Directional tunings independent of orientation in the primary visual cortex of the cat

A family of moving ‘random-line’ patterns was developed and used to study the directional tuning of 91 single units in cat primary visual cortex (V1). The results suggest that, in addition to the well-known orientation-dependent mechanism, there is also some kind of orientationindependent mechanism underlying the direction selectivity. The directional tuning of the neurons varies in accordance with the increase of orientation or non-orientation element in the stimulus.     

用数量场描述方向敏感感受野

1965年Barlow ＆ Levick及我国科技工作者分别独立提出了原理大致相同的方向检测感受野模型,但其数学模型却一直未见有报道.本文用数量场方法给出了其数学模型,这是一个Gabor函数模型,并据此解释了方向检测感受野的方向调谐曲线以及方向检测通道数量问题.     

Neural Tuning Functions Underlie Both Generalization and Interference

In sports, the role of backswing is considered critical for generating a good shot, even though it plays no direct role in hitting the ball. We recently demonstrated the scientific basis of this phenomenon by showing that immediate past movement affects the learning and recall of motor memories. This effect occurred regardless of whether the past contextual movement was performed actively, passively, or shown visually. In force field studies, it has been shown that motor memories generalize locally and that the level of compensation decays as a function of movement angle away from the trained movement. Here we examine if the contextual effect of past movement exhibits similar patterns of generalization and whether it can explain behavior seen in interference studies. Using a single force-field learning task, the directional tuning curves of both the prior contextual movement and the subsequent force field adaptive movements were measured. The adaptation movement direction showed strong directional tuning, decaying to zero by 90° relative to the training direction. The contextual movement direction exhibited a similar directional tuning, although the effect was always above 60%. We then investigated the directional tuning of the passive contextual movement using interference tasks, where the contextual movements that uniquely specified the force field direction were separated by ±15° or ±45°. Both groups showed a pronounced tuning effect, which could be well explained by the directional tuning functions for single force fields. Our results show that contextual effect of past movement influences predictive force compensation, even when adaptation does not require contextual information. However, when such past movement contextual information is crucial to the task, such as in an interference study, it plays a strong role in motor memory learning and recall. This work demonstrates that similar tuning responses underlie both generalization of movement direction during dynamic learning and contextual movements in both single force field and interference tasks.     

Directional tuning of visual cortex neurons in the cat before and after nembutal injection

Directional tuning was investigated in 40 neurons of the primary visual cortex (area 17) before and after Nembutal injection during acute experiments on immobilized cats. Preferred orientation (PO) in 50% of neurons was found to be stable after the drug, while the remainder showed a consistent shift in PO (averaging 53.6±8.0°) for a number of hours. Neurons with consistent PO more frequently showed a preference for horizontal and vertical stimulus orientation; cells with unstable tuning had a wider PO distribution. More refined directional detection (i.e., finer tuning) was noted in "stable" rather than in "unstable" neurons both before and after administering the drug. Under narcosis, directional tuning altered in 50% of cells — an effect more marked in "unstable" than in "stable" cells (68% as against 38%). Mean background discharge rate also fell by an average of 5.5-fold and induced firing rate declined 1.5-fold during narcosis, moreover.Institute of Higher Nervous Activity and Neurophysiology, Russian Academy of Sciences, Moscow. Translated from Neirofiziologiya, Vol. 23, No. 6, pp. 669–676, November–December, 1991.     

Orientation scanning effect monitored in neurons of the cat visual cortex

Incidence of the phenomenon of dynamic scanning of a portion of the orientation range during the development of neuronal response in cells of the primary visual cortex was monitored in immobilized cats using a technique involving time bins, having smoothed latencies and estimating only the highly significant portions of their response. It was found that this effect persisted in 13 out of 17 test neurons, actually remaining invariable in seven units, and modified in a further six cells owing to a shift in the starting point of the scanning process along the directional range, either extending the latter or producing a change in the direction of scanning. Directional tuning stabilized in 4 cells only following smoothing of latent periods. Findings indicate that dynamic changes in directional tuning are associated with a restructuring of the time course of response in most neurons, indicative of spatio-temporal directional coding.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translating for Neirofiziologiya, Vol. 19, No. 2, pp. 179–185, March–April, 1987.     

