Normative Theory of Visual Receptive Fields

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1 Normative Theory of Visual Receptive Fields Tony Lindeberg Computational Brain Science Lab, Department of Computational Science and Technology, KTH Royal Institute of Technology, SE Stockholm, Sweden. arxiv: v [q-bio.nc] 23 Jan 207 Abstract This article gives an overview of a normative computational theory of visual receptive fields, by which idealized shapes of early spatial, spatio-chromatic and spatiotemporal receptive fields can be derived in an axiomatic way based on structural properties of the environment in combination with assumptions about the internal structure of a vision system to guarantee consistent handling of image representations over multiple spatial and temporal scales. Interestingly, this theory leads to predictions about visual receptive field shapes with qualitatively very good similarity to biological receptive fields measured in the retina, the LGN and the primary visual cortex (V) of mammals. I. INTRODUCTION When light reaches a visual sensor such as the retina, the information necessary to infer properties about the surrounding world is not contained in the measurement of image intensity at a single point, but from the relations between intensity values at different points. A main reason for this is that the incoming light constitutes an indirect source of information depending on the interaction between geometric and material properties of objects in the surrounding world and on external illumination sources. Another fundamental reason why cues to the surrounding world need to be collected over regions in the visual field as opposed to at single image points is that the measurement process by itself requires the accumulation of energy over non-infinitesimal support regions over space and time. Such a region in the visual field for which a visual sensor or and a visual operator responds to visual input or a visual cell responds to visual stimuli is naturally referred to as a receptive field (Hubel and Wiesel [], [2]) (see figure ). If one considers the theoretical and algorithmic problem of designing a vision system that is going to make use of incoming reflected light to infer properties of the surrounding world, one may ask what types of image operations should be performed on the image data. Would any type of image operation be reasonable? Specifically regarding the notion of receptive fields one may ask what types of receptive field profiles would be reasonable? Is it possible to derive a theoretical model of how receptive fields ought to respond to visual data? Initially, such a problem might be regarded as intractable unless the question can be further specified. It is, however, possible to study this problem systematically using approaches that have been developed in the area of computer vision known as scale-space theory [3], [4], [5], [6], [7], [8], [9]. A paradigm that has been developed in this field is to impose structural constraints on the first stages of visual processing that reflect symmetry x_2 Fig.. A receptive field is a region in the visual field for which a visual sensor/neuron/operator responds to visual stimuli. This figure shows a set of partially overlapping receptive fields over the spatial domain with all the receptive fields having the same spatial extent. More generally, one can conceive distributions of receptive fields over space or space-time with the receptive fields of different size, different shape and orientation in space as well as different directions in space-time, where adjacent receptive fields may also have significantly larger relative overlap than shown in this schematic illustration. properties of the environment. Interestingly, it turns out to be possible to substantially reduce the class of permissible image operations from such arguments. The subject of this article is to describe how structural requirements on the first stages of visual processing as formulated in scale-space theory can be used for deriving idealized models of receptive fields and implications of how these theoretical results can be used when modelling biological vision. A main theoretical argument is that idealized models for linear receptive fields can be derived by necessity given a small set of symmetry requirements that reflect properties of the world that one may naturally require an idealized vision system to be adapted to. In this respect, the treatment bears similarities to approaches in theoretical physics, where symmetry properties are often used