Postnatal development of the motor representation in primary motor cortex.

The purpose of this study was to examine when the muscles and joints of the forelimb become represented in primary motor cortex (M1) during postnatal life and how local representation patterns change. We examined these questions in cats that were anesthetized (45-90 days, n = 14; adults, n = 3) and awake (n = 4; 52-86 days). We used intracortical microstimulation (45 ms duration train, 330 Hz, 0.2-ms balanced biphasic pulses, with a leading cathodic pulse; up to 100 microA). In young animals (less than day 70), we also used stimulus trains and pulses that could produce greater temporal summation (up to 200-ms train duration, down to 143-Hz stimulus frequency, up to 0.8-ms pulse width). Anesthetized animals were areflexic, and muscle tone was similar to that of the awake cats (i.e., relaxed, not weight or load bearing, with minimal resistance to passive stretch). We monitored the kinematic effects of microstimulation and changes in electromyographic (EMG) activity in forelimb muscles. There was an age-dependent reduction in the number of sites where microstimulation did not produce a motor effect (i.e., ineffective sites), from 95% in animals younger than 60 days to 33% between 81 and 90 days. In adults, 24% of sites were ineffective. Median current thresholds for evoking movements dropped from 79 microA in animals younger than day 60 to 38 and 28 microA in day 81-90 animals and adults, respectively. There was a proximal-to-distal development of the somatotopic organization of the motor map. Stimulation at the majority of sites in animals younger than day 71 produced shoulder and elbow movement. Wrist sites were first present by day 71, and digit sites by day 81. Sites at which multiple responses were evoked, between 1.0 and 1.5 times threshold, were present after day 71, and increased with age. A higher percentage of distal joints were co-represented with other joints, rather than being represented alone. We found that effective sites initially were scattered and new sites representing proximal and distal joints filled in the gaps between effective sites. During most of the period examined, development of the caudal M1 subregion lagged that of the rostral subregion (percent of effective sites; threshold currents), although these differences were minimal or absent in adults. Our results show that the M1 motor representation is absent at day 45 and, during the subsequent month, the motor map is constructed by progressively representing more distal forelimb joints.