

In order to investigate the nature (i.e. static or dynamic) of fusimotor drive to the flexor hallucis longus (FHL) and flexor digitorum longus (FDL) muscles during locomotion we recorded Ia and group II muscle spindle afferent responses to sinusoidal stretch (0.25 and 1 mm amplitude, respectively, 4–5 Hz) in a decerebrate cat preparation. FHL Ia and group II afferents generally had increased discharge rates and decreased modulation to stretch throughout the step cycle, compared to rest, suggesting raised static γ drive at all locomotor phases. Although the modulation of Ia afferents was reduced during locomotion, most (13 of 18) showed a clear increasing trend during homonymous muscle activity (extension). This was consistent with phasic dynamic γ drive to FHL spindles linked with α drive. In agreement with previous reports, FHL gave a single burst of EMG activity during the step cycle while FDL α drive had two components. One was related to extension while the other comprised a brief burst around the end of this phase. Typically FDL Ia and group II afferents also had elevated firing rates and reduced modulation at all locomotor phases, again implicating static γ drive. Half the afferents (seven Ia, three group II) showed increased discharge during extension, suggesting phasic static γ drive. There was no γ drive associated with the late FDL α burst. In conclusion, the γ drives to FHL and FDL differed during locomotion. FHL, which has the α drive of a classic extensor, received γ drive that closely resembled other extensors. The γ drive of FDL, which exhibits both extensor and flexor α synergies, did not match either muscle type. These observations are compatible with the view that fusimotor drive varies in different muscles during locomotion according to the prevailing sensorimotor requirements.     

Tonic and phasic phenomena underlying eye movements during sleep in the cat

Mammalian sleep is not a homogenous state, and different variables have traditionally been used to distinguish different periods during sleep. Of these variables, eye movement is one of the most paradigmatic, and has been used to differentiate between the so-called rapid eye movement (REM) and non-REM (NREM) sleep periods. Despite this, eye movements during sleep are poorly understood, and the behaviour of the oculomotor system remains almost unknown. In the present work, we recorded binocular eye movements during the sleep–wake cycle of adult cats by the scleral search-coil technique. During alertness, eye movements consisted of conjugated saccades and eye fixations. During NREM sleep, eye movements were slow and mostly unconjugated. The two eyes moved upwardly and in the abducting direction, producing a tonic divergence and elevation of the visual axis. During the transition period between NREM and REM sleep, rapid monocular eye movements of low amplitude in the abducting direction occurred in coincidence with ponto-geniculo-occipital waves. Along REM sleep, the eyes tended to maintain a tonic convergence and depression, broken by high-frequency bursts of complex rapid eye movements. In the horizontal plane, each eye movement in the burst comprised two consecutive movements in opposite directions, which were more evident in the eye that performed the abducting movements. In the vertical plane, rapid eye movements were always upward. Comparisons of the characteristics of eye movements during the sleep–wake cycle reveal the uniqueness of eye movements during sleep, and the noteworthy existence of tonic and phasic phenomena in the oculomotor system, not observed until now.     

Comparative analysis of cerebellar unit discharge patterns in the decerebrate cat during passive movements

Summary 1) The present experiments were undertaken to study how information about the parameters of a passive movement is processed at different neuronal levels of the cat cerebellar cortex. The analysis has been performed by recording extracellularly in the intermediate part of the cerebellar anterior lobe from presumed mossy fibres, presumed granule cells, and Purkinje cells with simple spikes and complex spikes. 2) The discharge patterns obtained during passive movements of the cat's forepaw were characterized by components which could be related to dynamic or static parameters of the movement. With respect to the occurrence of dynamic responses, patterns were classified according to a statistically derived measure in three different types. By using the same statistical measure, discharge patterns were additionally classified into two subgroups according to their response components reflecting static parameters. Within the patterns a clearcut relationship between dynamic and static components was observed. The corresponding distributions are shown and discussed. 3) A very interesting result of the classification of cerebellar discharge patterns is that the distribution of the different types depended on the level of integration within the cerebellar cortex. Patterns of the low scale integrated cerebellar input (mossy fibre-system), as well as those of granule cells (the first cerebellar computational niveau), reflected both static and dynamic movement parameters. At the Purkinje cell level (a level with a high degree of convergence) the discharge patterns are characterized predominantly by dynamic responses. 4) The interrelationship between complex- and simple spikes of Purkinje cells was tested by different methods: a) By analyzing the paired values of the mean complex-(CS) and simple spike (SS) discharge probabilities of 110 Purkinje cells a scatter was obtained, indicating an underlying hyperbolic relation (prob(CS) = a/(prob(SS))^b). Thus, a high CS discharge probability is accompanied by a low SS probability and vice versa, b) The timelocked complex- and simple spike responses were studied by comparing the similarity of their responses. All combinations of complex- and simple spike patterns were observed, ranging from a sign correct similarity to a mirror image similarity. The distribution of the measure for similarity shows that the mirror image predominated, c) The individual simple spike discharge probability is characterized by a pause evoked by the occurrence of a complex spike event. The simple spike discharge probabilities during an interval preceeding and following a complex spike event were compared. A post climbing pause coefficient was defined as a measure for the effectiveness of the complex spike event. No relationship between these coefficients and the above mentioned measure for similarity was found. Hence, for the Purkinje cell discharging with the simple spikes independent spike generating processes have to be assumed. 5) From these results it can be derived that cerebellar discharge patterns can be classified with respect to responses to static and dynamic parameters of passive limb movements. Based on this classification it appears that the distribution of responses to static and dynamic parameters depends on the computational level within the cerebellar cortex. If both static and dynamic parameters are conveyed by a single unit, a clear relationship between the response components could be observed. However, this effect was independently found at all cerebellar cortical computational levels indicating a functional principle of processing a pair of movement parameters. The interrelation of complex- and simple spike responses to passive movement was further studied. Since transients of complex- and simple spike patterns were observed ranging from two almost identical patterns to mirror image like patterns, it is assumed that under physiological conditions one of the tasks of the climbing fibre system consists of tuning the simple spike discharge according to the peripheral requirements.     