蛙下丘频率调谐特性方向选择性的建模和仿真

基于Ｆｅｎｇ等人的神经电生理实验结果,建立了关于豹蛙（Ｒａｎａ ｐｉｐｉｅｎ）半规隆凸（ｔｏｒｕｓ ｓｅｍｉｃｉｒｃｕｌａｒｉｓ）核团神经元频率调谐特性方向选择性反应的模型,并在此基础上讨论了模型各个参数对模型方向性反应输出的影响。结果提示：１）听觉系统两侧的相互抑制作用－双耳抑制主要决定频率调谐特性的尖锐性随方向变化的特性;２）前级神经元对不同方位外界声源刺激具有不同强度平均脉冲发放的特征则主要     

Matched filters, mate choice and the evolution of sexually selected traits

Background

Fundamental for understanding the evolution of communication systems is both the variation in a signal and how this affects the behavior of receivers, as well as variation in preference functions of receivers, and how this affects the variability of the signal. However, individual differences in female preference functions and their proximate causation have rarely been studied.

Methodology/Principal Findings

Calling songs of male field crickets represent secondary sexual characters and are subject to sexual selection by female choice. Following predictions from the “matched filter hypothesis” we studied the tuning of an identified interneuron in a field cricket, known for its function in phonotaxis, and correlated this with the preference of the same females in two-choice trials. Females vary in their neuronal frequency tuning, which strongly predicts the preference in a choice situation between two songs differing in carrier frequency. A second “matched filter” exists in directional hearing, where reliable cues for sound localization occur only in a narrow frequency range. There is a strong correlation between the directional tuning and the behavioural preference in no-choice tests. This second “matched filter” also varies widely in females, and surprisingly, differs on average by 400 Hz from the neuronal frequency tuning.

Conclusions/Significance

Our findings on the mismatch of the two “matched filters” would suggest that the difference in these two filters appears to be caused by their evolutionary history, and the different trade-offs which exist between sound emission, transmission and detection, as well as directional hearing under specific ecological settings. The mismatched filter situation may ultimately explain the maintenance of considerable variation in the carrier frequency of the male signal despite stabilizing selection.     

Dendritic Spikes Amplify the Synaptic Signal to Enhance Detection of Motion in a Simulation of the Direction-Selective Ganglion Cell

The On-Off direction-selective ganglion cell (DSGC) in mammalian retinas responds most strongly to a stimulus moving in a specific direction. The DSGC initiates spikes in its dendritic tree, which are thought to propagate to the soma with high probability. Both dendritic and somatic spikes in the DSGC display strong directional tuning, whereas somatic PSPs (postsynaptic potentials) are only weakly directional, indicating that spike generation includes marked enhancement of the directional signal. We used a realistic computational model based on anatomical and physiological measurements to determine the source of the enhancement. Our results indicate that the DSGC dendritic tree is partitioned into separate electrotonic regions, each summing its local excitatory and inhibitory synaptic inputs to initiate spikes. Within each local region the local spike threshold nonlinearly amplifies the preferred response over the null response on the basis of PSP amplitude. Using inhibitory conductances previously measured in DSGCs, the simulation results showed that inhibition is only sufficient to prevent spike initiation and cannot affect spike propagation. Therefore, inhibition will only act locally within the dendritic arbor. We identified the role of three mechanisms that generate directional selectivity (DS) in the local dendritic regions. First, a mechanism for DS intrinsic to the dendritic structure of the DSGC enhances DS on the null side of the cell''s dendritic tree and weakens it on the preferred side. Second, spatially offset postsynaptic inhibition generates robust DS in the isolated dendritic tips but weak DS near the soma. Third, presynaptic DS is apparently necessary because it is more robust across the dendritic tree. The pre- and postsynaptic mechanisms together can overcome the local intrinsic DS. These local dendritic mechanisms can perform independent nonlinear computations to make a decision, and there could be analogous mechanisms within cortical circuitry.     

