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 orientation-independent 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.     

Frequency-intensity characteristics of cricket cercal interneurons: units with high-pass functions

Interneurons in the cercal sensory system of crickets respond in a cell-specific manner if the cercal hair sensilla are stimulated by air-particle oscillations at frequencies below about 2000 Hz. We investigated the filter properties of several of these interneurons, and tested the effect of stimulus intensity (typically 0.3–50 mm s⁻¹ peak-to-peak air-particle velocity) on the frequency response in the range 5–600 Hz. We focus on three interneurons (the lateral and medial giant interneurons and interneuron 9-3a) of Acheta domesticus which are characterized by a relatively high sensitivity above ca. 50–200 Hz. The responses of the medial giant interneuron usually increase monotonically with frequency and intensity. Interneuron 9-3a and the lateral giant interneuron exhibit saturation or response decrement at high frequencies and intensities. The lateral giant interneuron has an additional peak of sensitivity below about 40 Hz. Small individual variations in the relative locations of the two response areas of this interneuron within the frequency-intensity field are responsible for a large variability obtained if frequency-response curves are determined for particular intensities. Stimulus frequency does not affect the principal directional preferences of the three interneurons. Nevertheless, if tested individually, the lateral giant interneuron and interneuron 9-3a exhibit small changes of directional tuning. Accepted: 12 November 1997     

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.     

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.     

Tuning the cochlea: wave-mediated positive feedback between cells

Frequency analysis by the mammalian cochlea is traditionally thought to occur via a hydrodynamically coupled ‘travelling wave’ along the basilar membrane. A persistent difficulty with this picture is how sharp tuning can emerge. This paper proposes, and models, a supplementary or alternative mechanism: it supposes that the cochlea analyses sound by setting up standing waves between parallel rows of outer hair cells. In this scheme, multiple cells mutually interact through positive feedback of wave-borne energy. Analytical modelling and numerical evaluation presented here demonstrate that this can provide narrow-band frequency analysis. Graded cochlear tuning will then rely on the distance between rows becoming greater as distance from the base increases (as exhibited by the actual cochlea) and on the wave’s phase velocity becoming slower. In effect, tuning is now a case of varying the feedback delay between the rows, and a prime candidate for a wave exhibiting suitably graded phase velocity—a short-wavelength ‘squirting wave’—is identified and used in the modelling. In this way, resonance between rows could supply both amplification and high Q, characteristics underlying the ‘cochlear amplifier’—the device whose action has long been evident to auditory science but whose anatomical basis and mode of operation are still obscure.     

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.     

Interval-counting neurons in the anuran auditory midbrain: factors underlying diversity of interval tuning

In anurans, the temporal patterning of sound pulses is the primary information used for differentiating between spectrally similar calls. One class of midbrain neurons, referred to as ‘interval-counting’ cells, appears to be particularly important for discriminating among calls that differ in pulse repetition rate (PRR). These cells only respond after several pulses are presented with appropriate interpulse intervals. Here we show that the range of selectivity and sharpness of interval tuning vary considerably across neurons. Whole-cell recordings revealed that neurons showing temporally summating excitatory postsynaptic potentials (EPSPs) with little or no inhibition or activity-dependent enhancement of excitation exhibited low-pass or band-pass tuning to slow PRRs. Neurons that showed inhibition and rate-dependent enhancement of excitation, however, were band-pass or high-pass to intermediate or fast PRRs. Surprisingly, across cells, interval tuning based on membrane depolarization and spike rate measures were not significantly correlated. Neurons that lacked inhibition showed the greatest disparities between these two measures of interval tuning. Cells that showed broad membrane potential-based tuning, for example, varied considerably in their spike rate-based tuning; narrow spike rate-based tuning resulted from ‘thresholding’ processes, whereby only the largest depolarizations triggered spikes. The potential constraints associated with generating interval tuning in this manner are discussed.     

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     

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.     

