胼胝体输入对猫皮层17/18交界区细胞方向和取向选择性的影响

用抑制性神经递质GABA阻断胼胝体输入、用微机控制的运动光棒作为视觉刺激,用金属电极胞外记录技术,研究猫皮层17/18区交界附近细胞方向选择性和取向选择性的变化.在被检测的48个细胞中,50%细胞的方向选择性强度,54.2%细胞的取向选择性强度发生了改变;约20%细胞的最优反应方向或.及最优取向发生了10-30°的偏移;共有56.2%细胞的方向选择性、58.3%细胞的取向选择性受到明确的影响.这些结果表明胼胝体对皮层细胞视觉反应的贡献是多方面的.     

视觉细胞反应中方向选择性和取向选择性成分的分离

运动的光刺激常被用于研究视觉神经细胞的方向和取向选择性.但是,在细胞的反应中,两种因素同时存在且混杂在一起; 以往的各种分析方法均因未能有效地解决这一问题而不够准确,对此我们给出了理论和实例证明.本工作引入正余弦函数系对实验数据作最优回归分析.拟合程度好于以往的方法.并且从根本上分离了反应中的方向和取向选择性成分.使对这两种性质的准确定量成为可能.本文对此方法的内在含义作出了解释,并提出了对方向、取向选择性及最优方向、取向的理解.     

猫上丘视觉神经元的取向和方向选择性

基于对上丘细胞对运动光棒刺激反应特性的定量分析,首次明确提出猫上丘视觉神经元具有一定的取向选择性,并从形态基础和生理功能的角度讨论了这种选择性存在的意义。另外还发现上丘的方向和取向选择性均弱于初级视皮层,且对向视野上、下方运动的光棒较为敏感。     

冷冻取消胼胝体传递对猫皮层17/18区边界细胞部分电生理特性的影响

用IBM-XT微机控制产生各种取向的可移动正弦光栅作为视觉刺激,用金属微电极在猫皮层一侧17/18区边界附近进行胞外记录,神经信号由微机实时采集并进行数据处理;对11只猫的62个细胞进行了冷冻取消另一侧胼胝体向记录侧传递前后细胞的取向选择性,最佳取向和最大反应的测定与比较.发现冷冻后有至少50%细胞的取向选择性强度发生了变化,其中选择性减少与增加的数目基本相等;有20-30%细胞的最佳取向发产了10°至20°的变化;有一半以上细胞的最大反应发生了变化,其中三分之二细胞的反应减少,其余的增加.通过部分细胞的恢复实验证明以上变化基本是可逆的,说明胼胝体对这些特性有一定影响.     

猫初级视皮层神经元不依赖于图形取向的方向调谐

设计并利用运动“随机”线条图研究了猫初级视皮层(V1)共91个细胞的方向调谐特性.结果表明, V1区神经元的方向选择性除了存在基于图形成分取向的机制以外,还存在不基于取向因素的机制.当刺激图形中的取向或非取向因素增强时,神经元的方向调谐也会相应转变.     

纹外皮层PMLS区反馈投射对初级视皮层神经元反应特性的影响

神经系统中存在大量下行投射,与上行输入一起形成复杂的前馈与反馈回路,调控神经信号的传导和处理,但目前对皮层内反馈投射的功能作用认识还比较薄弱.通过微量注射抑制性神经递质γ-氨基丁酸(γ-aminobutyric acid,GABA),使猫纹外皮层后内侧外上雪氏区(area posteromedial lateral suprasylvian,PMLS)局部可逆性失活,使用胞外记录方法,研究初级视皮层17区神经元反应特性的变化.实验结果显示,PMLS区失活后,17区细胞对运动刺激的反应总体减弱,反应的相对稳定性基本不变,最高发放率/自发之比有所下降.与此同时,细胞的方向选择性指数减小,朝向选择性无显著变化.除少数"双向"反应细胞外,绝大部分细胞的最优方向基本不变.进一步分析发现,细胞对各个方向刺激的反应普遍下降,最优方向上的下降程度最大,是导致方向选择性减弱的主要原因.这些结果表明,PMLS区反馈投射可增强初级视皮层的方向选择性,而对朝向选择性影响有限.这一作用特点体现了PMLS区在皮层中偏重处理运动视觉信息的功能.     

