猫初级视皮层17区神经元的动态反应特性及21a区的反馈投射对17区信息处理的调制影响
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摘要
对哺乳类动物视觉皮层如何编码视觉刺激的消失的研究并不多见。在这里,我们研究了猫视皮层17区的88个神经元从刺激开始以及刺激消失后的动态反应特性。在62.5%的神经元中(88个中的55个,31个简单细胞,24个复杂细胞),我们观察到由静止光栅诱发的可信的撤刺激瞬时反应(offset transient response),它与给刺激瞬时反应(onset transient response)有相似的最优方位,方位敏感强度,累积检测能力,峰反应幅度和峰潜伏期。但是,这种撤刺激瞬时反应显著地依赖于刺激呈现的时间,意味着神经元回路需要更长的时间来共激活以产生撤刺激瞬时反应。简单细胞和复杂细胞的撤刺激瞬时反应的特性并不一样。复杂细胞的给刺激瞬时反应和撤刺激瞬时反应的峰潜伏期比简单细胞的有更强的线性关系,有更高的撤刺激瞬时反应的峰反应幅度,并且给刺激瞬时反应和撤刺激瞬时反应的产生都不依赖于光栅的空间相位。我们所做的线性消褪光栅的实验证明,引起撤刺激瞬时反应的是感受野亚区内局部的光强变化。
人类和其他一些哺乳类对水平-垂直的视觉刺激的敏感度要高于倾斜方位的刺激(这一现象被称为“倾斜效应”)。虽然产生这种效应的神经机制目前尚不很清楚,但是一种可能性是:参与处理水平-垂直方位信息的神经元数量要大于参与处理倾斜方位的。猫视皮层21a区接受17区输入并有反馈投射至17区。本实验室以往的研究发现猫视皮层17区和21a区内对水平-垂直方位敏感的皮层区域明显大于对倾斜方位敏感的皮层区域。而且在21a区的倾斜效应更为明显。本文利用内源信号光学成像技术结合定点微量注射药物的方法,探索高级视皮层21a区的反馈调制对初级视皮层17区的倾斜效应得影响。我们首次发现,增加21a区的活动水平可以使17区的水平-垂直与倾斜方位的敏感区的不均一分布的差值水平从4.9%上升到18.8%,已经接近21a区的23.2%的水平,而降低21a区的活动水平则可以减弱17区的这种不均一分布。然而,在21a区加药不能改变17区全方位功能图的基本图式。对比加药前和恢复后,加药前和微量注射谷氨酸以及加药前和微量注射GABA的全方位功能图发现,大部分像素点的最优方位变化不大。这一结果意味着21a区的反馈调制可以改变大量17区神经元的最优方位,虽然每个神经元改变的幅度很小,但在总体上却可以引起17区的皮层区域最优方位的分布模式发生改变。
结果提示,从高级皮层的反馈调制可以自上而下地调动低级视皮层中更多的神经元来处理某一种特定的自下而上的视觉信息,使得视觉系统对该信息的敏感度较以前有更大的提高。反馈调制在信息处理上的这种偏好性,可能是高级视皮层的一种重要的生理功能。
Few researches have been done on how the mammalian visual cortex encodes the disappearance of visual stimuli. Here we study the dynamic response of 88 area 17 neurons in cats, from both the stimulus onset and offset. A reliable offset transient response to stationary grating could be seen in 62.5% of the neurons (55 in 88, 31 simple, 24 complex), showing a similar preferred orientation, orientation bias, cumulative detecability, peak response amplitude and latency with that of the onset transient response. However, this offset response, was significantly more dependent on the stimulus duration than the onset transient response, indicating that it needs longer time for the neurocircuit to coactivate in generating offset transient response. Simple and complex cells behaved differently in offset transient response. Complex cells exhibited a stronger linear relationship between the onset and offset response peak latency than simple cells, a greater offset peak response amplitude compared with that of the onset response, and were independent of grating spatial phase both in the onset and offset response in contrast to the simple cells. The linear fading-grating test demonstrated that it was the luminance change in receptive field subregions that generated offset transient response.
Visual system of human and some other mammalians are physiologically or psychologically more sensitive to vertical and horizontal visual stimuli than to those obliquely oriented (usually referred to as the 'oblique effect'). Although the neurobiological basis of this bias is not fully understood, one possibility is that more neural machinery is devoted to processing vertical and horizontal contours than to processing oblique ones. The over representation of vertical and horizontal orientations in an orientation map has been found in area 17 as well as area 21a in cat's visual cortex which receives input from and sends output to area 17. The over representation was greater in area 21a than in area 17. It is interesting to investigate whether feedback influence from area 21a could affect the unequal representation in area 17. Using a combination of intrinsic signal optic imaging and pharmacological techniques, we found for the first time that on average, increasing activity of area 21a enhanced the difference of over representation in area 17 from 4.9% to 18.8%, near the level found in area 21a (23.2%), while decreasing activity of area 21a reduced the over representation. However, the basic pattern of orientation map sampled before and during drug application maintained unchanged, and the preferred orientation of most pixels in the orientation map changed little between control and recovery, control and glutamate microinjection, as well as control and GABA microinjection. These findings imply that positive feedback modulation from area 21a alters the preferred orientation of a large number of neurons in area 17, although in a small degree of switch, thus making the global distribution of preferred orientation changed.
