Exteroceptive, proprioceptive, and sympathetic activity recorded with microelectrodes from human peripheral nerves.

With tungsten microelectrodes percutaneously inserted into human peripheral nerve fascicles, in vivo recordings of single-unit action potentials can be obtained from different types of myelinated and unmyelinated nerve fibers. This technique of microneurography has provided a great deal of information about (1) the sensory innervation of the human skin, with its mechanoreceptive, thermoreceptive, and nociceptive functions, (2) the proprioceptive innervation of human skeletal muscles, special attention being devoted to the motor control functions of the muscle spindles and the fusimotor system, and (3) the sympathetic innervation of autonomic effector organs in skin and muscle, with its significance for, for example, regulation of temperature and blood pressure. The technique has its main application not as a routine procedure in clinical diagnostic work but as a research tool in basic physiologic and pathophysiologic studies.

Thixotropy of rib cage respiratory muscles in normal subjects.

In this study, we searched for signs of thixotropic behavior in human rib cage respiratory muscles. If rib cage respiratory muscles possess thixotropic properties similar to those seen in other skeletal muscles in animals and humans, we expect resting rib cage circumference would be temporarily changed after deep rib cage inflations or deflations and that these aftereffects would be particularly pronounced in trials that combine conditioning deep inflations or deflations with forceful isometric contractions of the respiratory muscles. We used induction plethysmography to obtain a continuous relative measure of rib cage circumference changes during quiet breathing in 12 healthy subjects. Rib cage position at the end of the expiratory phase (EEP) was used as an index of resting rib cage circumference. Comparisons were made between EEP values of five spontaneous breaths immediately before and after six types of conditioning maneuvers: deep inspiration (DI); deep expiration (DE); DI combined with forceful effort to inspire (FII) or expire (FEI); and DE combined with forceful effort to inspire (FIE) or expire (FEE), both with temporary airway occlusion. The aftereffects of the conditioning maneuvers on EEP values were consistent with the supposition that human respiratory muscles possess thixotropic properties. EEP values were significantly enhanced after all conditioning maneuvers involving DI, and the aftereffects were particularly pronounced in the FII and FEI trials. In contrast, EEP values were reduced after DE maneuvers. The aftereffects were statistically significant for the FEE and FIE, but not DE, trials. It is suggested that respiratory muscle thixotropy may contribute to the pulmonary hyperinflation seen in patients with chronic obstructive pulmonary disease.

Conductivity ratios of the scalp-skull-brain head model in estimating equivalent dipole sources in human brain.

The dipole tracing (DT) method estimates the position and vector dipole moment of an equivalent current dipole by minimizing the mean squared error of the dipole potentials at the surface electrode positions. In the scalp-skull-brain/DT (SSB/DT) method, which we have developed, the head model consists of three compartments of uniform conductors corresponding to the scalp, skull and brain. The accuracy of the calculations are mainly dependent on the ratios of the conductivities of the three compartments. The best result was obtained with the conductivity ratios of 1:1/80:1 for the scalp, skull and brain compartments, respectively.

Microelectrode recordings from transected nerves in amputees with phantom limb pain.

Intraneural microelectrode recordings were made from the nerve supplying the phantom area in two patients suffering from phantom limb pain. Spontaneous activity was prominent in both cutaneous and muscle fascicle of the nerves. Tapping the neuromata which accentuated the phantom limb pain, induced afferent discharges with both short and long latencies, the latter from fibres with a conduction velocity of only 0.5 m/sec. Blocking the neuromata with lidocaine completely abolished the tap-induced afferent discharges and the tap-induced accentuation of the phantom pain. The spontaneous pain was, however, unchanged, as was the spontaneous activity recorded.

Segmentation of human spindle and EMG responses to sudden muscle stretch.

Grouping of the EMG response produced by quick stretches of contracting muscles has been thought to reflect 'long loop reflexes' through cerebral cortex adding to the segmental stretch reflex. Our recordings from human muscle spindle afferents responding to such stretches show that these discharges also tend to be grouped. EMG grouping may therefore be a consequence of successive segmental reflexes rather than of additional delays in long loop reflex arcs.

Generator mechanisms of epileptic potentials analyzed by dipole tracing method.