Proprioceptive modulation of hip flexor activity during the swing phase of locomotion in decerebrate cats

This study examined the influence of proprioceptive input from hip flexor muscles on the activity in hip flexors during the swing phase of walking in the decerebrate cat. One hindlimb was partially denervated to remove cutaneous input and afferent input from most other hindlimb muscles. Perturbations to hip movement were applied either by 1) manual resistance or assistance to swing or by 2) resistance to hip flexion using a device that blocked hip flexion but allowed leg extension. Electromyographic recordings were made from the iliopsoas (IP), sartorius, and medial gastrocnemius muscles. When the hip was manually assisted into flexion, there was a reduction in hip flexor burst activity. Conversely, when hip flexion was manually resisted or mechanically blocked during swing, the duration and amplitude of hip flexor activity was increased. We also found some specificity in the role of afferents from individual hip flexor muscles in the modulation of flexor burst activity. If the IP muscle was detached from its insertion, little change in the response to blocking flexion was observed. Specific activation of IP afferent fibers by stretching the muscle also did not greatly affect flexor activity. On the other hand, if conduction in the sartorius nerves was blocked, there was a diminished response to blocking hip flexion. The increase in duration of the flexor bursts still occurred, but this increase was consistently lower than that observed when the sartorius nerves were intact. From these results we propose that during swing, feedback from hip flexor muscle afferents, particularly those from the sartorius muscles, enhances flexor activity. In addition, if we delayed the onset of flexor activity in the contralateral hindlimb, blocking hip flexion often resulted in the prolongation of ipsilateral flexor activity for long periods of time, further revealing the reinforcing effects of flexor afferent feedback on flexor activity. This effect was not seen if conduction in the sartorius nerves was blocked. In conclusion, we have found that hip flexor activity during locomotion can be strongly modulated by modifying proprioceptive feedback from the hip flexor muscles.     

Short-term effects of muscular denervation and fasciotomy on global limb variables during locomotion in the decerebrate cat

The motor system is capable of preserving the trajectories during locomotion of task level variables such as limb length and limb orientation in the face of paralysis of major muscle groups. This compensation is accomplished by the adjustment of the kinematics of joints other than the one most affected by the paralysis. The conservation of these task level variables could be accomplished quickly by feedback regulation or intrinsic mechanics, or by a longer-term adaptive process. We investigated the immediate effects of denervation of the triceps surae muscles in one limb of stepping, decerebrate cats to determine whether task level variables were preserved by short-term regulatory or intrinsic mechanisms. We further investigated the effects of disruption of the crural fascia in conjunction with denervation of the triceps surae muscles to determine whether the system consisting of multi-articular muscles of the thigh and crural fascia provided some contribution toward the preservation of limb length and orientation. Denervation led to substantial increases in ankle yield during stance, as previously observed, but also to significant decreases in limb length during early stance. Disruption of the crural fascia did not lead to increased ankle yield but, instead, to evidence for decreased propulsion. The results suggest that the preservation of task level variables observed in other studies does not result from online error correction or intrinsic properties of the musculoskeletal system but, by inference, from longer-term neural adaptation.     