Direction-selective dendritic action potentials in rabbit retina

Dendritic spikes that propagate toward the soma are well documented, but their physiological role remains uncertain. Our in vitro patch-clamp recordings and two-photon calcium imaging show that direction-selective retinal ganglion cells (DSGCs) utilize orthograde dendritic spikes during physiological activity. DSGCs signal the direction of image motion. Excitatory subthreshold postsynaptic potentials are observed in DSGCs for motion in all directions and provide a weakly tuned directional signal. However, spikes are generated over only a narrow range of motion angles, indicating that spike generation greatly enhances directional tuning. Our results indicate that spikes are initiated at multiple sites within the dendritic arbors of DSGCs and that each dendritic spike initiates a somatic spike. We propose that dendritic spike failure, produced by local inhibitory inputs, might be a critical factor that enhances directional tuning of somatic spikes.     

Directional and frequency response characteristics in the descending octaval nucleus of the toadfish (Opsanus tau)

This study is a continuation of a long-term investigation of the auditory circuit in the oyster toadfish, Opsanus tau. Input from the auditory periphery projects to the ipsilateral descending octaval nucleus (DON). Ipsilateral and contralateral DONs project to the auditory midbrain, where a previous study indicated that both frequency tuning and directional sharpening are present. To better understand the transformation of auditory information along the auditory pathway, we have examined over 400 units in the DON to characterize frequency and directional information encoded in the dorsolateral division of the nucleus. Background activity was primarily low (<10 spikes/s) or absent. The maximum coefficient of synchronization was equivalent to the periphery (R = 0.9) and substantially better than in the midbrain. The majority of DON units (79%) responded best to stimulus frequencies of 84-141 Hz and were broadly tuned. DON cells retain or enhance the directional character of their peripheral input (s); however, characteristic axes were distributed in all quadrants around the fish, providing further evidence that binaural computations may first occur in the DON of this species.     

Directional sensitivity of tuberous electroreceptors: polarity preferences and frequency tuning

This paper examines the directionality of tuberous electroreceptor responses and relates them to a polarity bias seen for passive electrolocation by electric fish (Hypopomus). We recorded from Burst Duration Coders (BDCs) while stimulating with 1 kHz single period sine waves with electric fields oriented horizontally in different directions. Electroreceptors have figure-8 directional sensitivity profiles with two, usually unequal lobes of sensitivity separated by 180°. For most units the larger lobe points inward, while for a few, the lobes are symmetrical or the larger lobe points outward. The differences correlate with differences in frequency tuning of the receptors. We can alter, and even reverse, the directional asymmetry of a single unit by changing the frequency of the stimulus. Two general response profiles result, with two corresponding classes of tuning curves. The degree of asymmetry varies with position on the body surface. The asymmetries and the effects of stimulus frequency and of tuning can be modeled with a linear/non-linear/linear cascade filter. The behavioral preference for approaching the head end ( + ) of an electrode is difficult to understand in light of the asymmetry of responses we report for amplitude-coding BDCs but can be understood by reference to the time-coding Pulse Marker (PM) receptors.Abbreviations BDC Burst Duration Coder - EOD electric organ discharge - nALL anterior lateral line nerve - PM Pulse Marker     

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.     

Specificity of sensorimotor learning and the neural code: Neuronal representations in the primary motor cortex

Human studies show that the learning of a new sensorimotor mapping that requires adaptation to directional errors is local and generalizes poorly to untrained directions. We trained monkeys to learn new visuomotor rotations for only one target in space and recorded neuronal activity in the primary motor cortex before, during and after learning. Similar to humans, the monkeys showed poor transfer of learning to other directions, as observed by behavioral aftereffects for untrained directions. To test for internal representations underlying these changes, we compared two features of neuronal activity before and after learning: changes in firing rates and changes in information content. Specific elevations of firing rate were only observed in a subpopulation of cells in the motor cortex with directional properties corresponding to the locally learned rotation; namely cells only showed plasticity if their preferred direction was near the training one. We applied measures from information theory to probe for learning-related changes in the neuronal code. Single cells conveyed more information about the direction of movement and this specific improvement in encoding was correlated with an increase in the slope of the neurons' tuning curve. Further, the improved information after learning enabled a more accurate reconstruction of movement direction from neuronal populations. Our findings suggest a neural mechanism for the confined generalization of a newly acquired internal model by showing a tight relationship between the locality of learning and the properties of neurons. They also provide direct evidence for improvement in the neural code as a result of learning.     

