Modular neuromuscular control of human locomotion by central pattern generator

The central pattern generators (CPG) in the spinal cord are thought to be responsible for producing the rhythmic motor patterns during rhythmic activities. For locomotor tasks, this involves much complexity, due to a redundant system of muscle actuators with a large number of highly nonlinear muscles. This study proposes a reduced neural control strategy for the CPG, based on modular organization of the co-active muscles, i.e., muscle synergies. Four synergies were extracted from the EMG data of the major leg muscles of two subjects, during two gait trials each, using non-negative matrix factorization algorithm. A Matsuoka׳s four-neuron CPG model with mutual inhibition, was utilized to generate the rhythmic activation patterns of the muscle synergies, using the hip flexion angle and foot contact force information from the sensory afferents as inputs. The model parameters were tuned using the experimental data of one gait trial, which resulted in a good fitting accuracy (RMSEs between 0.0491 and 0.1399) between the simulation and experimental synergy activations. The model׳s performance was then assessed by comparing its predictions for the activation patterns of the individual leg muscles during locomotion with the relevant EMG data. Results indicated that the characteristic features of the complex activation patterns of the muscles were well reproduced by the model for different gait trials and subjects. In general, the CPG- and muscle synergy-based model was promising in view of its simple architecture, yet extensive potentials for neuromuscular control, e.g., resolving redundancies, distributed and fast control, and modulation of locomotion by simple control signals.     

Possible contributions of CPG activity to the control of rhythmic human arm movement

There is extensive modulation of cutaneous and H-reflexes during rhythmic leg movement in humans. Mechanisms controlling reflex modulation (e.g., phase- and task-dependent modulation, and reflex reversal) during leg movements have been ascribed to the activity of spinal central pattern generating (CPG) networks and peripheral feedback. Our working hypothesis has been that neural mechanisms (i.e., CPGs) controlling rhythmic movement are conserved between the human lumbar and cervical spinal cord. Thus reflex modulation during rhythmic arm movement should be similar to that for rhythmic leg movement. This hypothesis has been tested by studying the regulation of reflexes in arm muscles during rhythmic arm cycling and treadmill walking. This paper reviews recent studies that have revealed that reflexes in arm muscles show modulation within the movement cycle (e.g., phase-dependency and reflex reversal) and between static and rhythmic motor tasks (e.g., task-dependency). It is concluded that reflexes are modulated similarly during rhythmic movement of the upper and lower limbs, suggesting similar motor control mechanisms. One notable exception to this pattern is a failure of contralateral arm movement to modulate reflex amplitude, which contrasts directly with observations from the leg. Overall, the data support the hypothesis that CPG activity contributes to the neural control of rhythmic arm movement.     

Gait acts as a gate for reflexes from the foot

During human gait, electrical stimulation of the foot elicits facilitatory P2 (medium latency) responses in TA (tibialis anterior) at the onset of the swing phase, while the same stimuli cause suppressive responses at the end of swing phase, along with facilitatory responses in antagonists. This phenomenon is called phase-dependent reflex reversal. The suppressive responses can be evoked from a variety of skin sites in the leg and from stimulation of some muscles such as rectus femoris (RF). This paper reviews the data on reflex reversal and adds new data on this topic, using a split-belt paradigm. So far, the reflex reversal in TA could only be studied for the onset and end phases of the step cycle, simply because suppression can only be demonstrated when there is background activity. Normally there are only 2 TA bursts in the step cycle, whereas TA is normally silent during most of the stance phase. To know what happens in the stance phase, one needs to have a means to evoke some background activity during the stance phase. For this purpose, new experiments were carried out in which subjects were asked to walk on a treadmill with a split-belt. When the subject was walking with unequal leg speeds, the walking pattern was adapted to a gait pattern resembling limping. The TA then remained active throughout most of the stance phase of the slow-moving leg, which was used as the primary support. This activity was a result of coactivation of agonistic and antagonistic leg muscles in the supporting leg, and represented one of the ways to stabilize the body. Electrical stimulation was given to a cutaneous nerve (sural) at the ankle at twice the perception threshold. Nine of the 12 subjects showed increased TA activity during stance phase while walking on split-belts, and 5 of them showed pronounced suppressions during the first part of stance when stimuli were given on the slow side. It was concluded that a TA suppressive pathway remains open throughout most of the stance phase in the majority of subjects. The suggestion was made that the TA suppression increases loading of the ankle plantar flexors during the loading phase of stance.     