as main arguments in the formulation of physical theories of the world. The treatment that will follow will be general in the sense that spatial, spatio-chromatic and spatio-temporal receptive fields are encompassed by the same unified theory. This paper is a condensed summary of a more general theoretical framework for receptive fields derived and presented in [9], [0], [], [2] and in turn developed to enable a consistent handling of receptive field responses in terms of provable covariance or invariance properties under natural image transformations (see figure 2). x_ II. STRUCTURAL REQUIREMENTS In the following, we shall describe a set of structural requirements that can be stated concerning (i) spatial

2 external illumination c) Scale covariance under spatial scaling transformations: Under a uniform scaling transformation of the spatial domain x = S s x, we assume that spatial scale covariance should hold viewing distance viewing direction relative motion spatial sampling temporal sampling position in 3-D orientation in 3-D motion in 3-D L (x ; s ) = L(x; s) T Ss(s) S s f = S s T s f (5) for some transformation s = S s (s) of the scale parameter s. d) Affine covariance under affine transformations: Under an affine transformation of the spatial domain x = Ax, we assume that affine covariance should hold L (x ; s ) = L(x; s) T A(s) A f = A T s f (6) Fig. 2. Basic factors that influence the formation of images for a two-dimensional camera that observes objects in the three-dimensional world. In addition to the position, orientation and motion of the object in 3-D, the perspective projection onto the image plane is affected by the viewing distance, viewing direction and relative motion of the camera in relation to the object, the spatial and temporal sampling characteristics of the image sensor as well the usually unknown external illumination field in relation to the geometry of the scene and the camera. geometry, (ii) spatio-temporal geometry, (iii) the image measurement process with its close relationship to the notion of scale, (iv) internal representations of image data that are to be computed by a general purpose vision system and (v) the parameterization of image intensity with regard to the influence of illumination variations. A. Static image data over a spatial domain For time-independent data f over an N-dimensional spatial image domain, we would like to define a family of image representations L( ; s) = T s f () over a possibly multi-dimensional scale parameter s for some family of image operators T s that satisfies: a) Linearity and shift-invariance: To make as few irreversible decisions as possible at the earliest stages, we assume linearity T s (a f + a 2 f 2 ) = a T s f + b T s f 2. (2) Specifically, this property implies that scale-space properties that we derive for the zero-order image representation L will transfer to any spatial derivative of L. To ensure that the visual interpretation of an object should be the same irrespective of its position in the image plane, we assume shift invariance T s (S x f) = S x (T s f). (3) Together, the assumptions about linearity and shiftinvariance imply that T s will be a convolution operator. b) Semi-group structure over spatial scales: To ensure that transformations between any pairs of spatial scale levels are of the same form (algebraic closedness), we assume a semi-group structure over spatial scales T s T s2 = T s+s 2. (4) for some transformation s = A(s) of the scale parameter. e) Non-creation of new structure with increasing scale: To formalize the requirement that new structures should not be created from finer to coarser scales, we assume non-enhancement of local extrema: If at some scale s 0 a point x 0 is a local maximum (minimum) for the mapping x L(x; s 0 ), then ( s L)(x; s) 0 at any spatial maximum, ( s L)(x; s) 0 at any spatial minimum. This condition implies a strong condition on the class of possible smoothing kernels T ( ; s). B. Time-dependent image data over space-time For a time-dependent spatial domain, we do first inherit the structural properties regarding a spatial domain and complement the spatial scale parameter s by a temporal scale parameter τ. In addition, we assume: f) Scale covariance under temporal scaling transformations: Under a scaling transformation of the temporal domain t = S τ t, we assume that temporal scale covariance should hold L (x, t ; s, τ ) = L(x, t; s, τ) T S(s,τ) (τ) S τ f = S τ T s,τ f (7) for some transformation S τ (τ) of the temporal scale parameter τ. g) Galilean covariance under