Humphrey A L, Saul A B
Department of Neurobiology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania 15261, USA.
J Neurophysiol. 1998 Dec;80(6):2991-3004. doi: 10.1152/jn.1998.80.6.2991.
Strobe rearing reduces direction selectivity in area 17 by altering spatiotemporal receptive-field structure. J. Neurophysiol. 80: 2991-3004, 1998. Direction selectivity in simple cells of cat area 17 is linked to spatiotemporal (S-T) receptive-field structure. S-T inseparable receptive fields display gradients of response timing across the receptive field that confer a preferred direction of motion. Receptive fields that are not direction selective lack gradients; they are S-T separable, displaying uniform timing across the field. Here we further examine this link using a developmental paradigm that disrupts direction selectivity. Cats were reared from birth to 8 mo of age in 8-Hz stroboscopic illumination. Direction selectivity in simple cells was then measured using gratings drifting at different temporal frequencies (0.25-16 Hz). S-T structure was assessed using stationary bars presented at different receptive-field positions, with bar luminance being modulated sinusoidally at different temporal frequencies. For each cell, plots of response phase versus bar position were fit by lines to characterize S-T inseparability at each temporal frequency. Strobe rearing produced a profound loss of direction selectivity at all temporal frequencies; only 10% of cells were selective compared with 80% in normal cats. The few remaining directional cells were selective over a narrower than normal range of temporal frequencies and exhibited weaker than normal direction selectivity. Importantly, the directional loss was accompanied by a virtual elimination of S-T inseparability. Nearly all cells were S-T separable, like nondirectional cells in normal cats. The loss was clearest in layer 4. Normally, inseparability is greatest there, and it correlates well (r = 0.77) with direction selectivity; strobe rearing reduced inseparability and direction selectivity to very low values. The few remaining directional cells were inseparable. In layer 6 of normal cats, most direction-selective cells are only weakly inseparable, and there is no consistent relationship between the two measures. However, after strobe rearing, even the weak inseparability was eliminated along with direction selectivity. The correlated changes in S-T structure and direction selectivity were confirmed using conventional linear predictions of directional tuning based on responses to counterphasing bars and white noise stimuli. The developmental changes were permanent, being observed up to 12 yr after strobe rearing. The deficits were remarkably specific; strobe rearing did not affect spatial receptive-field structure, orientation selectivity, spatial or temporal frequency tuning, or general responsiveness to visual stimuli. These results provide further support for a critical role of S-T structure in determining direction selectivity in simple cells. Strobe rearing eliminates directional tuning by altering the timing of responses within the receptive field.
频闪饲养通过改变时空感受野结构降低了17区的方向选择性。《神经生理学杂志》80: 2991 - 3004, 1998年。猫17区简单细胞的方向选择性与时空(S-T)感受野结构有关。S-T不可分离的感受野在感受野上显示出反应时间的梯度,赋予了一个偏好的运动方向。非方向选择性的感受野缺乏梯度;它们是S-T可分离的,在整个视野中显示出均匀的时间。在这里,我们使用一种破坏方向选择性的发育范式进一步研究这种联系。猫从出生到8个月大在8赫兹频闪照明下饲养。然后使用以不同时间频率(0.25 - 16赫兹)漂移的光栅测量简单细胞的方向选择性。使用在不同感受野位置呈现的静止条来评估S-T结构,条的亮度以不同时间频率进行正弦调制。对于每个细胞,将反应相位与条位置的图拟合为直线,以表征每个时间频率下的S-T不可分离性。频闪饲养在所有时间频率下都导致方向选择性的严重丧失;只有10%的细胞具有选择性,而正常猫中这一比例为80%。少数剩余的方向选择性细胞在比正常更窄的时间频率范围内具有选择性,并且表现出比正常更弱的方向选择性。重要的是,方向选择性的丧失伴随着S-T不可分离性的几乎完全消除。几乎所有细胞都是S-T可分离的,就像正常猫中的非方向选择性细胞一样。这种丧失在第4层最为明显。正常情况下,那里的不可分离性最大,并且与方向选择性密切相关(r = 0.77);频闪饲养将不可分离性和方向选择性降低到非常低的值。少数剩余的方向选择性细胞是不可分离的。在正常猫的第6层,大多数方向选择性细胞只是弱不可分离的,并且这两种测量之间没有一致的关系。然而,在频闪饲养后,即使是弱不可分离性也与方向选择性一起被消除了。使用基于对反相条和白噪声刺激的反应的方向调谐的传统线性预测,证实了S-T结构和方向选择性相关的变化。这些发育变化是永久性的,在频闪饲养后长达12年都能观察到。这些缺陷非常具有特异性;频闪饲养不影响空间感受野结构、方向选择性、空间或时间频率调谐,或对视觉刺激的一般反应性。这些结果为S-T结构在确定简单细胞方向选择性中的关键作用提供了进一步的支持。频闪饲养通过改变感受野内的反应时间消除了方向调谐。