Mechanisms of abdominal muscle activation during vomiting

The possible contribution of spinal reflexes to abdominal muscle activation during vomiting was assessed in decerebrate cats. The activity of these muscles is partly controlled by bulbospinal expiratory neurons in the caudal ventral respiratory group (VRG). In a previous study it was found that the abdominal muscles are still active during vomiting after midsagittal lesion of the axons of these neurons between C1 and the obex (A.D. Miller, L.K. Tan, and I. Suzuki. J. Neurophysiol. 57: 1854-1866, 1987). The present experiments indicate that this postlesion activity was due to spinal stretch reflexes because 1) such midsagittal lesions eliminate abdominal muscle nerve activity during fictive vomiting in paralyzed cats in which there are no abdominal stretch reflexes, 2) the abdominal muscles are activated during vomiting by spinal reflexes after upper thoracic cord transections, and 3) the normal 100-ms delay between diaphragmatic and abdominal activation during vomiting is reduced to approximately 20-25 ms after both types of lesions, which is consistent with postlesion abdominal reflex activation. Our results also suggest that, during normal vomiting, abdominal stretch and tension reflexes have only a minor role if any and abdominal muscle activation is probably mediated primarily or exclusively by expiratory neurons in the caudal ventral respiratory group. However, our finding that phrenic activity is reduced both during vomiting after thoracic transections and during fictive vomiting after paralysis is consistent with a contribution of reflex activity from abdominal and/or intercostal muscles to phrenic discharge during normal vomiting.     

Reflex control of posterior shoulder muscles from arm afferents in healthy people

In order to position the hand during functional tasks, control of the shoulder is required. Heteronymous reflexes from the upper limb to shoulder muscles are used to assist in this control. To investigate this further, the radial and ulnar nerves were stimulated at elbow level whilst surface electromyographic activity of posterior deltoid, infraspinatus and latissimus dorsi muscles were recorded. In addition, the cutaneous branch of the radial nerve and the skin of the fifth digit were stimulated in order to investigate any cutaneous contribution to reflex activity. Reflexes were evoked in all three of these shoulder muscles from hand and/or forearm afferents. However, the reflexes differed; whereas both excitatory and inhibitory reflexes were evoked in posterior deltoid and infraspinatus, the reflexes in latissimus dorsi were mainly excitatory. Cutaneomuscular reflexes were seldom evoked here, but when they were present they were generally evoked at longer latencies than the reflexes evoked by mixed nerve stimulation. The results suggest a role for reflexes originating from the forearm and/or hand in the control of the shoulder.     

Contribution of stretch reflexes to locomotor control: a modeling study

It is known that the springlike properties of muscles provide automatic load compensation during weight bearing. How crucial is sensory control of the motor output given these basic properties of the locomotor system? To address this question, a neuromuscular model was used to test two hypotheses. (1) Stretch reflexes are too weak and too delayed to contribute significantly to weight-bearing. (2) The important contributions of sensory input involve state-dependent processing. We constructed a two-legged planar locomotor model with 9 segments, driven by 12 musculotendon actuators with Hill-type force-velocity and monotonic force-length properties. Electromyographic (EMG) profiles of the simulated muscle groups during slow level walking served as actuator activation functions. Spindle Ia and tendon organ Ib sensory inputs were represented by transfer functions with a latency of 35 ms, contributing 30% to the net EMG profile and gated to be active only when the receptor-bearing muscles were contracting. Locomotor stability was assessed by parametric variations of actuator maximum forces during locomotion in open-loop ("deafferented") trials and in trials with feedback control based on either sensory-evoked stretch reflexes or finite-state rules. We arrived at the following conclusions. (1) In the absence of sensory control, the intrinsic stiffness of limb muscles driven by a stereotyped rhythmical pattern can produce surprisingly stable gait. (2) When the level of central activity is low, the contribution of stretch reflexes to load compensation can be crucial. However, when central activity provides adequate load compensation, the contribution of stretch reflexes is less significant. (3) Finite-state control can greatly extend the adaptive capability of the locomotor system.     