猫视皮层17,18区神经元对错觉轮廓的反应

研究了轻度麻醉下猫视皮层17, 18区细胞对错觉轮廓刺激的反应特性, 比较了对错觉轮廓有明显反应的细胞对真实轮廓和错觉轮廓刺激的感受野特性的异同. 共记录了猫视皮层17, 18区200个方位/方向选择性细胞, 其中有42%的细胞是错觉轮廓反应细胞. 将这些细胞对真实轮廓和错觉轮廓的反应进行比较, 尽管错觉轮廓反应细胞对移动光棒和错觉轮廓光棒的方位/方向调制曲线十分相似, 但对移动错觉棒和移动光棒的反应模式(潜伏期和反应时程)不同. 对由光栅组成的错觉轮廓而言, 细胞的反应大小与组成光栅的相位无关, 并且细胞对组成错觉轮廓光栅的最优空间频率比对普通移动光栅的最优空间频率要高得多, 说明细胞确实是对轮廓本身反应, 而不是对组成轮廓的光栅的末端反应. 某些速度调制类型的细胞对移动错觉棒反应的最优速度比对移动光棒的最优速度要低得多. 进一步验证了猫视皮层17, 18区部分细胞能对错觉轮廓反应, 并且观察到这些细胞对错觉轮廓和真实轮廓有不同的感受野反应特性, 提示视觉系统对两种刺激图形的检测机制可能存在着差异.     

猫前内侧上雪氏区视神经元对运动随机线条的反应

对于运动信息在脑内的加工,一种观点认为分两阶段进行,低级视皮层只对运动图形内部成分的取向进行调谐,高级视皮层整合低级视皮层的输入,对图形整体的运动方向敏感。用网格（plaid）作为刺激的实验表明,在较低级皮层区,细胞多表现为成分方向选择性（Component-motion Selectivity),即对刺激中的取向因素敏感：而较高视皮层的细胞多表现为整体方向选择性（Pattern-motion Selecitivity),对运动整体的方向敏感,从而支持运动信息加工的“两阶段”理论。实验中,用一系列运动随机线条刺激（random line patterns)。研究猫前内侧上雪氏区（Anteriormedial lateral suprasylvian area,AMLS)神经元的方向调谐特性。结果表明多数细胞为整体方向选择性,且随线长增加此类细胞比例下降,而成分方向选择性细胞的比例有所增加,呈现由整体方向选择性向中间类型（Unclassified),由中间类型向成分方向选择性变化的趋势,提示整体或成分方向选择性可能并非细胞的固有特性,而是可以随刺激取向因素的变化而改变的。     

GABA 及荷包牡丹碱对金黄地鼠上丘视觉神经元反应性质的影响

金黄地鼠上丘的浅层细胞为视觉神经元,在记录处电泳γ-氨基丁酸(GABA)或其拮抗剂荷包牡丹碱(bicuculline简写为Bicu)可以改变细胞对视觉刺激的反应特性。GABA 降低细胞对闪光的反应强度,Bicu 则提高细胞对闪光的反应强度,并削弱外周抑制。电泳 GABA 使76.6%(n=60)的细胞反应下降,而 Bicu 使90.0%的细胞反应增强。对自发活动也有类似的影响,Bicu 还使65.0%(n=60)的细胞感受野增大.此外这两种药物对细胞的方位选择性也有一定影响。     

