Emergence of spatiotemporal receptive fields and its application to motion detection

A model of motion sensitivity as observed in some cells of area V1 of the visual cortex is proposed. Motion sensitivity is achieved by a combination of different spatiotemporal receptive fields, in particular, spatial and temporal differentiators. The receptive fields emerge if a Hebbian learning rule is applied to the network. Similar to a Linsker model the network has a spatially convergent, linear feedforward structure. Additionally, however, delays omnipresent in the brain are incorporated in the model. The emerging spatiotemporal receptive fields are derived explicitly by extending the approach of MacKay and Miller. The response characteristic of the network is calculated in frequency space and shows that the network can be considered as a spacetime filter for motion in one direction. The emergence of different types of receptive field requires certain structural constraints regarding the spatial and temporal arborisation. These requirements can be derived from the theoretical analysis and might be compared with neuroanatomical data. In this way an explicit link between structure and function of the network is established.     

Development of spatiotemporal receptive fields of simple cells: I. Model formulation

A model for the development of spatiotemporal receptive fields of simple cells in the visual cortex is proposed. The model is based on the 1990 hypothesis of Saul and Humphrey that the convergence of four types of input onto a cortical cell, viz. non-lagged ON and OFF inputs and lagged ON and OFF inputs, underlies the spatial and temporal structure of the receptive fields. It therefore explains both orientation and direction selectivity of simple cells. The response properties of the four types of input are described by the product of linear spatial and temporal response functions. Extending the 1994 model of one of the authors (K.D. Miller), we describe the development of spatiotemporal receptive fields as a Hebbian learning process taking into account not only spatial but also temporal correlations between the different inputs. We derive the correlation functions that drive the development both for the period before and after eye-opening and demonstrate how the joint development of orientation and direction selectivity can be understood in the framework of correlation-based learning. Our investigation is split into two parts that are presented in two papers. In the first, the model for the response properties and for the development of direction-selective receptive fields is presented. In the second paper we present simulation results that are compared with experimental data, and also provide a first analysis of our model. Received: 18 June 1997 / Accepted: 16 September 1997     

Development of spatiotemporal receptive fields of simple cells: II. Simulation and analysis

In part I of this article a correlation based model for the developmental process of spatiotemporal receptive fields has been introduced. In this model the development is described as an activity-dependent competition between four types of input from the lateral geniculate nucleus onto a cortical cell, viz. non-lagged ON and OFF and lagged ON and OFF inputs. In the present paper simulation results and a first analysis are presented for this model. We study the developmental process both before and after eye-opening and compare the results with experimental data from reverse correlation measurements. The outcome of the developmental process is determined mainly by the spatial and the temporal correlations between the different inputs. In particular, if the mean correlation between non-lagged and lagged inputs is weak, receptive fields with a widely varying degree of direction selectivity emerge. However, spatiotemporal receptive fields may show rotation of their preferred orientation as a function of response delay. Even if the mean correlation between two types of temporal input is not weak, direction-selective receptive fields may emerge because of an intracortical interaction between different cortical maps. In an environment of moving lines or gratings, direction-selective receptive fields develop only if the distribution of the directions of motion presented during development shows some anisotropy. In this case, a continuous map of preferred direction is also shown to develop. Received: 18 June 1997 / Accepted: 16 September 1997     

Oscillations in plant membrane transport: model predictions, experimental validation, and physiological implications

Although oscillations in membrane-transport activity are ubiquitous in plants, the ionic mechanisms of ultradian oscillations in plant cells remain largely unknown, despite much phenomenological data. The physiological role of such oscillations is also the subject of much speculation. Over the last decade, much experimental evidence showing oscillations in net ion fluxes across the plasma membrane of plant cells has been accumulated using the non-invasive MIFE technique. In this study, a recently proposed feedback-controlled oscillatory model was used. The model adequately describes the observed ion flux oscillations within the minute range of periods and predicts: (i) strong dependence of the period of oscillations on the rate constants for the H+ pump; (ii) a substantial phase shift between oscillations in net H+ and K+ fluxes; (iii) cessation of oscillations when H+ pump activity is suppressed; (iv) the existence of some 'window' of external temperatures and ionic concentrations, where non-damped oscillations are observed: outside this range, even small changes in external parameters lead to progressive damping and aperiodic behaviour; (v) frequency encoding of environmental information by oscillatory patterns; and (vi) strong dependence of oscillatory characteristics on cell size. All these predictions were successfully confirmed by direct experimental observations, when net ion fluxes were measured from root and leaf tissues of various plant species, or from single cells. Because oscillatory behaviour is inherent in feedback control systems having phase shifts, it is argued from this model that suitable conditions will allow oscillations in any cell or tissue. The possible physiological role of such oscillations is discussed in the context of plant adaptive responses to salinity, temperature, osmotic, hypoxia, and pH stresses.     