A test of a dual central pattern generator hypothesis for subcortical control of locomotion.

This study was designed to examine the nature of neural circuits involved in subcortical inter-limb coordination and reflex modulation mechanisms of locomotion. These circuits, called central pattern generators (CPGs), are believed to receive tonic input and generate rhythmically alternating sets of commands. Although CPGs have been theorized to exist in humans, their potential dual role in inter-limb coordination and reflex modulation is unclear. In the present study, nine participants walked on a treadmill, timing their heel-strikes to a metronome which varied the phase lag from 0.5 to 1.0 pi radians (0.1 pi intervals). A stimulus was delivered to the sural nerve and reflexes were measured in the ipsilateral and contralateral lower extremities through electromyography. The similarity between phase lag conditions for both temporal coordination (i.e., relative timing aspects between muscles and/or limbs) and reflex intensities suggested that they may be controlled by the same subcortical circuitry. Two plausible explanations exist: (1) a single CPG coordinates muscular contractions and phasically alters proprioceptive reflex modulation, as well as cutaneous input, using feed-forward control; (2) two separate circuits are strongly entrained, producing synchronous outputs for inter-limb coordination and reflex modulation. The out-of-phase task used in this study was limited in discerning such a difference, if it exists.     

Can Treadmill Perturbations Evoke Stretch Reflexes in the Calf Muscles?

Disinhibition of reflexes is a problem amongst spastic patients, for it limits a smooth and efficient execution of motor functions during gait. Treadmill belt accelerations may potentially be used to measure reflexes during walking, i.e. by dorsal flexing the ankle and stretching the calf muscles, while decelerations show the modulation of reflexes during a reduction of sensory feedback. The aim of the current study was to examine if belt accelerations and decelerations of different intensities applied during the stance phase of treadmill walking can evoke reflexes in the gastrocnemius, soleus and tibialis anterior in healthy subjects. Muscle electromyography and joint kinematics were measured in 10 subjects. To determine whether stretch reflexes occurred, we assessed modelled musculo-tendon length and stretch velocity, the amount of muscle activity, as well as the incidence of bursts or depressions in muscle activity with their time delays, and co-contraction between agonist and antagonist muscle. Although the effect on the ankle angle was small with 2.8±1.0°, the perturbations caused clear changes in muscle length and stretch velocity relative to unperturbed walking. Stretched muscles showed an increasing incidence of bursts in muscle activity, which occurred after a reasonable electrophysiological time delay (163–191 ms). Their amplitude was related to the muscle stretch velocity and not related to co-contraction of the antagonist muscle. These effects increased with perturbation intensity. Shortened muscles showed opposite effects, with a depression in muscle activity of the calf muscles. The perturbations only slightly affected the spatio-temporal parameters, indicating that normal walking was retained. Thus, our findings showed that treadmill perturbations can evoke reflexes in the calf muscles and tibialis anterior. This comprehensive study could form the basis for clinical implementation of treadmill perturbations to functionally measure reflexes during treadmill-based clinical gait analysis.     