The finding suggests that feedback from higher-order cortex can recruit more neural machinery in lower-order cortex in order to process certain kind of information, thus making an important contribution to higher level of visual function.
人类和其他一些哺乳类对水平-垂直的视觉刺激的敏感度要高于倾斜方位的刺激(这一现象被称为“倾斜效应”)。虽然产生这种效应的神经机制目前尚不很清楚,但是一种可能性是:参与处理水平-垂直方位信息的神经元数量要大于参与处理倾斜方位的。猫视皮层21a区接受17区输入并有反馈投射至17区。本实验室以往的研究发现猫视皮层17区和21a区内对水平-垂直方位敏感的皮层区域明显大于对倾斜方位敏感的皮层区域。而且在21a区的倾斜效应更为明显。本文利用内源信号光学成像技术结合定点微量注射药物的方法,探索高级视皮层21a区的反馈调制对初级视皮层17区的倾斜效应得影响。我们首次发现,增加21a区的活动水平可以使17区的水平-垂直与倾斜方位的敏感区的不均一分布的差值水平从4.9%上升到18.8%,已经接近21a区的23.2%的水平,而降低21a区的活动水平则可以减弱17区的这种不均一分布。然而,在21a区加药不能改变17区全方位功能图的基本图式。对比加药前和恢复后,加药前和微量注射谷氨酸以及加药前和微量注射GABA的全方位功能图发现,大部分像素点的最优方位变化不大。这一结果意味着21a区的反馈调制可以改变大量17区神经元的最优方位,虽然每个神经元改变的幅度很小,但在总体上却可以引起17区的皮层区域最优方位的分布模式发生改变。
结果提示,从高级皮层的反馈调制可以自上而下地调动低级视皮层中更多的神经元来处理某一种特定的自下而上的视觉信息,使得视觉系统对该信息的敏感度较以前有更大的提高。反馈调制在信息处理上的这种偏好性,可能是高级视皮层的一种重要的生理功能。
Few researches have been done on how the mammalian visual cortex encodes the disappearance of visual stimuli. Here we study the dynamic response of 88 area 17 neurons in cats, from both the stimulus onset and offset. A reliable offset transient response to stationary grating could be seen in 62.5% of the neurons (55 in 88, 31 simple, 24 complex), showing a similar preferred orientation, orientation bias, cumulative detecability, peak response amplitude and latency with that of the onset transient response. However, this offset response, was significantly more dependent on the stimulus duration than the onset transient response, indicating that it needs longer time for the neurocircuit to coactivate in generating offset transient response. Simple and complex cells behaved differently in offset transient response. Complex cells exhibited a stronger linear relationship between the onset and offset response peak latency than simple cells, a greater offset peak response amplitude compared with that of the onset response, and were independent of grating spatial phase both in the onset and offset response in contrast to the simple cells. The linear fading-grating test demonstrated that it was the luminance change in receptive field subregions that generated offset transient response.
Visual system of human and some other mammalians are physiologically or psychologically more sensitive to vertical and horizontal visual stimuli than to those obliquely oriented (usually referred to as the 'oblique effect'). Although the neurobiological basis of this bias is not fully understood, one possibility is that more neural machinery is devoted to processing vertical and horizontal contours than to processing oblique ones. The over representation of vertical and horizontal orientations in an orientation map has been found in area 17 as well as area 21a in cat's visual cortex which receives input from and sends output to area 17. The over representation was greater in area 21a than in area 17. It is interesting to investigate whether feedback influence from area 21a could affect the unequal representation in area 17. Using a combination of intrinsic signal optic imaging and pharmacological techniques, we found for the first time that on average, increasing activity of area 21a enhanced the difference of over representation in area 17 from 4.9% to 18.8%, near the level found in area 21a (23.2%), while decreasing activity of area 21a reduced the over representation. However, the basic pattern of orientation map sampled before and during drug application maintained unchanged, and the preferred orientation of most pixels in the orientation map changed little between control and recovery, control and glutamate microinjection, as well as control and GABA microinjection. These findings imply that positive feedback modulation from area 21a alters the preferred orientation of a large number of neurons in area 17, although in a small degree of switch, thus making the global distribution of preferred orientation changed.
The finding suggests that feedback from higher-order cortex can recruit more neural machinery in lower-order cortex in order to process certain kind of information, thus making an important contribution to higher level of visual function.
引文
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