Specific detection of the Old World screwworm fly,Chrysomya bezziana,in bulk fly trap catches using real‐time PCR

The Old World screwworm fly (OWS), Chrysomya bezziana Villeneuve (Diptera: Calliphoridae), is a myiasis‐causing blowfly of major concern for both animals and humans. Surveillance traps are used in several countries for early detection of incursions and to monitor control strategies. Examination of surveillance trap catches is time‐consuming and is complicated by the presence of morphologically similar flies that are difficult to differentiate from Ch. bezziana, especially when the condition of specimens is poor. A molecular‐based method to confirm or refute the presence of Ch. bezziana in trap catches would greatly simplify monitoring programmes. A species‐specific real‐time polymerase chain reaction (PCR) assay was designed to target the ribosomal DNA internal transcribed spacer 1 (rDNA ITS1) of Ch. bezziana. The assay uses both species‐specific primers and an OWS‐specific Taqman^® MGB probe. Specificity was confirmed against morphologically similar and related Chrysomya and Cochliomyia species. An optimal extraction protocol was developed to process trap catches of up to 1000 flies and the assay is sensitive enough to detect one Ch. bezziana in a sample of 1000 non‐target species. Blind testing of 29 trap catches from Australia and Malaysia detected Ch. bezziana with 100% accuracy. The probability of detecting OWS in a trap catch of 50 000 flies when the OWS population prevalence is low (one in 1000 flies) is 63.6% for one extraction. For three extractions (3000 flies), the probability of detection increases to 95.5%. The real‐time PCR assay, used in conjunction with morphology, will greatly increase screening capabilities in surveillance areas where OWS prevalence is low.     

Analysis and interpretation of quadratic models of receptive fields

In this protocol, we present a procedure to analyze and visualize models of neuronal input-output functions that have a quadratic, a linear and a constant term, to determine their overall behavior. The suggested interpretations are close to those given by physiological studies of neurons, making the proposed methods particularly suitable for the analysis of receptive fields resulting from physiological measurements or model simulations.     

Electrostatic binding of proteins to membranes. Theoretical predictions and experimental results with charybdotoxin and phospholipid vesicles.

We previously applied the Poisson-Boltzmann equation to atomic models of phospholipid bilayers and basic peptides to calculate their electrostatic interactions from first principles (Ben-Tal, N., B. Honig, R. M. Peitzsch, G. Denisov, and S. McLaughlan. 1996. Binding of small basic peptides to membranes containing acidic lipids. Theoretical models and experimental results. Biophys. J. 71:561-575). Specifically, we calculated the molar partition coefficient, K (the reciprocal of the lipid concentration at which 1/2 the peptide is bound), of simple basic peptides (e.g., pentalysine) with phospholipid vesicles. The theoretical predictions agreed well with experimental measurements of the binding, but the agreement could have been fortuitous because the structure(s) of these flexible peptides is not known. Here we use the same theoretical approach to calculate the membrane binding of two small proteins of known structure: charybdotoxin (CTx) and iberiotoxin (IbTx); we also measure the binding of these proteins to phospholipid vesicles. The theoretical model describes accurately the dependence of K on the ionic strength and mol % acidic lipid in the membrane for both CTx (net charge +4) and IbTx (net charge +2). For example, the theory correctly predicts that the value of K for the binding of CTx to a membrane containing 33% acidic lipid should decrease by a factor of 10(5) when the salt concentration increases from 10 to 200 mM. We discuss the limitations of the theoretical approach and also consider a simple extension of the theory that incorporates nonpolar interactions.     

Activity-dependent matching of excitatory and inhibitory inputs during refinement of visual receptive fields

The receptive field (RF) of single visual neurons undergoes progressive refinement during development. It remains largely unknown how the excitatory and inhibitory inputs on single developing neurons are refined in a coordinated manner to allow the formation of functionally correct circuits. Using whole-cell voltage-clamp recording from Xenopus tectal neurons, we found that RFs determined by excitatory and inhibitory inputs in more mature tectal neurons are spatially matched, with each spot stimulus evoking balanced synaptic excitation and inhibition. This emerges during development through a gradual reduction in the RF size and a transition from disparate to matched topography of excitatory and inhibitory inputs to the tectal neurons. Altering normal spiking activity of tectal neurons by either blocking or elevating GABA(A) receptor activity significantly impeded the developmental reduction and topographic matching of RFs. Thus, appropriate inhibitory activity is essential for the coordinated refinement of excitatory and inhibitory connections.     