Orientation and spatiotemporal tuning of cells in the primary visual cortex of an Australian marsupial,the wallaby<Emphasis Type="Italic"> Macropus eugenii</Emphasis>

The metatherians (marsupials) have been separated from eutherians (placentals) for approximately 135 million years. It might, therefore, be expected that significant independent evolution of the visual system has occurred. The present paper describes for the first time the orientation, direction and spatiotemporal tuning of neurons in the primary visual cortex of an Australian marsupial, the wallaby Macropus eugenii. The stimuli consisted of spatial sinusoidal gratings presented within apertures covering the classical receptive fields of the cells. The neurons can be classified as those with clear ON and OFF zones and those with less well-defined receptive field structures. Seventy-percent of the total cells encountered were strongly orientation selective (tuning functions at half height were less than 45 degrees ). The preferred orientations were evenly distributed throughout 360 degrees for cells with uniform receptive fields but biased towards the vertical and horizontal for cells with clear ON-OFF zones. Many neurons gave directional responses but only a small percentage of them (4%) showed motion opponent properties (i.e. they were excited by motion in one direction and actively inhibited by motion in the opposite direction). The median peak temporal tuning for cells with clear ON-OFF zones and those without were 3 Hz and 6 Hz, respectively. The most common peak spatial frequency tuning for the two groups were 2 cycles per degree and 0.5 cycles per degree, respectively. Spatiotemporal tuning was not always the same for preferred and antipreferred direction motion. In general, the physiology of the wallaby cortex was similar to well studied eutherian mammals suggesting either convergent evolution or a highly conserved architecture that stems from a common therian ancestor.     

Directional selectivity and frequency tuning of midbrain cells in the oyster toadfish, Opsanus tau

Single-unit recordings were made from areas in the midbrain (torus semicircularis) of the oyster toadfish. We evaluated frequency tuning and directional responses using whole-body oscillation to simulate auditory stimulation by particle motion along axes in the horizontal and mid-sagittal planes. We also tested for bimodality in responses to auditory and hydrodynamic stimuli. One recording location in each animal was marked by a neurobiotin injection to confirm the recording site. Recordings were made in nucleus centralis, nucleus ventrolateralis, and the deep cell layer. Most units were frequency-selective with best frequencies between 50 and 141 Hz. Suppression of activity was apparent in 10% of the cells. Bimodality was common, including inhibition and suppression of background activity by auditory or hydrodynamic stimulation. The majority of the cells were directionally selective with directional response patterns that were sharpened compared with those of primary saccular afferents. The best directional axes were arrayed widely in spherical space, covering most azimuths and elevations. This representation is adequate for the computation of the motional axis of an auditory stimulus for sound source localization.Abbreviations BF best frequency - DCL deep cell layer - DON descending octaval nucleus - DRP directional response pattern - FFT fast Fourier transform - LL lateral lemniscus - NC nucleus centralis - NVL nucleus ventrolateralis - PVC periventricular cells - R coefficient of synchronization - TS torus semicircularis - Z Rayleigh statistic     

Optic flow representation in the optic lobes of Diptera: modeling the role of T5 directional tuning properties

An evolutionarily conserved system of small retinotopic neurons in dipteran insects, called bushy T-cells, provides information about directional motion to large collator neurons in the lobula plate. Physiological and anatomical features of these cells provide the basis for a model that is used to investigate requirements for generating optic flow selectivity in collators while allowing for evolutionary variations. This account focuses on the role of physiological tuning properties of T5 neurons. Various flow fields are defined as inputs to retinotopic arrays of T5 cells, the responses of which are mapped onto collators using innervation matrices that promote selectivity for flow type and position. Properties known or inferred from physiological and anatomical studies of neurons contributing to motion detection are incorporated into the model: broad tuning to local motion direction and the representation of each visual sampling unit by a quartet of small-field T5-like neurons with orthogonal preferred directions. The model predicts hitherto untested response properties of optic flow selective collators, and predicts that selectivity for a given flow field can be highly sensitive to perturbations in physiological properties of the motion detectors.     