初级视皮层神经元响应反应与撤反应的朝向选择性

朝向选择性是初级视皮层(17区或V1)神经元的基本性质,在图形感知中起着关键作用.同时这些神经元对于持续时间大于100 ms的视觉刺激具有清晰的响应反应(Onset responses)和撤反应(Offset responses).以往的研究只关注响应反应的朝向选择性,而忽视了对撤反应的朝向选择性研究.我们比较了响应与撤反应的朝向调谐性质,大多数细胞的撤反应与响应反应基本上具有相似的最优朝向,而撤反应的朝向调谐宽度有窄于响应反应的趋势,撤反应的最优延迟普遍滞后于响应反应的最优延迟.撤反应的朝向选择性略强于响应反应和具有显著长的反应延迟提示,皮层内的反馈输入可能在形成撤反应的朝向选择性中起着作用.本研究揭示了撤反应的朝向选择性在刺激朝向的连续表征和主体在形状知觉的后期对朝向的精细区分中起着作用.     

Quantitative determination of orientational and directional components in the response of visual cortical cells to moving stimuli

The response characteristic of visual cortical cells to moving oriented stimuli consists mainly of directional (D) and orientational (O) components superimposed to a spontaneous activity (S). Commonly used polar plot diagrams reflect the maximal responses for different orientations and directions of stimulus movement with a periodicity of 360 degrees in the visual field. Fast Fourier analysis (FFT) is applied to polar plot data in order to determine the intermingled S, D, and O components. The zero order gain component of the spectrum corresponds to a (virtual) spontaneous activity. The first order component is interpreted as the strength of the direction selectivity and the second order component as the strength of the orientation specificity. The axes of the preferred direction and optimal orientation are represented by the respective phase values. Experimental data are well described with these parameters and relative changes of the shape of a polar plot can be detected with an accuracy better than 1%. The results are compatible with a model of converging excitatory and inhibitory inputs weighted according to the zero to second order components of the Fourier analysis. The easily performed quantitative determination of the S, D, and O components allows the study of pharmacologically induced changes in the dynamic response characteristics of single visual cortical cells.     

Quantification of directional and orientational selectivities of visual neurons to moving stimuli

Directional and orientational components usually coexist and are mixed in the cell's overall responses when moving optical stimuli are used to study the response characteristics of visual neurons. While these two properties were quantified with all the previous methods for data analysis, their effects could not be efficiently separated from each other, and thus the analyses were imperfect. In this paper, theoretical evidence and examples are provided to show the defects of the old methods. In order to separate the two components completely, we propose to apply optimal regression analysis with the sine-cosine function series as the fundamental variables. Based on this separation, we defined the orientational selectivity as variation of response strength with orientation and performed integration and averaging to quantify the two properties [cf. Eqs. (5) and (6)]. The present method has the advantages of completeness and accuracy, and can detect some details which would have been missed by other methods. An explanation of the intrinsic implications of the method and our comprehension of directional and orientational selectivities and preferred direction and orientation are also given. Received: 4 January 1993/Accepted in revised form: 1 July 1993     

猫后内侧上雪氏区视神经元对运动棒反应的取向和方向特征

猫后内侧上雪区（ｐｏｓｔｅｒｏｍｅｄｉａｌｌａｔｅｒａｌｓｕｐｒａｓｙｌｖｉａｎａｒｅａ,ＰＭＬＳ）的绝大多数神经元（１７１／２００）对运动棒的取向调谐,６２％（１２４／２００）细胞的取向调谐宽度（半高波宽）小于９０°：按方向选择性和取向选择性可分辨出几类特征明显的细胞类型：１、强取向和强方向选择性细胞;２、强取向调谐的双向选择细胞;３、弱取向调谐的强方向选择细胞;４、无取向无方向选择性细胞;以及５、特征不明显的或中间类型细胞。它们与最近光学记录揭示的鹰猴中颞叶视区（ｍｉｄｄｌｅｔｅｍｐｏｒａｌｖｉｓｕａｌａｒｅａ,ＭＴ）的组织有很好的吻合。     