The potential for sexual selection in males: Effect of sex ratio and spatiotemporal distribution of receptive females

Summary An animal mating system characterized by male-male competition and active searching for sexually receptive females was modelled to study how varying sex ratio and spatiotemporal distribution of receptive females can affect the variance in male mating success (i. e. potential for sexual selection) in males. The temporal distribution of female receptivity periods appeared to be the variable that had the most pronounced effect on the potential for sexual selection in males. The potential for sexual selection increased monotonically as the degree of asynchrony of female reproduction increased. Female spatial distribution and sex ratio were important only when female reproduction was asynchronous. Then, the potential for sexual selection in males was at its peak when females were overdispersed in space and the sex ratio was female biased. Some of the results derived from the model analysis contrast with predictions from previous studies. The deviating results are most likely caused by different assumptions about modes of mate acquisition in males.     

Spatiotemporal organization of simple-cell receptive fields in area 18 of cat''''s cortex

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Contextual modulation of V1 receptive fields depends on their spatial symmetry

The apparent receptive field characteristics of sensory neurons depend on the statistics of the stimulus ensemble—a nonlinear phenomenon often called contextual modulation. Since visual cortical receptive fields determined from simple stimuli typically do not predict responses to complex stimuli, understanding contextual modulation is crucial to understanding responses to natural scenes. To analyze contextual modulation, we examined how apparent receptive fields differ for two stimulus ensembles that are matched in first- and second-order statistics, but differ in their feature content: one ensemble is enriched in elongated contours. To identify systematic trends across the neural population, we used a multidimensional scaling method, the Procrustes transformation. We found that contextual modulation of receptive field components increases with their spatial extent. More surprisingly, we also found that odd-symmetric components change systematically, but even-symmetric components do not. This symmetry dependence suggests that contextual modulation is driven by oriented On/Off dyads, i.e., modulation of the strength of intracortically-generated signals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Extents of visual, auditory and bimodal receptive fields of single neurons in the feline visual associative cortex

Extracellular microelectrode recordings were carried out on 150 neurons in the anterior ectosylvian sulcal region of halothane-anesthetized, immobilized, artificially ventilated cats. Fifty-nine neurons were visual, 60 were auditory and 31 were bimodal visual-auditory. As the extent of the receptive fields has never been exactly determined, we introduced a quasi-objective, computer-based, statistical method in order to estimate the receptive field sizes in the anterior half of the perimeter. The visual, auditory and bimodal cells had very large receptive fields, often with portions extending well into the ipsilateral hemifield. The mean extents of the visual and auditory receptive fields in the horizontal plane were 75.75 degrees (N=59, SD: +/- 28.620, range: 15-135 degrees), and 132.5 degrees (N=60, SD: +/- 46.72 degrees, range: 15-165 degrees) respectively. These data suggest that a single visual neuron can carry information from the whole visual field of the right eye and a single auditory unit can carry information of azimuths throughout the whole area of the horizontal plane studied. The mean extent of the bimodal receptive fields in the horizontal plane was 82.1 degrees (N=31, SD: +/- 24.24 degrees, range: 30-135 degrees). In 21 of the 31 bimodal cells we observed a facilitatory interaction between visual and auditory stimuli. The mean extent of the facilitatory interactions in these cells was 75.75 degrees (N=21, SD: +/- 24.56 degrees, range: 45-135 degrees).     

Direct extraction of curvature-based metric shape from stereo by view-modulated receptive fields

