对记忆目标的扫视:平稳跟踪和虚幻刺激运动的影响。
Saccades to remembered targets: the effects of smooth pursuit and illusory stimulus motion.
作者信息
Zivotofsky A Z, Rottach K G, Averbuch-Heller L, Kori A A, Thomas C W, Dell'Osso L F, Leigh R J
机构信息
Department of Neurology, Case Western Reserve University, Cleveland, Ohio 44106, USA.
出版信息
J Neurophysiol. 1996 Dec;76(6):3617-32. doi: 10.1152/jn.1996.76.6.3617.
- Measurements were made in four normal human subjects of the accuracy of saccades to remembered locations of targets that were flashed on a 20 x 30 deg random dot display that was either stationary or moving horizontally and sinusoidally at +/-9 deg at 0.3 Hz. During the interval between the target flash and the memory-guided saccade, the "memory period" (1.4 s), subjects either fixated a stationary spot or pursued a spot moving vertically sinusoidally at +/-9 deg at 0.3 Hz. 2. When saccades were made toward the location of targets previously flashed on a stationary background as subjects fixated the stationary spot, median saccadic error was 0.93 deg horizontally and 1.1 deg vertically. These errors were greater than for saccades to visible targets, which had median values of 0.59 deg horizontally and 0.60 deg vertically. 3. When targets were flashed as subjects smoothly pursued a spot that moved vertically across the stationary background, median saccadic error was 1.1 deg horizontally and 1.2 deg vertically, thus being of similar accuracy to when targets were flashed during fixation. In addition, the vertical component of the memory-guided saccade was much more closely correlated with the "spatial error" than with the "retinal error"; this indicated that, when programming the saccade, the brain had taken into account eye movements that occurred during the memory period. 4. When saccades were made to targets flashed during attempted fixation of a stationary spot on a horizontally moving background, a condition that produces a weak Duncker-type illusion of horizontal movement of the primary target, median saccadic error increased horizontally to 3.2 deg but was 1.1 deg vertically. 5. When targets were flashed as subjects smoothly pursued a spot that moved vertically on the horizontally moving background, a condition that induces a strong illusion of diagonal target motion, median saccadic error was 4.0 deg horizontally and 1.5 deg vertically; thus the horizontal error was greater than under any other experimental condition. 6. In most trials, the initial saccade to the remembered target was followed by additional saccades while the subject was still in darkness. These secondary saccades, which were executed in the absence of visual feedback, brought the eye closer to the target location. During paradigms involving horizontal background movement, these corrections were more prominent horizontally than vertically. 7. Further measurements were made in two subjects to determine whether inaccuracy of memory-guided saccades, in the horizontal plane, was due to mislocalization at the time that the target flashed, misrepresentation of the trajectory of the pursuit eye movement during the memory period, or both. 8. The magnitude of the saccadic error, both with and without corrections made in darkness, was mislocalized by approximately 30% of the displacement of the background at the time that the target flashed. The magnitude of the saccadic error also was influenced by net movement of the background during the memory period, corresponding to approximately 25% of net background movement for the initial saccade and approximately 13% for the final eye position achieved in darkness. 9. We formulated simple linear models to test specific hypotheses about which combinations of signals best describe the observed saccadic amplitudes. We tested the possibilities that the brain made an accurate memory of target location and a reliable representation of the eye movement during the memory period, or that one or both of these was corrupted by the illusory visual stimulus. Our data were best accounted for by a model in which both the working memory of target location and the internal representation of the horizontal eye movements were corrupted by the illusory visual stimulus. We conclude that extraretinal signals played only a minor role, in comparison with visual estimates of the direction of gaze, in planning eye movements to remembered targ
- 对4名正常人类受试者进行了测量,测量他们对记忆中目标位置进行扫视的准确性。目标在一个20×30度的随机点显示器上闪烁,该显示器要么静止,要么以0.3赫兹的频率水平正弦移动,幅度为±9度。在目标闪烁与记忆引导扫视之间的间隔期,即“记忆期”(1.4秒)内,受试者要么注视一个固定点,要么追踪一个以0.3赫兹的频率垂直正弦移动、幅度为±9度的点。2. 当受试者注视固定点时,向先前在静止背景上闪烁的目标位置进行扫视,水平方向的中位数扫视误差为0.93度,垂直方向为1.1度。这些误差大于对可见目标的扫视误差,可见目标扫视误差的中位数水平为0.59度,垂直为0.60度。3. 当受试者平稳追踪在静止背景上垂直移动的点时目标闪烁,水平方向的中位数扫视误差为1.1度,垂直方向为1.2度,因此与注视期间目标闪烁时的准确性相似。此外,记忆引导扫视的垂直分量与“空间误差”的相关性比与“视网膜误差”的相关性高得多;这表明,在规划扫视时,大脑已经考虑到了记忆期内发生的眼球运动。4. 当在水平移动背景上试图注视固定点时目标闪烁并进行扫视,这种情况会产生较弱的主要目标水平移动的邓克类型错觉,水平方向的中位数扫视误差增加到3.2度,但垂直方向为1.1度。5. 当受试者平稳追踪在水平移动背景上垂直移动的点时目标闪烁,这种情况会产生强烈的目标对角运动错觉,水平方向的中位数扫视误差为4.0度,垂直方向为1.5度;因此水平误差大于任何其他实验条件下的误差。6. 在大多数试验中,对记忆目标的初始扫视之后,受试者仍处于黑暗中时会有额外的扫视。这些在没有视觉反馈的情况下执行的二次扫视使眼睛更接近目标位置。在涉及水平背景移动的范式中,这些校正水平方向比垂直方向更明显。7. 对两名受试者进行了进一步测量,以确定在水平面内记忆引导扫视的不准确是由于目标闪烁时的定位错误、记忆期内追踪眼球运动轨迹的错误表征,还是两者兼而有之。8. 无论在黑暗中是否进行校正,扫视误差的大小因目标闪烁时背景位移的约30%而发生定位错误。扫视误差的大小也受到记忆期内背景净移动的影响,对于初始扫视,约相当于背景净移动的25%,对于在黑暗中最终达到的眼位,则约为13%。9. 我们制定了简单的线性模型来测试关于哪些信号组合最能描述观察到的扫视幅度的特定假设。我们测试了大脑对目标位置进行准确记忆并在记忆期内对眼球运动进行可靠表征的可能性,或者这些中的一个或两个是否被虚幻的视觉刺激破坏。我们的数据最能由一个模型解释,在该模型中,目标位置的工作记忆和水平眼球运动的内部表征都被虚幻的视觉刺激破坏。我们得出结论,与注视方向的视觉估计相比,视网膜外信号在规划对记忆目标的眼球运动中仅起次要作用。