Galilean transformations: To guarantee a consistent visual interpretation under different relative motions between the object and the observer as modelled by Galilean transformations f = G f f (x, t ) = f(x, t) with x = x+v t, (8) we assume that L (x, t ; s, τ ) = L(x, t; s, τ ) T Gv(s,τ) G v f = G v T s,τ f (9) should hold for some of transformation G v (s, τ) of the spatio-temporal scale parameters (s, τ). h) Semi-group structure over temporal scales in the case of a non-causal temporal domain: To ensure that transformations between any pairs of spatial scale levels are of the same form (algebraic closedness), we assume a semi-group structure over temporal scales T τ T τ2 = T τ+τ 2. (0)

3 i) Cascade structure over temporal scales in the case of a time-causal temporal domain: Since it can be shown that the assumption of a semi-group structure over temporal scales leads to undesirable temporal dynamics in terms of e.g. longer temporal delays for a time-causal temporal domain [3, Appendix A], we do for a timecausal temporal domain instead assume a weaker cascade smoothing property over temporal scales for the temporal smoothing kernel over temporal scales L( ; τ 2 ) = h( ; τ τ 2 ) L( ; τ ), () where the temporal kernels h(t; τ) should for any triplets of temporal scale values and temporal delays τ, τ 2 and τ 3 obey the transitive property h( ; τ τ 2 ) h( ; τ 2 τ 3 ) = h( ; τ τ 3 ). (2) j) Non-enhancement of local space-time extrema in the case of a non-causal temporal domain: In the case of a non-causal temporal domain, to formalize the requirement that new structures should not be created from finer to coarser spatio-temporal scales, we assume nonenhancement of local extrema: If at some scale (s 0, τ 0 ) a point (x 0, t 0 ) is a local maximum (minimum) for the mapping (x, t) L(x, t; s 0, τ 0 ), then α ( s L)(x, t; s, τ) + β ( τ L)(x, t; s, τ) 0 at any spatio-temporal maximum α ( s L)(x, t; s, τ) + β ( τ L)(x, t; s, τ) 0 at any spatio-temporal minimum should hold in any positive spatio-temporal direction define from any non-negative linear combinations of α and β. This condition implies a strong condition on the class of possible smoothing kernels T (, ; s, τ). k) Non-creation of new local extrema or zerocrossings for a purely temporal signal in the case of a non-causal temporal domain: In the case of a timecausal temporal domain, we do instead require that for a purely temporal signal f(t), the transformation from a finer temporal scale τ to a coarser temporal scale τ 2 must not increase the number of local extrema or zero-crossings in the signal. III. IDEALIZED RECEPTIVE FIELD FAMILIES A. Spatial image domain Based on the above assumptions in Section II-A, it can be shown [9] that when complemented with certain regularity assumptions in terms of Sobolev norms, they imply that spatial scale-space representation L as determined by these must satisfy a diffusion equation of the form s L = 2 T (Σ 0 L) δ T 0 L (3) for some positive semi-definite covariance matrix Σ 0 and some translation vector δ 0. In terms of convolution kernels, this corresponds to Gaussian kernels of the form g(x; Σ s, δ s ) = (2π) N/2 e (x δs) T Σ s (x δs)/2, det Σ s (4) which for a given Σ s = sσ 0 and a given δ s = s δ 0 satisfy (3). If we additionally require these kernels to be mirror symmetric through the origin, then we obtain affine Gaussian kernels g(x; Σ) = T (2π) N/2 det Σ e x Σ x/2. (5) Their spatial derivatives constitute a canonical family for expressing receptive fields over a spatial domain. Specifically, for a two-dimensional image domain this leads to spatial receptive fields of the form T (x, x 2 ; s, Σ) = ϕ m m2 ϕ (g(x, x 2 ; s, Σ)), (6) where ϕ and ϕ constitute directional derivative operators in two orthogonal directions, preferably aligned to the eigendirections of the affine covariance matrix Σ. B. Spatio-temporal image domain Over a non-causal spatio-temporal domain, corresponding arguments as in Section III-A lead to a similar form of diffusion equation as in Equation (3), while expressed over the joint space-time domain p = (x, t) and with δ 0 interpreted as a local drift velocity. After splitting the composed affine Gaussian spatio-temporal smoothing kernel corresponding to (4) while expressed over the joint space-time domain into separate smoothing operations over space and time, and combining that result with the results from corresponding theoretical analysis for a time-causal spatio-temporal domain in [9], [2], the resulting spatio-temporal derivative kernels constituting the spatio-temporal extension