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# NEW DEVELOPMENTS OF PHOTODETECTORS FOR THE LAKE BAIKAL NEUTRINO EXPERIMENT

## 1 Introduction

The lake Baikal Neutrino Experiment has history of more than 20 years, starting from small short experiments with a few PMTs in the early 80s to the present large scale longterm operating neutrino telescope NT-200$`^\mathrm{?}`$, which has been put into operation on April 6th 1998. The telescope’s effective area for muons is 2000-10000 depending on a muon energy. The rate of events due to atmospheric neutrinos is about 1 per two days.

## 2 Neutrino telescope NT-200

The lake Baikal Neutrino Telescope NT-200 is located in the southern part of the great Siberian lake Baikal at 3.6km from the shore and at the depth of 1km. The schematic view of NT-200 and optical modules attached at a string are presented in fig.1. The telescope consists of 192 optical modules at 8 vertical strings arranged at an umbrella like frame. Optical modules are grouped in pairs and swithched in coincidence with 15ns time window. So two optical modules in one pair define one optical channel resulting in rather low optical channel background counting rate of 100–300Hz. Two pairs of optical modules form so called svyazka. The detector electronics system is hierarchical: from the lowest level to the highest one - optical module electronics, svyazka electronics module, string and detector electronics modules. In the latter detector trigger signals are formed and all information from string electronics modules are received and sent to the shore station. Three underwater electrical cables connect the detector with the shore station. The detector is operated from the shore station.

## 3 Quasar-370 phototube

The Quasar-370 phototube $`^{\mathrm{?},\mathrm{?},\mathrm{?},\mathrm{?}}`$ is a hybrid phototube which consists of an electro-optical preamplifier with large area hemispherical photocathode and small conventional type PMT, see fig.2. Photoelectrons from a large 37cm diameter hemispherical photocathode with $`2\pi `$ acceptance are accelerated by 25 kV to a fast, high gain luminescent screen ($`YSO`$ scintillator is usually used). The light flashes in luminescent screen induced by photoelectrons are read out by small PMT with 3cm diameter photocathode. The latter has been developed especially for this kind of application by INR and MELZ factory $`^\mathrm{?}`$. As a result one photoelectron from the hemispherical photocathode yields typically 25 photoelectrons in the small PMT.This high gain first stage results in an excellent single photoelectron resolution. Due to the fast acceleration of primary photoelectrons by 25 kV high voltage and mushroom shaped glass envelope the time jitter can be kept rather low. Last but not the least the tube is almost insensitive to the Earth’s magnetic field. Averaged over more than 200 tubes the mean values for single photoelectron resolution and time resolution is 70% and 2ns respectively. Quasar-370 phototube is characterized by conspicuously low level of afterpulses in comparison with conventional PMTs. The level of afterpulses in Quasar-370 tube is substantially less than 1%. Another advantage of Quasar-370 phototube in contrary to conventional PMTs is the lack of prepulses due to the fact that the first stage of the phototube is optically separated from tube’s photocathode. It was shown $`^{\mathrm{?},\mathrm{?},\mathrm{?}}`$ that Quasar-370 parameters depend strongly on characteristics of a scintillator in a luminescent screen. So recently we have developed a number of modifications of Quasar-370 tube with new scintillators. The most promising results we have got with $`ScBO_3`$, $`YAP`$ and $`LSO`$. Unfortunately the latter one has one substantial drawback for using in Quasar-370 tube. It has rather high value of $`Z_{eff}`$. It is important to note here we need scintillators with $`Z_{eff}`$ as low as possible to suppress effectively late pulses $`^\mathrm{?}`$ due to photoelectron backscattering effect. With these new scintillators we have reached $`1`$ns (FWHM) time jitter and $`40`$% single photoelectron resolution. It is noteworthy to mention here about very much intriguing scintillator$`^\mathrm{?}`$ \- $`ZnO:Ga`$ with less than 1ns decay time and light yield of 40% of $`NaI:Tl`$. At present we try to work with this scintillator and to reproduce results of $`^\mathrm{?}`$. The success of this work would undoubtedly be a kind of a breakthrough in designing of very fast and effective hybrid phototubes.

