Premotor System Research Articles

Distributed Neural Systems Underlying the Timing of Movements

Timing is essential to the execution of skilled movements, yet our knowledge of the neural systems underlying timekeeping operations is limited. Using whole-brain functional magnetic resonance imaging, subjects were imaged while tapping with their right index finger in synchrony with tones that were separated by constant intervals [Synchronization (S)], followed by tapping without the benefit of an auditory cue [Continuation (C)]. Two control conditions followed in which subjects listened to tones and then made pitch discriminations (D). Both the S and the C conditions produced equivalent activation within the left sensorimotor cortex, the right cerebellum (dorsal dentate nucleus), and the right superior temporal gyrus (STG). Only the C condition produced activation of a medial premotor system, including the caudal supplementary motor area (SMA), the left putamen, and the left ventrolateral thalamus. The C condition also activated a region within the right inferior frontal gyrus (IFG), which is functionally interconnected with auditory cortex. Both control conditions produced bilateral activation of the STG, and the D condition also activated the rostral SMA. These results suggest that the internal generation of precisely timed movements is dependent on three interrelated neural systems, one that is involved in explicit timing (putamen, ventrolateral thalamus, SMA), one that mediates auditory sensory memory (IFG, STG), and another that is involved in sensorimotor processing (dorsal dentate nucleus, sensorimotor cortex).

Chapter 34 Cerebellar somatotopic representation and cerebro-cerebellar interconnections in ataxic patients

Different methods of functional neuroimaging were used for studying somatotopic encoding of function in the cerebellum and for investigating cerebro-cerebellar interconnections in patients with cerebellar degeneration. fMRI showed, that the center of activation for hand function was located in the intermediate hemispheric portion of Larsell lobules H IV-V. Foot movements activated areas medial and anterior to the corresponding hand areas within Larsell lobules II-III. Changed function in motor cortices could be demonstrated in patients with cerebellar degeneration as compared to normal controls by recording movement-related cortical potentials (BP). In patients the motor potential was almost lacking and transcranial magnetic stimulation demonstrated enhancement of inhibitory mechanisms (prolonged postexcitatory inhibition) in the motor cortex. PET-findings suggested, that both effects are correlated to increased activity of inhibitory interneurons. Cerebellar patients showed increased activation in relation to movements in the SMA and basal ganglia and reduced activation in the cerebellum and lateral premotor areas. It could be speculated, that compensatory mechanisms are the reason for a stronger activation of the medial premotor system, including SMA, in patients with cerebellar degeneration. On the basis of our results it appears, that the cerebellum facilitates the lateral premotor system areas much more than it does the medial areas.

Regional cerebral blood flow during a self-paced sequential finger opposition task in patients with cerebellar degeneration.

The brain regions controlling self-paced sequential finger movements in patients with cerebellar degeneration were studied by measuring changes in regional cerebral blood flow (rCBF) in eight patients using bolus injections of H2(15)O and PET. The results were compared with those obtained in eight normal age-matched control subjects. Patients and control subjects performed a self-paced sequential finger opposition task with the right hand, completing a sequence of movements every 4-6 s. Both groups had strong increases in the adjusted rCBF contralaterally in the primary motor cortex (M1) and ventral premotor area (PMv), in the caudal supplementary motor area (SMA) and cingulate motor area (CMA), and bilaterally in the prefrontal cortex (PFC), the lobus parietalis inferior (LPI), putamen and cerebellum. The cerebellum, PMv, rostral CMA, PFC and LPI were more active in the control subjects than in the patients, and the M1, SMA, caudal CMA and putamen were more active in the patients than in the control subjects. The reduced activity of the cerebellar neurons in the patients produced a complex pattern of rCBF increases and decreases in other brain regions. Our results suggest that for the preparation and execution of sequential finger movements, patients with cerebellar degeneration use a medial premotor system, including the SMA and caudal CMA, as well as the M1 and putamen, rather than the PMv, PFC, LPI and rostral CMA.

A model of the saccadic sensorimotor system of salamanders.

A model of the saccadic system of salamanders on the basis of electrophysiological and anatomical results is presented. The model includes centers found to be significant for the guidance of saccades in these comparatively simple vertebrates. In particular, these are the optic tectum, the bulbar reticular formation and the motor system. The latter consists of two pairs of neck-muscles, an epaxial and a hypaxial one driven by their respective motoneurons. The model includes a visual, a sensori-motor, and a motor level. At the sensory level, the retinal coordinates are transferred to the optic tectum to establish an orthogonal map of visual angles. A secondary visual map of the ipsilateral eye with a pointsymmetrical organization exists in addition. The premotor system of the tectum was modelled according to an ensemble-coding principle. Thus, local activation of the visual map results in recruitment of an appropriate number of tectal premotor units. Simulation of the model reproduces correct metric properties of salamander saccades under varying stimulus presentations.