of the spatial receptive field model (6) can be reparameterised and summarized on the following form (see [9], [0], [], [2]): T (x, x 2, t; s, τ; v, Σ) where = ϕ m m2 ϕ n t (g(x v t, x 2 v 2 t; s, Σ) h(t; τ)) (7) x = (x, x 2 ) denote the spatial coordinates, t denotes time, s denotes the spatial scale, τ denotes the temporal scale, v = (v, v 2 ) T denotes a local image velocity, Σ denotes a spatial covariance matrix determining the shape of a spatial affine Gaussian kernel, m and m 2 denote orders of spatial differentiation, n denotes the order of temporal differentiation, ϕ = cos ϕ x + sin ϕ x2, ϕ = sin ϕ x cos ϕ x2 denote spatial directional derivative operators in two orthogonal directions ϕ and ϕ, t = v x + v 2 y + t is velocity-adapted temporal derivative operator aligned to the direction of the local image velocity v = (v, v 2 ) T, g(x; s, Σ) = 2πs Σ x/2s det Σ e xt is an affine Gaussian kernel with its size determined by the spatial scale parameter s and its shape by the spatial covariance matrix Σ, g(x v t, x 2 v 2 t; s, Σ) denotes a spatial affine Gaussian kernel that moves with image velocity v = (v, v 2 ) in space-time and

4 h(t; τ ) is a temporal smoothing kernel over time corresponding to a Gaussian kernel in the case of non-causal time or a cascade of first-order integrators or equivalently truncated exponential kernels coupled in cascade over a time-causal temporal domain. This family of spatio-temporal scale-space kernels can be seen as a canonical family for linear receptive fields over a spatio-temporal domain. For the case of a time-causal temporal domain, the result states that truncated exponential kernels of the form t/µ k t 0, µk e (8) hexp (t; µk ) = 0 t < 0. coupled in cascade constitute the natural temporal smoothing kernels and leading to a composed convolution kernel of the form hcomposed ( ; µ) = K k= hexp ( ; µk ). T (x, t; s, τ ) Tx (x, t; s, τ ) Txx (x, t; s, τ ) Tt (x, t; s, τ, δ) Txt (x, t; s, τ ) Ttt (x, t; s, τ ) (9) Two natural ways of distributing the discrete time constants µk over temporal scales are studied in detail in [2], [3] corresponding to either a uniform or a logarithmic distribution in terms of the composed variance τk = K X µ2k. (20) k= Specifically, it is shown in [2] that in the case of a logarithmic distribution of the discrete temporal scale levels, it is possible to consider an infinite number of temporal scale levels that cluster infinitely dense near zero temporal scale τ0 τ0 τ0 (2)... 6, 4, 2, τ0, c2 τ0, c4 τ0, c6 τ0,... c c c so that a scale-invariant limit kernel Ψ(t; τ, c) can be defined obeying self-similarity and scale covariance over temporal scales and with a Fourier transform of the form Y Ψ (ω; τ, c) =. k + ic c2 τ ω k= Fig. 3. Space-time separable kernels Txm tn (x, t; s, τ ) = xm tn (g(x; s) h(t; τ )) up to order two obtained as the composition of Gaussian kernels over the spatial domain x and a cascade of truncated exponential kernels over the temporal domain t with a logarithmic distribution of the intermediate temporal scale levels (s =, τ =, K = 7, c = 2). (Horizontal axis: space x. Vertical axis: time t.) T (x, t; s, τ, v) Tx (x, t; s, τ, v) Tt (x, t; s, τ, δ) (22) Figure 3 and Figure 4 show spatio-temporal kernels over a +-dimensional spatio-temporal domain using approsimations of the time-causal limit kernel for temporal smoothing over the temporal domain and the Gaussian kernel for spatial smoothing over the spatial domain. Txx (x, t; s, τ, v) Txt (x, t; s, τ, v) Ttt (x, t; s, τ, v) IV. C OMPUTATIONAL MODELLING OF BIOLOGICAL RECEPTIVE FIELDS In two comprehensive reviews, DeAngelis et al. [4], [5] present overviews of spatial and temporal response properties of (classical) receptive fields in the central visual pathways. Specifically, the authors point out the limitations of defining receptive fields in the spatial domain only and emphasize the need to characterize receptive fields in the joint space-time domain, to describe how a neuron processes the visual image. Conway and Livingstone [6] and Johnson et al. [7] show results of corresponding investigations concerning spatio-chromatic receptive fields. Fig. 4. Velocity-adapted spatio-temporal kernels Txm tn (x, t; s, τ, v) = xm tn (g(x vt; s) h(t; τ )) up to order two obtained as the composition of Gaussian kernels over the spatial domain x and a cascade of truncated exponential kernels over the temporal domain t with a logarithmic distribution of the intermediate temporal scale levels (s =, τ =, K = 7, c = 2, v = 0.5). (Horizontal axis: space x. Vertical axis: time t.)