## 4 Two-channel optical module

The pairwise ideology pursued in NT-200 neutrino telescope has many advantages. The ideology allows to suppress effectively an individual optical module background counting rate due to water luminescence and a phototube’s dark current, to eliminate phenomena deteriorating phototube’s time resolution namely prepulses, late pulses and afterpulses. Moreover such an approach facilitates very much designing of trigger system, data acquisition system etc. Unfortunately there is just one but essential shortcoming. Namely it’s too expensive to have two optical modules for one optical channel. To overcome this problem we have developed two-channel optical module based on two-channel version of Quasar-370 phototube$`^\mathrm{?}`$.

Two-channel Quasar-370 phototube uses the same electro-optical preamplifier and new two-anode small PMT with mesh dynode system instead of conventional PMT. This new two-anode small PMT has following characteristics: time jitter – $`\sigma =340`$ps; Peak-to valley ratio of single photoelectron sharge distribution – 1.5; cross-talks between channels – 1%. Basing on two-channel Quasar-370 phototube we have developed two-channel optical module which incorporates besides Quasar-370 phototube two fast transimpedance preamplifiers for each anode signals and uses the same glass pressure vessel and the same penetrators as conventional Baikal optical module. In the course of the last expedition at the lake Baikal we have tested one pilot sample of two-channel optical module in frame of a new experimental string designed to test technological innovations for future neutrino telescopes at the lake. In the test measurements at the lake we used ordinary but slightly modified NT-200 front-end electronics. Output signals of each channel are switched in coincidence just in the same way as in NT-200. It results in the same local trigger counting rate of 100-300Hz. Fig.3 presents the time dependence of muon trigger rate for two conventional Baikal optical channels and new optical channel based on two-channel optical module. One can see that the new optical channel has comparable sensitivity to muons with ordinary optical channels of NT-200. It opens new possibilities for NT-200 further extension plans.

## 5 Other developments

A new version of Quasar-370 phototube (Quasar-370D) with semiconductor diode as a photoelectron multiplying element instead of a system of luminescent screen and small PMT has been developed. We have manufactured two pilot samples which are under studies now. Modifications of Quasar-370 phototubes (Quasar-370G)$`^\mathrm{?}`$ which are able to withstand high current due to night sky background operate successfully in the wide angle atmospheric air Cherenkov detectors TUNKA$`^\mathrm{?}`$ and SMECA$`^\mathrm{?}`$. Quasar-370L is low background version of Quasar-370 phototube. It has $`U^{238}`$ and $`Th^{232}`$ content of about $`10^8`$g/g. Quasar-370L is aimed at using in low background experiments. Another development is concerned with phototubes with high quantum efficiency (more than 40% in the range 450-550nm) photocathodes and it’s results will be reported elsewhere.

## Conclusion

The lake Baikal Neutrino Experiment successful operation proves high performance and high reliability of a number of photodetectors developed for this kind of application. New developments are focused at future neutrino experiments at the lake Baikal and other experiments in high energy physics, cosmic ray and astroparticle physics.

## Acknowledgments

This work was supported by the Russian Ministry of Research (contract 102-11(00)-p), the German Ministry of Education and Research and the Russian Fund of Basic Research (grants 99-02-1837a,01-02-31013 and 00-15-96794) and by the Russian Federal Program ”Integration”(project 346). The author is indebted very much to his colleagues from KATOD and MELZ laboratoties and Baikal Collaboration. The author would like to thank particularly Dr. V.Ch.Lubsandorzhieva for many invaluable advices and help in preparation of this paper.

## References


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# Information Optimization in Coupled Audio–Visual Cortical Maps

## I Introduction

In the struggle of biological organisms to survive and reproduce, processing of information is of central importance. Sensory signals provide valuable information about the external world, such as the locations of predators and preys. Localization of sources is facilitated by topographic maps of neurons to various parts of the brainmaps , reflecting the spatial arrangements of signals around the animal. The barn owl has to rely extensively on sounds to find its prey in the dark, and has consequently developed precise ‘auditory space maps.’

