School of Engineering, University of Glasgow, Glasgow, UK.
Systems Biology Laboratory, Melbourne School of Engineering, University of Melbourne, Melbourne, Australia.
J Physiol. 2017 Nov 1;595(21):6751-6770. doi: 10.1113/JP274288. Epub 2017 Oct 1.
A human controlling an external system is described most easily and conventionally as linearly and continuously translating sensory input to motor output, with the inevitable output remnant, non-linearly related to the input, attributed to sensorimotor noise. Recent experiments show sustained manual tracking involves repeated refractoriness (insensitivity to sensory information for a certain duration), with the temporary 200-500 ms periods of irresponsiveness to sensory input making the control process intrinsically non-linear. This evidence calls for re-examination of the extent to which random sensorimotor noise is required to explain the non-linear remnant. This investigation of manual tracking shows how the full motor output (linear component and remnant) can be explained mechanistically by aperiodic sampling triggered by prediction error thresholds. Whereas broadband physiological noise is general to all processes, aperiodic sampling is associated with sensorimotor decision making within specific frontal, striatal and parietal networks; we conclude that manual tracking utilises such slow serial decision making pathways up to several times per second.
The human operator is described adequately by linear translation of sensory input to motor output. Motor output also always includes a non-linear remnant resulting from random sensorimotor noise from multiple sources, and non-linear input transformations, for example thresholds or refractory periods. Recent evidence showed that manual tracking incurs substantial, serial, refractoriness (insensitivity to sensory information of 350 and 550 ms for 1st and 2nd order systems respectively). Our two questions are: (i) What are the comparative merits of explaining the non-linear remnant using noise or non-linear transformations? (ii) Can non-linear transformations represent serial motor decision making within the sensorimotor feedback loop intrinsic to tracking? Twelve participants (instructed to act in three prescribed ways) manually controlled two systems (1st and 2nd order) subject to a periodic multi-sine disturbance. Joystick power was analysed using three models, continuous-linear-control (CC), continuous-linear-control with calculated noise spectrum (CCN), and intermittent control with aperiodic sampling triggered by prediction error thresholds (IC). Unlike the linear mechanism, the intermittent control mechanism explained the majority of total power (linear and remnant) (77-87% vs. 8-48%, IC vs. CC). Between conditions, IC used thresholds and distributions of open loop intervals consistent with, respectively, instructions and previous measured, model independent values; whereas CCN required changes in noise spectrum deviating from broadband, signal dependent noise. We conclude that manual tracking uses open loop predictive control with aperiodic sampling. Because aperiodic sampling is inherent to serial decision making within previously identified, specific frontal, striatal and parietal networks we suggest that these structures are intimately involved in visuo-manual tracking.
描述人类对外部系统的控制,最容易和最常用的方法是线性且连续地将感觉输入转换为运动输出,而不可避免的输出残余是非线性的,与输入无关,归因于感觉运动噪声。最近的实验表明,持续的手动跟踪涉及反复的不敏感(在一定时间内对感觉信息不敏感),暂时的 200-500 毫秒对感觉输入无响应的时间段使控制过程本质上是非线性的。这一证据要求重新检验随机感觉运动噪声在多大程度上需要解释非线性残余。这项对手动跟踪的研究表明,周期性采样触发的预测误差阈值如何能够从机制上解释完整的运动输出(线性成分和残余)。虽然宽带生理噪声是所有过程的普遍现象,但周期性采样与特定额、纹状体和顶叶网络中的感觉运动决策有关;我们的结论是,手动跟踪每秒会利用几次这样的缓慢串行决策路径。
人类操作员可以通过线性翻译感觉输入到运动输出来充分描述。运动输出也总是包括由于来自多个来源的随机感觉运动噪声以及非线性输入转换(例如阈值或不应期)而产生的非线性残余。最近的证据表明,手动跟踪会产生大量的、串行的不应期(对一阶和二阶系统的感觉信息分别为 350 和 550 毫秒不敏感)。我们的两个问题是:(i)使用噪声或非线性转换来解释非线性残余的相对优点是什么?(ii)非线性转换是否可以代表跟踪内在的感觉运动反馈回路中的串行运动决策?12 名参与者(被指示以三种规定的方式行事)手动控制两个受周期性多正弦干扰的系统(一阶和二阶)。使用三种模型(连续线性控制(CC)、带有计算噪声谱的连续线性控制(CCN)和由预测误差阈值触发的间歇控制(IC))分析操纵杆功率。与线性机制不同,间歇控制机制解释了大部分总功率(线性和残余)(77-87% 与 8-48%,IC 与 CC)。在不同条件下,IC 使用与指令和以前测量的、模型独立的值分别一致的阈值和开环间隔分布;而 CCN 需要偏离宽带、信号相关噪声的噪声谱变化。我们的结论是,手动跟踪使用带有周期性采样的开环预测控制。由于间歇采样是先前确定的特定额、纹状体和顶叶网络内串行决策的固有组成部分,我们认为这些结构与视觉手动跟踪密切相关。