5 In the following, we will describe how the above mentioned spatial and spatio-temporal scale-space concepts can be used for modelling the spatial, spatio-chromatic and spatio-temporal response properties of biological receptive fields. Indeed, it will be shown that the Gaussian and time-causal scale-space concepts lead to predictions of receptive field profiles that are qualitatively very similar to all the receptive field types presented in (DeAngelis et al. [4], [5]) and schematic simplifications of most of the receptive fields shown in (Conway and Livingstone [6]) and (Johnson et al. [7]). hxxt (x, t; s, τ ) hxxtt (x, t; s, τ ) A. Receptive fields in the LGN Regarding visual receptive fields in the lateral geniculate nucleus (LGN), DeAngelis et al. [4], [5] report that most neurons (i) have approximately circular centersurround organization in the spatial domain and that (ii) most of the receptive fields are separable in spacetime. There are two main classes of temporal responses for such cells: (i) a non-lagged cell is defined as a cell for which the first temporal lobe is the largest one (figure 5(left)), whereas (ii) a lagged cell is defined as a cell for which the second lobe dominates (figure 5(right)). When using a time-causal temporal smoothing kernel, the first peak of a first-order temporal derivative will be strongest, whereas second peak of a second-order temporal derivative will be strongest (see [2, Figure 2]). Thus, according to this theory, non-lagged LGN cells can be seen as corresponding to first-order temporal derivatives, whereas lagged LGN cells can be seen as corresponding to second-order temporal derivatives. The spatial response, on the other hand, shows a high similarity to a Laplacian of a Gaussian, leading to an idealized receptive field model of the form (Lindeberg [0, equation (08)]) hlgn (x, y, t; s, τ ) = ±( xx + yy ) g(x, y; s) tn h(t; τ ). (23) Figure 5 shows results of modelling separable receptive fields in the LGN in this way, using a cascade of truncated exponential kernels of the form (9). B. Receptive fields in V Concerning the neurons in the primary visual cortex (V), DeAngelis et al. [4], [5] describe that their receptive fields are generally different from the receptive fields in the LGN in the sense that they are (i) oriented in the spatial domain and (ii) sensitive to specific stimulus velocities. Cells (iii) for which there are precisely localized on and off subregions with (iv) spatial summation within each subregion, (v) spatial antagonism between on- and off-subregions and (vi) whose visual responses to stationary or moving spots can be predicted from the spatial subregions are referred to as simple cells (Hubel and Wiesel [], [2]). Figure 6 shows an example of the spatial dependency of a simple cell that can be well modelled by a firstorder affine Gaussian derivative over image intensities. Figure 7 shows corresponding results for a color-opponent receptive field of a simple cell in V that can be modelled Fig. 5. Computational modelling of space-time separable receptive field profiles in the lateral geniculate nucleus (LGN) as reported by DeAngelis et al. [4] using idealized spatio-temporal receptive fields of the form T (x, t; s, τ ) = xα tβ (g(; s) h(t; τ )) according to (7) and with the temporal smoothing function h(t; τ ) modelled as a cascade of first-order integrators/truncated exponential kernels of the form (9). (left) a non-lagged cell modelled using first-order temporal derivatives (right) a lagged cell modelled using second-order temporal derivatives. Parameter values: (a) hxxt : σx = 0.5 degrees, σt = 40 ms. (b) hxxtt : σx = 0.6 degrees, σt = 60 ms. (Horizontal dimension: space x. Vertical dimension: time t.) x g(x, y; Σ) Fig. 6. Example of a receptive field profile over the spatial domain in the primary visual cortex (V) as reported by DeAngelis et al. [4], [5]. (middle) Receptive field profile of a simple cell over image intensities as reconstructed from cell recordings, with positive weights represented as green and negative weights by red. (left) Stylized simplification of the receptive field shape. (right) Idealized model of the receptive field from a first-order directional derivative of an affine Gaussian kernel x g(x, y; Σ) = x g(x, y; λx, λy ) according to (5), here with λx = 0.2 and λy = 2 in units of degrees of visual angle, and with positive weights with respect to image intensities represented by white and negative values by violet. ϕ g(x, y; Σ) Fig. 7. Example of a colour-opponent receptive field profile over the spatial domain for a double-opponent simple cell in the primary visual cortex (V) as measured by Johnson et al. [7]. (left) Responses to Lcones corresponding to long wavelength red cones, with positive weights represented by red and negative weights by blue. (middle) Responses to M-cones corresponding to medium wavelength green cones, with positive weights represented by red and negative weights by blue. (right) Idealized model of the receptive field from a first-order directional derivative of an affine Gaussian kernel ϕ g(x, y; Σ) according to (5) over a red-green colour-opponent channel for σ = λ = 0.6, σ2 = λ2 = 0.2 in units of degrees of visual angle, α = 67 degrees and with positive weights for the red-green colour-opponent channel represented by red and negative values by green.