By extensive experiments, Knudsen and collaborators have shown that the optic tectum of the barn owl has both visual and aural maps of space that are in close registryK82 , K83 . The visual signal plays a crucial role in aligning the aural map; experimental manipulations of the owl’s sensory experience reveal the plasticity of these maps in young animals, and the instructive role played by the visual experience. (A recent review, with specific references can be found in Ref. Krev .) The current study was motivated by experiments in which owls are fitted with prismatic spectacles that shift the visual fields by a preset degree in the horizontal directionBK . In young owls, the receptive auditory maps were found to shift to remain in registry with the visual maps, which stayed unchanged.

There is at least one theoretical attempt to explain the registry of neural maps through a ‘value–dependent learning,’ where synaptic connections in a network are enhanced after ‘foveation towards an auditory stimulus’Rucci . In this paper we take a more abstract approach to the coupling of audio–visual maps, and search for neural connections (receptive fields) that maximize the information gained from the sensory signals. In earlier studiesBRZ , BZ , Bialek and one of us formulated an approach to optimization of information in the visual system, and in computations with neural spike trainsspikes .

Here, we extend the methods of Ref. BRZ  for computing receptive fields in the visual system, to finding the optimal connectivities in an audio-visual cortex, such as the owl’s optic tectum. We find that the shape and registry of the aural map is established by the correlations between the audio and visual signals. In response to an artificial shift of the visual field (as with the prismatic spectacles), the visual receptive field is unchanged. While the aural receptive field shifts in the adaptive direction, its shape changes due to the costs of rewiring the neurons.

The general formalism for our calculations is set up in Sec. II.1, which reviews the methodology introduced in Ref. BRZ . The essence of this approach is the assumption that neural connections act as linear filters of the incoming signals, and also introduce noise in the outputs. If the (correlated) input signals, and the random noise, are taken from Gaussian probability distributions, the outputs are also Gaussian distributed. The Shannoninformation  information content of the resulting outputs is easily calculated. The task is to find filter functions that maximize this information, subject to biologically motivated costs, and for given correlations of the input signals. In Ref. BRZ  this approach was used to obtain receptive fields in the visual system. In Sec. II.2, we generalize this formalism to coupled audio-visual signals.

A necessary input to the calculations is the correlations between the audio and visual signals, as discussed in Sec. II.3. Since it is clearly much easier to localize objects by sight that sound, it is reasonable that the information carried by the visual channel should far exceed the aural one. The two sources of information are however quite likely to be correlated, resulting in couplings between the corresponding filters. In the experiments on barn owls, the prismatic glasses shift the visual field and hence modify the correlations between the signals. We examine how such shifts change the filter functions (neural connectivities) that optimize the information content in the outputs.

As argued in Sec. II.4, the disparities in the strengths of visual and aural signals simplify the search for optimal filters. In particular, we find that the visual receptive fields are relatively robust and unchangeable, while the shape of the aural receptive field is the product of two terms: One reflects the correlations between sights and sounds, and shifts along with external displacement of these signals; the second is associated with the costs of making connections to distant neurons. This result is further interpreted in the final section (Sec. III), where some implications for experiments, as well as directions for future extensions and generalizations, are also discussed.

## II Analysis of Information

### II.1 General Formalism

The processing of information by neural connections in the cortex is modelled in Ref. BRZ  as follows: After passing through intermediate stations, sensory signals arrive as a set of inputs $`\{s_J\}`$. Further processing takes place by neurons that sample the information from a subset of these inputs, and produce an appropriate output. For ease of calculation, the outputs are represented as a linear transformation of the inputs, according to

$$O_i=\underset{J}{}F_{iJ}s_J+\eta _i.$$
(1)
The filtering of information is thus parameterized by the matrix $`\{F_{iJ}\}`$, and is also assumed to introduce an unavoidable noise $`\eta _i`$. There are of course many possible sensory inputs, which can be taken from a joint probability distribution $`P_{in}[s_J]`$. Equation (1) is thus a transformation from one set of random variables (the inputs) to another (the outputs); the latter described by the joint probability distribution function $`P_{out}[O_i]`$. The amount of information associated with a given probability distribution is quantifiedinformation  (up to a baseline and units) by $`[P]\mathrm{ln}P`$, where the averages are taken with the corresponding probability. The task of finding optimal filters is thus to come up with the matrix $`F`$ that maximizes $`[P_{out}]`$ for specified input and noise probabilities.