Any computation of metric surface structure from horizontal disparities depends on the viewing geometry, and analysing this dependence allows us to narrow down the choice of viable schemes. For example, all depth-based or slant-based schemes (i.e. nearly all existing models) are found to be unrealistically sensitive to natural errors in vergence. Curvature-based schemes avoid these problems and require only moderate, more robust view-dependent corrections to yield local object shape, without any depth coding. This fits the fact that humans are strikingly insensitive to global depth but accurate in discriminating surface curvature. The latter also excludes coding only affine structure. In view of new adaptation results, our goal becomes to directly extract retinotopic fields of metric surface curvatures (i.e. avoiding intermediate disparity curvature).To find a robust neural realisation, we combine new exact analysis with basic neural and psychophysical constraints. Systematic, step-by-step ‘design’ leads to neural operators which employ a novel family of ‘dynamic’ receptive fields (RFs), tuned to specific (bi-)local disparity structure. The required RF family is dictated by the non-Euclidean geometry that we identify as inherent in cyclopean vision. The dynamic RF-subfield patterns are controlled via gain modulation by binocular vergence and version, and parameterised by a cell-specific tuning to slant. Our full characterisation of the neural operators invites a range of new neurophysiological tests. Regarding shape perception, the model inverts widely accepted interpretations: It predicts the various types of errors that have often been mistaken for evidence against metric shape extraction.     

Competition of polysaccharides with sugars for the pyranose and the furanose sites in the labellar sugar receptor cell of the blowfly, Phormia regina

In the labellar sugar receptor cell of the blowfly, Phormia regina, soluble starch and dextran T500 inhibited the response to sucrose, to maltose or to glucose, but did not inhibit that to fructose. On the other hand, inulin inhibited the response to fructose, but did not inhibit that to sucrose. These results suggest that both soluble starch and dextran T500 compete with sucrose, with maltose or with glucose for the pyranose site (P site), and that inulin competes with fructose for the furanose site (F site) in a single sugar receptor cell. Each inhibition constant (K_i) was estimated to be 0.6–0.7% for soluble starch. about 4.5% for dextran T500, and about 1.3% for inulin.     

Mathematical principles in afferent visual neurons: Differentiation,integration and transient proportionality related to receptive fields and shift-effect

The dual reciprocal and antagonistic organization of B- and D-neurons of the afferent visual system is obtained using differentiation and integration as mathematical equivalents of visual information processing by an impulse frequency code. The spatial and temporal derivatives lead to the transient responses. A constant and a time-dependent term proportional to the luminance distribution describe the sustained response components and the shift-effect of retinal on- and off-center ganglion cells. Receptive field properties of lateral geniculate cells and their antagonistic shift-effect are obtained by passing the retinal output, i.e. the difference between B- and D-neurons' activity, once again through the same operations. However, the factor of proportionality is applied to the retina alone. The surprisingly small difference between retinal and geniculate receptive field properties on the one hand and the dramatic change from a synergistic to an antagonistic shift-effect on the other hand are thereby explained. The theory offers an understanding of a a possible functional significance of the shift-effect as a mechanism of transientrestoration of visual information, which prevents the system from total fading by means of shifts of the retinal image, normally produced by eye movements.     

Comparison of LFP-based and spike-based spectro-temporal receptive fields and cross-correlation in cat primary auditory cortex

Multi-electrode array recordings of spike and local field potential (LFP) activity were made from primary auditory cortex of 12 normal hearing, ketamine-anesthetized cats. We evaluated 259 spectro-temporal receptive fields (STRFs) and 492 frequency-tuning curves (FTCs) based on LFPs and spikes simultaneously recorded on the same electrode. We compared their characteristic frequency (CF) gradients and their cross-correlation distances. The CF gradient for spike-based FTCs was about twice that for 2-40 Hz-filtered LFP-based FTCs, indicating greatly reduced frequency selectivity for LFPs. We also present comparisons for LFPs band-pass filtered between 4-8 Hz, 8-16 Hz and 16-40 Hz, with spike-based STRFs, on the basis of their marginal frequency distributions. We find on average a significantly larger correlation between the spike based marginal frequency distributions and those based on the 16-40 Hz filtered LFP, compared to those based on the 4-8 Hz, 8-16 Hz and 2-40 Hz filtered LFP. This suggests greater frequency specificity for the 16-40 Hz LFPs compared to those of lower frequency content. For spontaneous LFP and spike activity we evaluated 1373 pair correlations for pairs with >200 spikes in 900 s per electrode. Peak correlation-coefficient space constants were similar for the 2-40 Hz filtered LFP (5.5 mm) and the 16-40 Hz LFP (7.4 mm), whereas for spike-pair correlations it was about half that, at 3.2 mm. Comparing spike-pairs with 2-40 Hz (and 16-40 Hz) LFP-pair correlations showed that about 16% (9%) of the variance in the spike-pair correlations could be explained from LFP-pair correlations recorded on the same electrodes within the same electrode array. This larger correlation distance combined with the reduced CF gradient and much broader frequency selectivity suggests that LFPs are not a substitute for spike activity in primary auditory cortex.