Frequency-intensity characteristics of cricket cercal interneurons: broadband units

The bilateral pairs of cercal interneurons 10-2a and 10-3a in the cricket terminal ganglion are supposed to constitute a functional system for measuring the direction of air-borne signals, based on their phase-locked responses and selective directional sensitivity. The purpose of this study was to obtain information on the frequency and intensity characteristics and thus the potential working range of this system. By recording intracellularly from the axons of the interneurons we measured responses for stimuli of varying frequency, intensity, and direction. Typically, the stimulus frequency range examined extended from 5 to 600 Hz, at intensities of 0.03–30 mm s⁻¹ (peak-to-peak air-particle velocity). The results show that interneurons 10-2a and 10-3a preserved their level of activity, response type, and direction tuning in the whole frequency range tested. Stimulus-response cross-correlograms revealed that spike trains were synchronized with stimulus waves at even higher frequencies, at least up to 1000 Hz. At a given air-particle velocity in the range of about 2–2.5 logarithmic units, the spike number responses of the interneurons were nearly constant over a wide frequency range. Directional diagrams appeared to be independent of stimulus frequency, both in orientation and in amplitude. Accepted: 14 October 1998     

Contribution of GABAA receptor-mediated inhibition to the determination of the velocity tuning of low pass-tuned neurons of the Superior Colliculus

The participation of intrinsic inhibitory networks in providing the velocity selectivity of neurons of the superior colliculus (SC) of the Syrian hamster was tested using iontophoretic application of bicuculline methiodide, a GABA_A receptor competitive antagonist. The impulse activity of 22 low pass-tuned (LP) cells was recorded extracellularly. Following application of bicuculline, 10 cells exhibited an increase in the velocity selectivity, while the other 12 units showed decreases in their tuning. We assume that SC intrinsic inhibitory networks contributing to the velocity tuning of neurons of this structure are driven in a dissimilar way by afferent volleys arriving from the retina through “fast” Y and “slow” W channels. Neirofiziologiya/Neurophysiology, Vol. 39, Nos. 4/5, pp. 385–387, July–October, 2007.     

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     

Dependence of the threshold of sinusoidal grating displacement on its spatial frequency characteristics

The threshold of detection of sinusoidal grating displacement was studied under conditions of the absence of apparent movement. It was shown that the threshold rises with the pattern size and depends on the number of grating cycles at a constant stimulus magnitude. The stimulus size in 2–3 periods is optimal for its localization. The results suggest that objects in the visual field are localized by mechanisms tuned to a certain spatial frequency and selective to a certain image size. The size of the analyzed part of the visual field requires respective frequency tuning of the mechanism.     

A comparison of resonance tuning with positive versus negative sensory feedback

We used a computational model of rhythmic movement to analyze how the connectivity of sensory feedback affects the tuning of a closed-loop neuromechanical system to the mechanical resonant frequency (ω_r). Our model includes a Matsuoka half-center oscillator for a central pattern generator (CPG) and a linear, one-degree-of-freedom system for a mechanical component. Using both an open-loop frequency response analysis and closed-loop simulations, we compared resonance tuning with four different feedback configurations as the mechanical resonant frequency, feedback gain, and mechanical damping varied. The feedback configurations consisted of two negative and two positive feedback connectivity schemes. We found that with negative feedback, resonance tuning predominantly occurred when ω_r was higher than the CPG’s endogenous frequency (ω_CPG). In contrast, with the two positive feedback configurations, resonance tuning only occurred if ω_r was lower than ω_CPG. Moreover, the differences in resonance tuning between the two positive (negative) feedback configurations increased with increasing feedback gain and with decreasing mechanical damping. Our results indicate that resonance tuning can be achieved with positive feedback. Furthermore, we have shown that the feedback configuration affects the parameter space over which the endogenous frequency of the CPG or resonant frequency the mechanical dynamics dominates the frequency of a rhythmic movement.     

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.     

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 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