Orientational and directional selectivities of visual neurons in the superior colliculus of the cat

Based on quantitative analyses of the response characteristics of visual neurons in the superior colliculus to moving optical bar stimuli, it is demonstrated for the first time that the visual neurons in superior colliculus of the cat have, to some extent, orientational selectivity. The significance of this selectivity is discussed in reference to its morphological substrate and physiological functions. In addition, both the directional and orientational selectivities in the superior colliculus are relatively weak when compared with those in the primary visual cortex, and the majority of the neurons prefer upward or downward motion in the visual field.     

How to unconfound the directional and orientational information in visual neuron's response

When drifting bars or gratings are used as visual stimuli, information about orientation specificity (which has a period of 180°) and direction specificity (which has a period of 360°) is inherently confounded in the response of visual cortical neurons, which have long been known to be selective for both the orientation of the stimulus and the direction of its movement. It is essential to unconfound or separate these two components of the response as they may respectively contribute to form and motion perception, two of the main streams of information processing in the mammalian brain. Wörgötter and Eysel (1987) recently proposed the Fourier transform technique as a method of unconfounding the two components, but their analysis was incomplete. Here we formally develop the mathematical tools for this method to calculate the peak angles, bandwidths, and relative strengths, the three most important elements of a tuning curve, of both the orientational and the directional components, based on the experimentally-recorded neuron's response polar-plot. It will be shown that, in the 1-D Fourier decomposition of the polar-plot along its angular dimension, (1) the odd harmonics contain only the directional component, while the even harmonics are contributed to by both the orientational and the directional components; (2) the phases and the amplitudes of all the harmonics are related, respectively, to the peak angle and the bandwidth of the individual component. The basic assumption used here is that the two components are linearly additive; this in turn is immediately testable by the method itself.     

The spatial substructure of visual receptive fields in the cat's superior colliculus

Although the direction selective properties of the superficial layer cells of the cat's superior colliculus have been extensively studied, the mechanisms underlying this property remain controversial. With the aim to understand the mechanism(s) underlying directional selectivity of collicular neurons we examined the substructure of their visual receptive fields. 1. The strength of cell responses and the direction selectivity indices varied in relation to the location of the tested region within the receptive field and the amplitude of stimulus movement. 2. Decrease of the amplitude of motion resulted in a decrease of direction selectivity index both in the group of direction-selective cells and in the group of cells classified as direction nonselective but with a directional bias. 3. The decrease of direction selectivity for small amplitude movement resulted mainly from increase in the magnitude of response in the nonpreferred direction of movement. 4. These results suggest that the receptive fields of most collicular cells are composed of subregions with different response profiles and indicate that inhibitory mechanisms dictate direction selectivity of collicular cells.     

Sharpening of directional responses along the auditory pathway of the oyster toadfish, <Emphasis Type="Italic">Opsanus tau</Emphasis>

Our previous studies have shown that the peripheral auditory system of the toadfish encodes the direction of a sound source. Here, we compare directional responses of peripheral saccular afferents, cells in the descending octaval nucleus (DON) of the medulla, and the torus semicircularis (TS) of the midbrain. Recording locations in the brain were labeled with neurobiotin to confirm the site. To compare directional responses among cells, we calculated an index [sharpening ratio (SR)] that weights the relative strength of responses to the best direction for that cell and to the adjacent stimulus angles tested. Unsharpened saccular afferents tend to have a cosinusoidal directional response pattern (DRP) with an expected SR of 0.87. In DON, more than 60% of the cells exhibited directional sharpening (defined as SR <0.8). In TS, more than 80% of the cells exhibited directional sharpening. We conclude that directional auditory sharpening first occurs in DON and some additional sharpening occurs in the ascending pathway to the midbrain, particularly in azimuth. The sharpening of directional selectivity is likely to be an important component of the neural computations underlying directional hearing.