A new dipole tracing method, based on a realistic head model, was used to determine dipole locations and vector moments of interictal convexity sharp waves recorded (with conventional EEG technique) from the right fronto-temporal region in a patient with partial complex seizures. When the dipole locations in the head model were compared to MRI scans, the majority of the sharp wave dipoles were found to be located in the right hippocampal area. For individual sharp waves, the hippocampal dipoles moved along tracks corresponding to the vector moment directions, suggesting that the electrical sources of the convexity sharp waves were somato-dendritic currents which spread rapidly from one neuron group to the next in the hippocampal area. Previous long-term subdural recording had shown seizure onset in this area. After right-sided anterior temporal lobectomy including the hippocampus the patient has been seizure-free for three months.

Double blind cross over trial with deprenyl in amyotrophic lateral sclerosis.

In this paper we present results from a double blind cross over trial with deprenyl, a selective and irreversible monoamine oxidase-B (MAO-B) inhibitor, in 10 patients suffering from amyotrophic lateral sclerosis. The patients were randomised in such a way that half of the patients started with the active drug and half with the placebo treatment. Each patient was given 10 mg deprenyl (eldepryl, 10 mg tablets) per day for 12 weeks and then placebo for the same length of time. There was a drug free period of 12 weeks between the courses. The neurological status of the patients were evaluated every six weeks by using Norris, spinal and bulbar scores and it was observed that all cases deteriorated in their clinical status during the 36 weeks of the controlled study. MAO-B activity in blood platelets was completely inhibited during treatment with deprenyl. In the preliminary analysis performed so far, no obvious retardation in the progress of the disease could be observed with deprenyl treatment.

The fusimotor system. Its role in fatigue.

Several lines of evidence point to an important role of the fusimotor system in the "muscle-wisdom" phenomenon during peripheral fatigue of some human voluntary contractions: 1) muscle afferents provide a net amplification of skeletomotor output, with the only known afferent species capable of this being the muscle spindle; 2) muscle spindle firing rates decline during constant-force voluntary contractions, so fusimotor support to skeletomotor output decreases; 3) this waning support can be offset by application of high-frequency vibration to the fatiguing muscle, which excites spindle endings; and 4) the progressive decline in motor unit firing rates during maximal voluntary contractions is abolished by blocking muscle afferent inputs, and it is argued that, at least in the initial stages of a contraction, this must be due to a progressive withdrawal of spindle support.

Microneurography and applications to issues of motor control: Fifth Annual Stuart Reiner Memorial Lecture.

Among the hypotheses regarding fusimotor functions based on earlier animal experiments some are inconsistent, others are in conformity with microneurographic observations in man. The human data provide evidence against the following two theories: (1) the length follow-up servo theory; and (2) the theory that fusimotor neurons can be selectively activated to produce spindle sensitization and stretch reflex reinforcements. The human data support the theory of alpha-gamma coactivation. In particular, in the early phase of isometric voluntary contractions fusimotor-driven afferent spindle activity assists in autogenetic activation of alpha motoneurons and in reciprocal relaxation of antagonists. As muscle fatigue develops, the autogenetic reflex drive via the fusimotor route declines. The fusimotor bias during contraction provides for maintenance of spindle sensitivity to minute perturbations and for load-compensating reflex adjustments to such perturbations. Reflex overcorrections may lead to uncontrollable oscillations of the type seen in enhanced physiological tremor.

Human motor compensations for thixotropy-dependent changes in resting wrist joint position after large joint movements.

AIM: Resting tension of relaxed skeletal muscle fibres held at a given length varies with the immediate previous history of length changes and contractions. The primary aim of this study was to explore the motor control consequences of this history-dependency in healthy subjects. METHODS: Angular position and passive torque were recorded from the intact wrist joint. Integrated surface electromyography (IEMG) was recorded from wrist extensor and flexor muscles. RESULTS: In relaxed subjects, wrist joint position was displaced towards dorsiflexion after a single high-amplitude dorsiflexion movement combined with a strong flexor/extensor co-contraction (dorsiflexion conditioning), whereas after volarflexion conditioning there was a shift towards volarflexion. These after-effects could be abruptly cancelled by short periods ( approximately 5 s) of rapid flapping hand movements or forceful isometric co-contractions, findings indicative of muscle thixotropy. The IEMG-evaluated motor after-effects were as follows. A slowly subsiding wrist flexor contraction was needed to restore and maintain the original resting wrist position after dorsiflexion conditioning whereas a slowly subsiding extensor contraction was needed for the same goal after volarflexion conditioning. Furthermore, ongoing wrist extensor IEMG activity required to actively hold the wrist in a moderate dorsiflexed position or to resist a constant volar torque at resting position was temporarily reduced after dorsiflexion conditioning and enhanced (not significantly) after volarflexion conditioning. CONCLUSION: The results provide evidence that during voluntary maintenance of a desired wrist joint position the motor commands to the position-holding muscles are unconsciously adjusted to compensate for thixotropy-dependent variations in the resting tension of the muscles.