6 h xx(x, t; s, τ, v) h xxx(x, t; s, τ, v) Fig. 8. Computational modelling of simple cells in the primary visual cortex (V) as reported by DeAngelis et al. [4] using idealized spatiotemporal receptive fields of the form T (x, t; s, τ, v) = x α t β (g(x vt; s) h(t; τ)) according to equation (7), specifically in terms of velocity-adapted receptive fields corresponding to second- or third-order derivatives over space and with time-causal temporal smoothing based on truncated exponential kernels coupled in cascade (9). Parameter values: (left) h xx: σ x = 0.7 degrees, σ t = 50 ms, v = degrees/ms. (right) h xxx: σ x = 0.5 degrees, σ t = 80 ms, v = degrees/ms. (Horizontal axis: Space x in degrees of visual angle. Vertical axis: Time t in ms.) as a first-order affine Gaussian spatio-chromatic derivative over an R-G colour-opponent channel. Figure 8 shows the result of modelling the spatiotemporal receptive fields of simple cells in V in this way, using the general idealized model of spatio-temporal receptive fields in equation (7) in combination with a temporal smoothing kernel obtained by coupling a set of truncated exponential kernels in cascade. As can be seen from these figures, the proposed idealized receptive field models do rather well reproduce the qualitative shape of the neurophysiologically recorded biological receptive fields. V. SUMMARY We have described a normative theory for visual receptive fields, by which idealized receptive field shapes can be derived in an axiomatic way from a combination of mathematical and physical assumptions. These receptive field shapes are compatible with natural image transformations including: (i) variations in the sizes of objects in the world, (ii) variations in the viewing distance, (iii) variations in the viewing direction, (iv) relative motion between objects in the world and the observer, (v) the speed by which temporal events occur and (vi) locally multiplicative illumination variations. Invariance to local multiplicative illumination variations is achieved by applying the derivative-based receptive fields on a logarithmic brightness scale f log I [0, Section 2.3]. Additionally, the receptive fields from this theory lead to predictions about receptive field profiles in good agreement with receptive field shapes measured in the retina, the LGN and the primary visual cortex (V). ACKNOWLEDGMENTS The support from the Swedish Research Council (Contract No ) is gratefully acknowledged. REFERENCES [] D. H. Hubel and T. N. Wiesel, Receptive fields of single neurones in the cat s striate cortex, J Physiol, vol. 47, pp , 959. [2], Brain and Visual Perception: The Story of a 25-Year Collaboration. Oxford University Press, [3] T. Iijima, Observation theory of two-dimensional visual patterns, Papers of Technical Group on Automata and Automatic Control, IECE, Japan, Tech. Rep., 962. [4] J. J. Koenderink, The structure of images, Biological Cybernetics, vol. 50, pp , 984. [5] J. J. Koenderink and A. J. van Doorn, Generic neighborhood operators, IEEE Trans. Pattern Analysis and Machine Intell., vol. 4, no. 6, pp , Jun [6] T. Lindeberg, Scale-Space Theory in Computer Vision. Springer, 993. [7] L. M. J. Florack, Image Structure, ser. Series in Mathematical Imaging and Vision. Springer, 997. [8] B. ter Haar Romeny, Front-End Vision and Multi-Scale Image Analysis. Springer, [9] T. Lindeberg, Generalized Gaussian scale-space axiomatics comprising linear scale-space, affine scale-space and spatio-temporal scale-space, Journal of Mathematical Imaging and Vision, vol. 40, no., pp. 36 8, 20. [0], A computational theory of visual receptive fields, Biological Cybernetics, vol. 07, no. 6, pp , 203. [], Invariance of visual operations at the level of receptive fields, PLOS ONE, vol. 8, no. 7, p. e66990, 203. [2], Time-causal and time-recursive spatio-temporal receptive fields, Journal of Mathematical Imaging and Vision, vol. 55, no., pp , 206. [3], Temporal scale selection in time-causal scale space, Journal of Mathematical Imaging and Vision, 207, doi:0.007/s [4] G. C. DeAngelis, I. Ohzawa, and R. D. Freeman, Receptive field dynamics in the central visual pathways, Trends in Neuroscience, vol. 8, no. 0, pp , 995. [5] G. C. DeAngelis and A. Anzai, A modern view of the classical receptive field: Linear and non-linear spatio-temporal processing by V neurons, in The Visual Neurosciences, L. M. Chalupa and J. S. Werner, Eds. MIT Press, 2004, vol., pp [6] B. R. Conway and M. S. Livingstone, Spatial and temporal properties of cone signals in alert macaque primary visual cortex, Journal of Neuroscience, vol. 26, no. 42, pp , [7] E. N. Johnson, M. J. Hawken, and R. Shapley, The orientation selectivity of color-responsive neurons in Macaque V, The Journal of Neuroscience, vol. 28, no. 32, pp , 2008.