Pontine respiratory group neuron discharge is altered during fictive cough in the decerebrate cat

A network of neurons in the rostral dorsal lateral pons and pons/mescencephalic junction constitute the pontine respiratory group (PRG) and is essential for reflex cough. As a next step in understanding the role of the PRG in the expression of the cough reflex, we examined neuron firing rates during fictive cough in cats. Decerebrated, thoracotomized, paralyzed, cycle-triggered ventilated adult cats were used. Extracellular activity of many single neurons and phrenic and lumbar neurograms were monitored during fictive cough produced by mechanical stimulation of the intrathoracic trachea. Neurons were tested during control periods for respiratory modulation of firing rate by cycle-triggered histograms and statistical tests. Most respiratory modulated cells were continuously active with various superimposed respiratory patterns; major categories included inspiratory decrementing (I-Dec), expiratory decrementing (E-Dec) and expiratory augmenting (E-Aug). There were alterations in the discharge patterns of respiratory, as well as, non-respiratory modulated neurons during cough. The results suggest an involvement of the PRG in the configuration of the cough motor pattern.     

Simple spike discharge patterns of Purkinje cells in the paramedian lobule of the cerebellum during locomotion in the awake cat

Purkinje cells in the paravermal cortex of the paramedian lobule in the cerebellum of awake cats have been recorded during locomotion on a moving belt. The discharges of all 35 recorded cells exhibited frequency modulation that was time-locked to the step cycle. Most cells showed increased activity during late flexion or at the onset of the stance phase of the step in the ipsilateral forelimb. This observation is consistent with the hypothesis that the role played by the paravermal cortex during locomotion is in the coordination and timing of the transition between the stance and swing phases of the step.     

Resetting of resultant stiffness in ankle flexor and extensor muscles in the decerebrate cat

Summary Flexor (tibialis anterior, TA, and extensor digitorum longus, EDL) and extensor (soleus, SOL) muscles in the decerebrate cat were subjected to length changes and the force responses were measured. Resultant muscular stiffness, which arises from the mechanical reaction of muscle fibers contracting prior to the length change and from a change in force due to reflex action, was calculated by dividing the changes in force by the corresponding length changes. As shown previously in the premammillary preparation, resultant stiffness was usually higher in SOL than in TA or EDL. Following an intercollicular transection in some preparations, resultant stiffness increased markedly for TA but not substantially for SOL. During continuous electrical stimulation in the magnocellular red nucleus in premammillary preparations, resultant stiffness of SOL decreased for a wide range of forces while EDL responses were unaffected. These results show that reflex gain is not determined solely by the level of motoneuronal excitability but also by a descending control from the brainstem, and that the lower resultant stiffness in flexors compared to extensors in the decerebrate cat is set by this control system and not by inherent differences in the strength of autogenetic reflex pathways for the two muscles.     

Last-order interneurones controlling activity of elbow flexor motoneurones during forelimb fictive locomotion in the cat

Activity of C5-C7 last-order interneurones, which were identified by antidromic invasion from the elbow flexor motor nuclei, was examined during fictive locomotion. Three groups of neurones were found: one showing no rhythmic modulation of activity (non-modulated neurones), another rhythmically active mainly in phase with the target motoneurones (modulated neurones of group 1), the other out of phase with them (modulated neurones of group 2). Activity pattern suggested group 1 and group 2 neurones distributing excitation and inhibition to flexor motoneurones, respectively. Location of neurones seemed to be differentiated as well, group 1 located dorsally, group 2 ventrally in the gray matter.     

Tonic neck reflex of the decerebrate cat: a role for propriospinal neurons

In decerebrate, acutely labyrinthectomized cats we used neck rotation to study the role of direct upper cervical afferents to the cervical enlargement and of cervical and lumbar propriospinal neurons in the tonic neck reflex. Interruption of the dorsal columns between C4 and C5 had no qualitative effect on the dynamics of the reflex although gain usually increased. Direct upper cervical afferents to the cervical enlargement therefore have no unique role in producing the reflex. Many medially located propriospinal neurons in C4 were modulated by neck rotation. About 40% had axons, mostly crossed, that terminated in the cervical enlargement. The others projected more caudally, some as far as L3-L4 or even the lumbar enlargement. For a population of C4 neurons, including propriospinal neurons, we measured the response vector with combinations of roll and pitch stimuli. These vectors ranged from pitch to roll. Many propriospinal neurons in L3-L4, projecting to the lumbosacral enlargement, were also modulated by neck rotation with a variety of response vectors. Some of these neurons had an ascending projection. As in previous experiments, C4 neurons were modulated by neck rotation after spinal transection rostral to the C1 dorsal root entry zone; a wide variety of response vectors was observed. In contrast, almost no modulated L3-L4 neurons were found in the same experiments. The results suggest a role for propriospinal neurons in the tonic neck reflex. They also demonstrate that responses of lumbar neurons to neck rotation are much more dependent on supraspinal pathways than are those of cervical neurons.