Analysis of the gait generation principle by a simulated quadruped model with a CPG incorporating vestibular modulation

This study aims to understand the principles of gait generation in a quadrupedal model. It is difficult to determine the essence of gait generation simply by observation of the movement of complicated animals composed of brains, nerves, muscles, etc. Therefore, we build a planar quadruped model with simplified nervous system and mechanisms, in order to observe its gaits under simulation. The model is equipped with a mathematical central pattern generator (CPG), consisting of four coupled neural oscillators, basically producing a trot pattern. The model also contains sensory feedback to the CPG, measuring the body tilt (vestibular modulation). This spontaneously gives rise to an unprogrammed lateral walk at low speeds, a transverse gallop while running, in addition to trotting at a medium speed. This is because the body oscillation exhibits a double peak per leg frequency at low speeds, no peak (little oscillation) at medium speeds, and a single peak while running. The body oscillation autonomously adjusts the phase differences between the neural oscillators via the feedback. We assume that the oscillations of the four legs produced by the CPG and the body oscillation varying according to the current speed are synchronized along with the varied phase differences to keep balance during locomotion through postural adaptation via the vestibular modulation, resulting in each gait. We succeeded in determining a single simple principle that accounts for gait transition from walking to trotting to galloping, even without brain control, complicated leg mechanisms, or a flexible trunk.     

Motor unit recruitment when neuromuscular electrical stimulation is applied over a nerve trunk compared with a muscle belly: triceps surae

Neuromuscular electrical stimulation (NMES) can be delivered over a nerve trunk or muscle belly and can generate contractions by activating motor (peripheral pathway) and sensory (central pathway) axons. In the present experiments, we compared the peripheral and central contributions to plantar flexion contractions evoked by stimulation over the tibial nerve vs. the triceps surae muscles. Generating contractions through central pathways follows Henneman's size principle, whereby low-threshold motor units are activated first, and this may have advantages for rehabilitation. Statistical analyses were performed on data from trials in which NMES was delivered to evoke 10-30% maximum voluntary torque 2-3 s into the stimulation (Time(1)). Two patterns of stimulation were delivered: 1) 20 Hz for 8 s; and 2) 20-100-20 Hz for 3-2-3 s. Torque and soleus electromyography were quantified at the beginning (Time(1)) and end (Time(2); 6-7 s into the stimulation) of each stimulation train. H reflexes (central pathway) and M waves (peripheral pathway) were quantified. Motor unit activity that was not time-locked to each stimulation pulse as an M wave or H reflex ("asynchronous" activity) was also quantified as a second measure of central recruitment. Torque was not different for stimulation over the nerve or the muscle. In contrast, M waves were approximately five to six times smaller, and H reflexes were approximately two to three times larger during NMES over the nerve vs. the muscle. Asynchronous activity increased by 50% over time, regardless of the stimulation location or pattern, and was largest during NMES over the muscle belly. Compared with NMES over the triceps surae muscles, NMES over the tibial nerve produced contractions with a relatively greater central contribution, and this may help reduce muscle atrophy and fatigue when NMES is used for rehabilitation.     

Convergence in Reflex Pathways from Multiple Cutaneous Nerves Innervating the Foot Depends upon the Number of Rhythmically Active Limbs during Locomotion

Neural output from the locomotor system for each arm and leg influences the spinal motoneuronal pools directly and indirectly through interneuronal (IN) reflex networks. While well documented in other species, less is known about the functions and features of convergence in common IN reflex system from cutaneous afferents innervating different foot regions during remote arm and leg movement in humans. The purpose of the present study was to use spatial facilitation to examine possible convergence in common reflex pathways during rhythmic locomotor limb movements. Cutaneous reflexes were evoked in ipsilateral tibialis anterior muscle by stimulating (in random order) the sural nerve (SUR), the distal tibial nerve (TIB), and combined simultaneous stimulation of both nerves (TIB&SUR). Reflexes were evoked while participants performed rhythmic stepping and arm swinging movement with both arms and the leg contralateral to stimulation (ARM&LEG), with just arm movement (ARM) and with just contralateral leg movement (LEG). Stimulation intensities were just below threshold for evoking early latency (<80 ms to peak) reflexes. For each stimulus condition, rectified EMG signals were averaged while participants held static contractions in the stationary (stimulated) leg. During ARM&LEG movement, amplitudes of cutaneous reflexes evoked by combined TIB&SUR stimulation were significantly larger than simple mathematical summation of the amplitudes evoked by SUR or TIB alone. Interestingly, this extra facilitation seen during combined nerve stimulation was significantly reduced when performing ARM or LEG compared to ARM&LEG. We conclude that locomotor rhythmic limb movement induces excitation of common IN reflex pathways from cutaneous afferents innervating different foot regions. Importantly, activity in this pathway is most facilitated during ARM&LEG movement. These results suggest that transmission in IN reflex pathways is weighted according to the number of limbs directly engaged in human locomotor activity and underscores the importance of arm swing to support neuronal excitability in leg muscles.     