Readiness potentials preceding spontaneous motor acts: voluntary vs. involuntary control

Libet et al. (1983) developed a method to compare the onset time of a readiness potential (RP) with the onset time of the corresponding intention to perform a spontaneous voluntary motor act. In relation to the onset of the RP, the time of conscious intention to move followed 350 msec later. From these results Libet (1985) concluded that the cerebral initiation of a spontaneous voluntary act begins unconsciously. We investigated the alternative interpretation that with the instruction to pay attention to feelings of ‘wanting to move’, automatic and normally unconscious motor acts might have been brought to a level of conscious awareness. Therefore we conducted 3 kinds of experiments. In the first, RPs were measured from subjects performing unconscious movements. The second experiment was a replication of Libet's study while the third was a resting condition in which subjects looked for intentions to move introspectively. The results showed that RPs beginning approximately 500 msec before movement onset can be obtained with unconsciously as well as consciously performed spontaneous motor acts. The different scalp distributions of the two types of RP indicate that unconscious movements can be attributed to the activation of a contralateral process (lateral premotor system (LPS), primary motor cortex), whereas voluntary spontaneous motor acts seemed to be predominated by the medial premotor system (MPS). It is proposed that in the Libet situation focused attention on internal events led to the conscious detection of a normally unconscious process. This resulted in the activation of the MPS, especially the supplementary motor area (SMA), which released the starting signal for the execution of the motor act. We believe that the activation of the SMA and the urge to move occurred at the same time.

The Alien Hand Sign

The alien hand sign was first described by Brion and Jedynak as a "feeling of estrangement between the patient and one of his hands." The affected hand frequently shows a grasp reflex and an instinctive grasp reaction as well as elements of what Denny-Brown referred to as a "magnetic apraxia" associated with frontal lobe damage. Most notably, however, the affected hand is observed to perform apparently purposive behaviors that are perceived as being outside the volitional control of the patient. The patients interpret the behavior of their own affected limb as being controlled by an external agent. They do not feel that they are initiating or controlling the behavior of the hand and often express dismay at the hand's "extravolitional" activity. The patients attempt to control behavior of the alien hand with the unimpaired hand by forcibly restraining the affected limb, an act that may be termed "self-restriction." In this paper, we report an additional four cases of alien hand sign in right-handed subjects: two involving the right hand and two involving the left hand. In each case, the clinical findings were associated with extensive unilateral damage of the medial frontal cortex of the hemisphere contralateral to the affected hand. Furthermore, the alien movement gradually disappears over the course of 6-12 months after the stroke. These clinical case studies are presented and discussed in the context of the "dual premotoer systems hypothesis," an anatomicophysiological model that proposes that action is organized by two separate but interactive premotor brain systems corresponding to evolutionarily defined medial and lateral cortical moieties. It is hypothesized that the alien mode behavior results from unconstrained activity of the lateral premotor system in the damaged hemisphere. The residual volitional control in the limb occurs through the activity of the intact medial premotor system of the ipsilateral hemisphere. Recovery may occur through extension of these ipsilateral control mechanisms by compensatory changes in subcortical systems controlling hemispheric activation associated with adaptive behavior. This observation may be important in understanding mechanisms involved in motor recovery after stroke.

Premotor systems, motor learning, and ipsilateral control: Learning to get set

Premotor systems, motor learning, and ipsilateral control: Learning to get set

Supplementary motor area structure and function: Review and hypotheses

AbstractThough its existence has been known for well over 30 years, only recently has the supplementary motor area (SMA) and its role in the cortical organization of movement come to be examined in detail by neuroscientists. Evidence from a wide variety of investigational perspectives is reviewed in an attempt to synthesize a conceptual framework for understanding SMA function. It is suggested that the SMA has an important role to play in the intentional process whereby internal context influences the elaboration of action. It may be viewed as phylogenetically older motor cortex, derived from anterior cingulate periarchicortical limbic cortex, which, as a key part of a medial premotor system, is crucial in the “programming” and fluent execution of extended action sequences which are “projectional” in that they rely on model-based prediction. This medial system can be distinguished from a lateral premotor system postulated to have evolved over phylogeny from a different neural source. An anatomico-physiologic model of the medial premotor system is proposed which embodies the principles of cyclicity and reentrance in the process of selecting those neural components to become active in conjunction with the performance of a particular action. The postulated dynamic action of this model in the microgenesis of a discrete action is outlined. It is concluded that although there is a great deal to be learned about the SMA, a convergence of current evidence can be identified. Such evidence suggests that the SMA plays an important role in the development of the intention-to-act and the specification and elaboration of action through its mediation between medial limbic cortex and primary motor cortex.

