Pruszynski JA, Kurtzer I, Lillicrap TP, Scott SH. Temporal evolution of "automatic gain-scaling." J Neurophysiol 102: 992-1003, 2009. First published May 3, 2009 doi:10.1152/jn.00085.2009. The earliest neural response to a mechanical perturbation, the short-latency stretch response (R1: 20 -45 ms), is known to exhibit "automatic gain-scaling" whereby its magnitude is proportional to preperturbation muscle activity. Because gain-scaling likely reflects an intrinsic property of the motoneuron pool (via the size-recruitment principle), counteracting this property poses a fundamental challenge for the nervous system, which must ultimately counter the absolute change in load regardless of the initial muscle activity (i.e., show no gainscaling). Here we explore the temporal evolution of gain-scaling in a simple behavioral task where subjects stabilize their arm against different background loads and randomly occurring torque perturbations. We quantified gain-scaling in four elbow muscles (brachioradialis, biceps long, triceps lateral, triceps long) over the entire sequence of muscle activity following perturbation onset-the shortlatency response, long-latency response (R2: 50 -75 ms; R3: 75-105 ms), early voluntary corrections (120 -180 ms), and steady-state activity (750 -1250 ms). In agreement with previous observations, we found that the short-latency response demonstrated substantial gainscaling with a threefold increase in background load resulting in an approximately twofold increase in muscle activity for the same perturbation. Following the short-latency response, we found a rapid decrease in gain-scaling starting in the long-latency epoch (ϳ75-ms postperturbation) such that no significant gain-scaling was observed for the early voluntary corrections or steady-state activity. The rapid decrease in gain-scaling supports our recent suggestion that longlatency responses and voluntary control are inherently linked as part of an evolving sensorimotor control process through similar neural circuitry.
I N T R O D U C T I O NA prominent and well-studied feature of the short-latency stretch response (R1: 20 -45 ms postperturbation) is "automatic gain-scaling" whereby the same muscle-stretch will elicit larger responses when preperturbation muscle activity is increased (Bedingham and Tatton 1984;Marsden et al. 1976;Matthews 1986;Stein et al. 1995;Verrier 1985). This modulation is generally attributed to the intrinsic organization of the motoneuron pool (Capaday and Stein 1987;Houk et al. 1970;Kernell and Hultborn 1990;Marsden et al. 1976;Matthews 1986;Slot and Sinkjaer 1994) where motor units are recruited in order of their force-generating capability and resilience to fatigue (Cope and Clark 1991; Henneman 1957), a phenomenon termed the size-recruitment principle. Although gainscaling may be a useful short-term strategy (Bedingham and Tatton 1984;Marsden et al. 1976;Matthews 1986), ultimately the steady-state response to an additional load must be independent of preperturbation muscle activity (i.e., not show gain-sc...