Assessment of the activation modalities of gastrocnemius lateralis and tibialis anterior during gait: A statistical analysis

Aim of the study was to identify the different modalities of activation of gastrocnemius lateralis (GL) and tibialis anterior (TA) during gait at self-selected speed, by a statistical analysis of surface electromyographic signal from a large number (hundreds) of strides per subject. The analysis on fourteen healthy adults showed a large variability in the number of activation intervals, in their occurrence rate, and in the on-off instants, within different strides of the same walk. For each muscle, the assessment of the different modalities of activation (five for muscle) allowed to identify a single pattern, common for all the modalities and able to characterize the behavior of muscles during normal gait. The pattern of GL activity centered in two regions of the gait cycle: the transition between flat foot contact and push-off (observed in 100% of total strides) and the final swing (67.1 ± 15.9%). Two regions characterized also the pattern of TA activity: from pre-swing to following loading response (100%), and the mid-stance (30.5 ± 15.0%). This “normality” pattern represents the first attempt for the development in healthy young adults of a reference for dynamic EMG activity of GL and TA, in terms of variability of on-off muscular activity and occurrence rate during gait.     

Long loop reflexes: concepts and consequences

New experimental results on long latency reflexes in human limb muscles were presented by neurophysiologists and clinicians with great experience in this field. The scope of presentations reached from the pattern of long latency reflexes in leg muscles during stance and the modifications in various motor disorders to the firing behavior of single motor units during torque perturbations of the elbow and to the role of habituation in long latency reflexes. Finally, the effect of stereoencephalotomy on long latency reflexes in normals and patients with parkinsonian tremor was reviewed.     

Interlimb Reflexes Induced by Electrical Stimulation of Cutaneous Nerves after Spinal Cord Injury

Whether interlimb reflexes emerge only after a severe insult to the human spinal cord is controversial. Here the aim was to examine interlimb reflexes at rest in participants with chronic (>1 year) spinal cord injury (SCI, n = 17) and able-bodied control participants (n = 5). Cutaneous reflexes were evoked by delivering up to 30 trains of stimuli to either the superficial peroneal nerve on the dorsum of the foot or the radial nerve at the wrist (5 pulses, 300 Hz, approximately every 30 s). Participants were instructed to relax the test muscles prior to the delivery of the stimuli. Electromyographic activity was recorded bilaterally in proximal and distal arm and leg muscles. Superficial peroneal nerve stimulation evoked interlimb reflexes in ipsilateral and contralateral arm and contralateral leg muscles of SCI and control participants. Radial nerve stimulation evoked interlimb reflexes in the ipsilateral leg and contralateral arm muscles of control and SCI participants but only contralateral leg muscles of control participants. Interlimb reflexes evoked by superficial peroneal nerve stimulation were longer in latency and duration, and larger in magnitude in SCI participants. Interlimb reflex properties were similar for both SCI and control groups for radial nerve stimulation. Ascending interlimb reflexes tended to occur with a higher incidence in participants with SCI, while descending interlimb reflexes occurred with a higher incidence in able-bodied participants. However, the overall incidence of interlimb reflexes in SCI and neurologically intact participants was similar which suggests that the neural circuitry underlying these reflexes does not necessarily develop after central nervous system injury.     