Hypoglossal motoneuron responses to pulmonary and superior laryngeal afferent inputs

In decerebrate, paralyzed cats ventilated with a cycle-triggered pump, the inspiratory discharges of the hypoglossal (whole nerve or single fibers), phrenic, and recurrent laryngeal nerves were compared, and the effects of pulmonary and superior laryngeal inputs were observed. During lung inflations phase with neural inspiration, hypoglossal and recurrent laryngeal activities differed from phrenic with respect to (a) burst onset times: both preceded the phrenic; (b) overall pattern: phrenic, augmenting; hypoglossal, decrementing; recurrent laryngeal, plateau-like. When inflation was withheld, the phrenic pattern was not markedly changed, but both hypoglossal and recurrent laryngeal became augmenting; the marked increas of hypoglossal activity (both whole nerve and single fiber) indicated strong inhibition by lung afferents. Superior laryngeal electrical stimulation evoked excitation of the contralateral phrenic (latency 4.1 msec) and the ipsilateral whole hypoglossal (latency 5.3 msec), followed by bilateral inhibitions (durations 20–30 msec); most hypoglossal fibers showed only inhibition. We conclude that, although both hypoglossal and phrenic outputs are driven by the inspiratory pattern generator(s), their premotor systems differ with respect to influences from central and peripheral inputs.

The organization of flexion-evoking interneurons in the abdominal nerve cord of the crayfish, Procambarus clarkii.

Intracellular recording and dye injection was used to investigate the flexion-evoking interneurons in the isolated abdominal nerve cord of the crayfish, Procambarus clarkii. The interneurons described in this study have all of the characteristics of cells called flexion command fibers in earlier works. These interneurons show cell bodies in all of the abdominal ganglia, all are interganglionic, and those with posteriorly directed axons enter the terminal caudal ganglion. Some cells were found in as many as eight different preparations, supporting the idea that these interneurons are identified cells. In addition to presenting evidence for identities, evidence was obtained for serial homology in this premotor system. A summary of the general organization of the flexion premotor apparatus is offered. It consists primarily of parallel elements (command interneurons) that are either bipolar or monopolar whose axonal projections course to the terminal caudal ganglion or rostrally into the thoracic region, perhaps into the cerebral ganglion. When activated singly, these parallel elements do not recruit additional interneurons but there are data to suggest that several of these elements could act in concert and interact in a low-gain fashion to produce coordinated positioning behavior.

Exteroceptive and proprioceptive afferents of the trigeminal and facial motor nuclei in the mallard (Anas platyrhynchos L.).

Central pathways converging upon the trigeminofacial motor nuclei of the mallard were studied in order to elucidate neuroanatomically the presumed influence of primary sensory trigeminal afferents upon jaw muscle activity. The techniques used included the Fink-Heimer I method after lesions, and axonal transport labeling following injections of 3H-leucine or of HRP for retrograde identification of the neurons of origin. A general description is given of the trigeminofacial motor complex. Jaw closer muscles are innervated by trigeminal motor neurons, and facial motor neurons innervate the jaw depressor muscles. Two afferents premotor systems, one including the mesencephalic trigeminal nucleus (MesV) and the other the rhombencephalic reticular formation, are distinguished. The proprioceptive neurons of the mesencephalic trigeminal nucleus project upon the ipsilateral trigeminal motor nucleus and upon the nucleus supratrigeminalis. The latter cell group bilaterally projects upon the dorsal and intermediate parts of the facial motor nucleus and upon the dorsal and intermediate parts of the facial motor nucleus and upon part of the trigeminal motor nucleus. Exteroceptive information, relayed through the primary sensory trigeminal column (PrV and nTTD), ultimately reaches the motor nuclei via the reticular formation. The reticular formation forms the final link of three separate circuits: a telencephalic one entered through the principal trigeminal sensory nucleus, a cerebellar one via subnucleus oralis of the descending trigeminal system, and a direct one via subnucleus interpolaris. No direct connections between the principal trigeminal sensory nucleus or subnuclei of the descending trigeminal system and the motor nuclei of the trigeminal (NV) and facial (NVII) nerves have been observed, nor are such direct projections present in the outflow of the presumed telencephalic and cerebellar circuits, viz. of the archistriatum and the central cerebellar nuclei, respectively. The archistriatum projects via the occipitomesencephalic tract upon the lateral rhombencephalic reticular formation as far down as the rostral cervical cord, as well as upon the subnucleus interpolaris of the descending trigeminal system. Similarly, efferents from the central cerebellar nuclei reach the reticular formation, which in turn projects bilaterally upon the motor nuclei. Finally, commissural intermotor connections apparently are mediated by reticular cells surrounding the motor nuclei of NV or NVII, rather than emanating from these nuclei directly.

Feeding inHelisoma trivolvis: The Morphological and Physiological Bases of a Fixed Action Pattern

This report is a description of feeding in the pulmonate snail, Helisoma trivolvis and provides a detailed account of: (i) the behavior, (ii) the muscular system, (iii) the muscle activity patterns, (iv) the neuromusculai relationships, (v) the motoneurons to specific muscles, and finally, (vi) an introduction to the premotor system. Feeding is the result of the integration of sensory information onto components of a central program which is delived from connections within the buccal ganglia. This report emphasizes the analysis of the centrally programmed components of the feeding output by characterizing interactions among three classes of neurons (approximately 50 cells) in the buccal ganglia: protractor motoneurons, retractor motoneurons, and the neurons of an electrically coupled neural network which produces both the timing and driving of motoneuron activity.