Population vector code: a geometric universal as actuator

The population vector code relates directional tuning of single cells and global, directional motion incited by an assembly of neurons. In this paper three things are done. First, we analyze the population vector code as a purely geometric construct, focusing attention on its universality. Second, we generalize the algorithm on the basis of its geometrical realization so that the same construct that responds to sensation can function as an actuator for behavioral output. Third, we suggest at least a partial answer to the question of what many maps, neuronal representations of the outside sensory world in space-time, are good for: encoding vectorial input they enable a direct realization of the population vector code.     

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。     

Development of optokinetic neuronal responses in the pretectum and horizontal optokinetic nystagmus in unilaterally enucleated rats

Responses of single units to constant-velocity rotations of the visual surround (0.25-10 degrees/s) were studied in the pretectum of unilateral enucleated rats at different ages. Enucleation was performed either in the first postnatal week ("early" enucleated rats) or in the adult stage ("late" enucleated rats). Pretectal unitary responses were recorded in early enucleated animals at postnatal day 20-21, 36-49 and, in both experimental groups, in the adult stage. Optokinetic ocular nystagmus was studied in early and late enucleated rats in the adult stage. Gain of optokinetic nystagmus in temporo-nasal stimulus direction was not changed for visual surround rotations of up to 20 degrees/s compared to controls in monocular viewing conditions. At higher stimulus velocities, however, the gain dropped. In naso-temporal stimulus direction, optokinetic nystagmus was improved in gain for optokinetic pattern motions of up to 5-10 degrees/s. There were only minor differences in the gain behaviour of optokinetic nystagmus obtained from early or late enucleated rats. The optokinetic responses of pretectal neurons obtained from early and late enucleated rats were reduced in sensitivity by more than 50%. The response patterns of neurons recorded in the contralateral pretectum relative to the intact eye were shifted by a large amount from directional selective to directional nonselective response types. No such changes were obtained in the ipsilateral pretectum. In contrast to normal rats, there were very few directional selective units responding to temporo-nasal pattern motion. On the other hand, a large proportion of directional selective units responded to naso-temporal pattern motion. These latter units were found in both early and late enucleated rats. A similar response type has previously been described for intact young rats but not for adult rats. The velocity tuning curve of pretectal units studied in the adult stage was similar in shape in early and late enucleated rats and resembled that obtained from enucleated or intact young animals. Our results show that response sensitivity, direction and velocity tuning of pretectal units depend crucially on retinal afferent input originating from both eyes. The data suggest that the response characteristics of many of the pretectal units that are considered to be important for mediating optokinetic reflexes depend on interpretectal signal processing using commissural connections. There is very little evidence for an adaptative structural plasticity of the optokinetic system following loss of one eye. The reduced asymmetry observed in gain of optokinetic responses correlated in both early and late enucleated rats with the shifts observed in the distribution of pretectal unitary response patterns.     

Motor learning with unstable neural representations

It is often assumed that learning takes place by changing an otherwise stable neural representation. To test this assumption, we studied changes in the directional tuning of primate motor cortical neurons during reaching movements performed in familiar and novel environments. During the familiar task, tuning curves exhibited slow random drift. During learning of the novel task, random drift was accompanied by systematic shifts of tuning curves. Our analysis suggests that motor learning is based on a surprisingly unstable neural representation. To explain these results, we propose that motor cortex is a redundant neural network, i.e., any single behavior can be realized by multiple configurations of synaptic strengths. We further hypothesize that synaptic modifications underlying learning contain a random component, which causes wandering among synaptic configurations with equivalent behaviors but different neural representations. We use a simple model to explore the implications of these assumptions.