Modulation of the human nociceptive reflex by cyclic movements

During static conditions the nociceptive reflex is known to vary as a function of, for example, the stimulus position, stimulus intensity, and muscle contraction. The aim of the present human study was to investigate whether the reflex and the corresponding perception of pain are modulated by cyclic movements of the limb involved. Reflexes, evoked by nociceptive electric stimulation of the sural nerve, were recorded from the biceps femoris and the rectus femoris muscles in eight volunteers. Four different experiments were performed to compare the nociceptive reflex and pain score elicited during active isometric/dynamic flexion/extension of the knee joint. The amplitudes of the reflexes were largest for the dynamic conditions. The reflexes, evoked during dynamic extension and isometric contraction of the rectus femoris muscle, had the shortest latencies but the recordings from the biceps femoris muscle were larger than from the rectus femoris muscle. Knee joint angle recordings showed that the largest angle variations occurred for the dynamic conditions and were only marginally disturbed for the isometric conditions. A given stimulus intensity evoked the highest pain intensity during isometric contractions. This indicates that there would seem to be no causal relationship between the size of the nociceptive reflex and the pain intensity.     

Artificial neural network model for the generation of muscle activation patterns for human locomotion.

Skilled locomotor behaviour requires information from various levels within the central nervous system (CNS). Mathematical models have permitted researchers to simulate various mechanisms in order to understand the organization of the locomotor control system. While it is difficult to adequately characterize the numerous inputs to the locomotor control system, an alternative strategy may be to use a kinematic movement plan to represent the complex inputs to the locomotor control system based on the possibility that the CNS may plan movements at a kinematic level. We propose the use of artificial neural network (ANN) models to represent the transformation of a kinematic plan into the necessary motor patterns. Essentially, kinematic representation of the actual limb movement was used as the input to an ANN model which generated the EMG activity of 8 muscles of the lower limb and trunk. Data from a wide variety of gait conditions was necessary to develop a robust model that could accommodate various environmental conditions encountered during everyday activity. A total of 120 walking strides representing normal walking and ten conditions where the normal gait was modified in terms of cadence, stride length, stance width or required foot clearance. The final network was assessed on its ability to predict the EMG activity on individual walking trials as well as its ability to represent the general activation pattern of a particular gait condition. The predicted EMG patterns closely matched those recorded experimentally, exhibiting the appropriate magnitude and temporal phasing required for each modification. Only 2 of the 96 muscle/gait conditions had RMS errors above 0.10, only 5 muscle/gait conditions exhibited correlations below 0.80 (most were above 0.90) and only 25 muscle/gait conditions deviated outside the normal range of muscle activity for more than 25% of the gait cycle. These results indicate the ability of single network ANNs to represent the transformation between a kinematic movement plan and the necessary muscle activations for normal steady state locomotion but they were also able to generate muscle activation patterns for conditions requiring changes in walking speed, foot placement and foot clearance. The abilities of this type of network have implications towards both the fundamental understanding of the control of locomotion and practical realizations of artificial control systems for use in rehabilitation medicine.     

Cellular properties and modulation of the stomatogastric ganglion neurons of a stomatopod,Squilla oratoria

Cellular properties and modulation of the identified neurons of the posterior cardiac plate-pyloric system in the stomatogastric ganglion of a stomatopod, Squilla oratoria, were studied electrophysiologically. Each class of neurons involved in the cyclic bursting activity was able to trigger an endogenous, slow depolarizing potential (termed a driver potential) which sustained bursting. Endogenous oscillatory properties were demonstrated by the phase reset behavior in response to brief stimuli during ongoing rhythm. The driver potential was produced by membrane voltage-dependent activation and terminated by an active repolarization. Striking enhancement of bursting properties of all the cell types was induced by synaptic activation via extrinsic nerves, seen as increases in amplitude or duration of driver potentials, spiking rate during a burst, and bursting rate. The motor pattern produced under the influence of extrinsic modulatory inputs continued for a long time, relative to that in the absence of activation of modulatory inputs. Voltage-dependent conductance mechanisms underlying postinhibitory rebound and driver potential responses were modified by inputs. It is concluded that endogenous cellular properties, as well as synaptic circuitry and extrinsic inputs, contribute to generation of the rhythmic motor pattern, and that a motor system and its component neurons have been highly conserved during evolution between stomatopods and decapods.Abbreviations AB anterior burster neuron - CoG commissural ganglion - CPG central pattern generator - lvn lateral ventricular nerve - OG oesophageal ganglion - pcp posterior cardiac plate - PCP pcp constrictor neuron - PD pyloric dilator neuron - PY pyloric constrictor neuron - son superior oesophageal nerve - STG stomatogastric ganglion - stn stomatogastric nerve     