Participation of the stretch reflex in human physiological tremor.

Microneurographic studies were made of normal human muscle spindle afferent activity to evaluate the role played by these endings, and by the segmental stretch reflex, in various types of physiological tremor. Primary endings are sufficiently sensitive to respond to the minute tremulous movements normally seen in contracting muscles. Human muscle spindle endings appear to possess the same non-linear sensitivity to small stretches as has been reported for cat muscle spindles. The findings also support the notion that the segmental stretch reflex plays an important role in enhanced physiological tremor, in so far as it tends to synchronize the motor outflow in such a way as to produce rhythmical contractions properly timed to reinforce the mechanical resonant properties of muscle spindle endings, the stretch reflex also serves to damp the larger oscillations of finger or hand which result from a brisk tap or muscle twitch. There is no evidence for alpha-gamma co-activation in these enhanced physiological tremors.

Physiological tremor enhanced by manoeuvres affecting the segmental stretch reflex.

In view of recent evidence that physiological tremor can be enhanced by positive feedback via the segmental stretch reflex, several manoeuvres and procedures were employed to enhance the finger and hand tremor of healthy subjects--the purpose being to determine if tremorogenic effects, at least in part, are due to increase efficacy of the stretch reflex servo. Mechanical events during tremor (and during voluntary or electrically induced muscle twitches) were recorded together with EMG activity from wrist and finger flexor muscles and discharges from primary spindle endings in these muscles. Physiological tremor can be enhanced not only by manoeuvres which increase the gain of segmental stretch reflexes (Jendrassik manoeuvre) but also by manoeuvres which increase the contrast in spindle firing during stretch versus shortening phases of tremor, thus enhancing reflex modulation. Effects of the latter type can be achieved by procedures which alter mechanical twitch properties of extrafusal fibres (isoproterenol infusions and fatigue) and by procedures which involve application of spindle stimuli acting preferentially during stretch phases of tremor movements (muscle vibrations). Physiological tremor, which can be temporarily enhanced by an externally applied muscle perturbation, also becomes accentuated by those small "pseudo-myoclonic" jerks which occur in all normal subjects attempting to perform slow, smooth movements.

Human motor control consequences of thixotropic changes in muscular short-range stiffness.

1. The primary aim of the present study was to explore whether in healthy subjects the muscle contractions required for unrestrained voluntary wrist dorsiflexions are adjusted in strength to thixotropy-dependent variations in the short-range stiffness encountered in measurements of passive torque resistance to imposed wrist dorsiflexions. 2. After a period of rest, only the first movement in a series of passive wrist dorsiflexions of moderate amplitude exhibited clear signs of short-range stiffness in the torque response. During analogous types of voluntary movements, the extensor EMG during the first movement after rest showed a steep initial rise of activity, which apparently served to compensate for the short-range stiffness. 3. The passive torque resistance to minute repetitive wrist dorsiflexions (within the range of short-range stiffness) was markedly reduced after various types of mechanical agitation. During analogous low-amplitude voluntary wrist dorsiflexions the extensor EMG signals were weaker after than before agitation. 4. Mechanical agitation also led to enhancement of passive dorsiflexion movements induced by weak constant torque pulses. In an analogous way, the movement-generating capacity of weak voluntary extensor activations (as determined by EMG recordings) was greatly enhanced by mechanical agitation. 5. The signals from a force transducer probe pressed against the wrist flexor tendons--during passive wrist dorsiflexions--revealed short-range stiffness responses which highly resembled those observed in the torque measurements, suggesting that the latter to a large extent emanated from the stretched, relaxed flexor muscles. During repetitive stereotyped voluntary wrist dorsiflexions, a close correspondence was observed between the degree of short-range stiffness as sensed by the wrist flexor tension transducer and the strength of the initial extensor activation required for movement generation. 6. The results provide evidence that the central nervous system in its control of voluntary movements takes account of and compensates for the history-dependent degree of inherent short-range stiffness of the muscles antagonistic to the prime movers.