The contribution of sensory inputs to the pattern generation of breathing

Current concepts of the basic neural control system and its modulation by afferent inputs are reviewed. It is emphasized that, in analogy with locomotion, the central pattern generator (CPG) for automatic metabolic respiration does not depend on any afferent feedback from receptors sensitive to the movements of the "pump," or the streams of pumped air, for its production of a rhythmic motor output provides the CPG receives some "drive" inputs above threshold and adequate bias. The operation for a variety of reflexes and feedback loops is of fundamental importance, however, for adapting the breathing pattern to the varying requirements for gas exchange to the many behavioural, nonmetabolic demands on the breathing apparatus which are competing with its primary metabolic control functions. The presentation is focussed also on available evidence that the respiratory CPG exerts powerful modulations on the transmission in these reflex pathways controlling the pattern of breathing and adjusting it to the various metabolic and behavioural demands. Mechanisms for "gating," "phasic gain changes," and "phase-dependent reflex reversal" are exemplified.     

Tactile Modulation of Whisking via the Brainstem Loop: Statechart Modeling and Experimental Validation

Rats repeatedly sweep their facial whiskers back and forth in order to explore their environment. Such explorative whisking appears to be driven by central pattern generators (CPGs) that operate independently of direct sensory feedback. Nevertheless, whisking can be modulated by sensory feedback, and it has been hypothesized that some of this modulation already occurs within the brainstem. However, the interaction between sensory feedback and CPG activity is poorly understood. Using the visual language of statecharts, a dynamic, bottom-up computerized model of the brainstem loop of the whisking system was built in order to investigate the interaction between sensory feedback and CPG activity during whisking behavior. As a benchmark, we used a previously quantified closed-loop phenomenon of the whisking system, touched-induced pump (TIP), which is thought to be mediated by the brainstem loop. First, we showed that TIPs depend on sensory feedback, by comparing TIP occurrence in intact rats with that in rats whose sensory nerve was experimentally cut. We then inspected several possible feedback mechanisms of TIPs using our model. The model ruled out all hypothesized mechanisms but one, which adequately simulated the corresponding motion observed in the rat. Results of the simulations suggest that TIPs are generated via sensory feedback that activates extrinsic retractor muscles in the mystacial pad. The model further predicted that in addition to the touching whisker, all whiskers found on the same side of the snout should exhibit a TIP. We present experimental results that confirm the predicted movements in behaving rats, establishing the validity of the hypothesized interaction between sensory feedback and CPG activity we suggest here for the generation of TIPs in the whisking system.     

Inspiratory and expiratory patterns of the pectoralis major muscle during pulmonary defensive reflexes

Experiments wereconducted to determine the discharge pattern of the pectoralis majormuscle during pulmonary defensive reflexes in anesthetized cats(n = 15). Coughs andexpiration reflexes were elicited by mechanical stimulation of theintrathoracic trachea or larynx. Augmented breaths occurredspontaneously or were evoked by the same mechanical stimuli.Electromyograms (EMGs) were recorded from the diaphragm, rectusabdominis, and pectoralis major muscles. During augmented breaths, thepectoralis major had inspiratory EMG activity similar to that of thediaphragm, but during expiration reflexes the pectoralis major also hadpurely expiratory EMG activity similar to the rectus abdominis. Duringtracheobronchial cough, the pectoralis major had an inspiratory patternsimilar to that of the diaphragm in 10 animals, an expiratory patternsimilar to that of the rectus abdominis in 3 animals, and a biphasicpattern in 2 animals. The pectoralis major was active during both the inspiratory and expiratory phases during laryngeal cough. We conclude that, in contrast to the diaphragm or rectus abdominis muscles, thepectoralis major is active during both inspiratory and expiratory pulmonary defensive reflexes.