The Shannon information can be calculated easily for Gaussian distributed random variables. Let us consider the set of $`N`$ random variables $`\{x_i\}`$, taken from the probability

$$P\left[x_i\right]=\sqrt{\frac{detA}{(2\pi )^N}}\mathrm{exp}\left[\frac{1}{2}x_iA_{ij}x_j\right],$$
(2)
where summation over the repeated indices is implicit, and $`detA`$ indicates the determinant of the $`N\times N`$ matrix with elements $`A_{ij}`$. It is easy to check that, up to an unimportant additive constant of $`N/2`$,

$$[P]=\frac{1}{2}\mathrm{ln}detA=\frac{1}{2}\mathrm{ln}det\left[x_ix_j\right],$$
(3)
where we have noted that the pairwise averages are related to the inverse matrix by $`x_ix_j=A_{ij}^1`$. A linear filter as in Eq. (1), maps one set of Gaussian variables to new ones. Thus if we assume that the inputs $`\{s_J\}`$, and the (independent) noise $`\{\eta _i\}`$, are Gaussian distributed, we can calculate the information content of the output using Eq. (3), with

$$O_iO_j=F_{iJ}F_{kL}s_Js_L+\eta _i\eta _j.$$
(4)
We are interested in describing cortical maps related to visual or aural localization of objects. These locations vary continuously in space, and are topographically mapped to positions on a two-dimensional cortex. As such, it is convenient to promote the indices $`i`$ and $`J`$, used above to label output and input neurons, to continuous vectors in two dimensional space. For example, following Ref. BRZ , let us consider an image described by a scalar field $`s(\stackrel{}{x})`$ on a $`2`$dimensional surface with coordinates $`\stackrel{}{x}.`$ The image is sampled by an array of cells such that the output of the cell located at $`\stackrel{}{x}`$ is given by

$$O\left(\stackrel{}{x}\right)=d^2yF\left(\stackrel{}{x}\stackrel{}{y}\right)s\left(\stackrel{}{y}\right)+\eta \left(\stackrel{}{x}\right),$$
(5)
where the function $`F(\stackrel{}{r})`$ describes the receptive field of the cell. Assuming uncorrelated neural noise, $`\eta (\stackrel{}{x})\eta (\stackrel{}{x}^{})=N\delta ^2(\stackrel{}{x}\stackrel{}{x}^{})`$, and signal correlations $`s(\stackrel{}{x})s(\stackrel{}{x}^{})=S(\stackrel{}{x}\stackrel{}{x}^{})`$, the filter-dependent part of the output information is given by

$$=\frac{1}{2}\mathrm{ln}det\left[\delta ^2(\stackrel{}{x}\stackrel{}{x}^{})+d^2yd^2y^{}F\left(\stackrel{}{x}\stackrel{}{y}\right)F\left(\stackrel{}{x}^{}\stackrel{}{y}^{}\right)\frac{S\left(\stackrel{}{y}\stackrel{}{y}^{}\right)}{N}\right].$$
(6)
Note that we have assumed that the signal is translationally invariant, such that correlations only depend on the relative distance between their sources. This allows us to change basis to the Fourier components, $`\stackrel{~}{s}(\stackrel{}{k})d^2x\mathrm{exp}(i\stackrel{}{k}\stackrel{}{x})s(\stackrel{}{x})`$, which are uncorrelated for different wave-vectors $`\stackrel{}{k}`$. The overall information is then obtained from a sum of independent contributions, and using $`_\stackrel{}{k}Ad^2k/(2\pi )^2`$ where $`A`$ is the cortical area, equal to

$$=\frac{A}{2}\frac{d^2k}{(2\pi )^2}\mathrm{ln}\left[1+\left|F(\stackrel{}{k})\right|^2𝒮(\stackrel{}{k})\right],$$
(7)
where $`F(\stackrel{}{k})`$ and $`\stackrel{~}{S}(\stackrel{}{k})`$ are Fourier transforms of the receptive field $`F(\stackrel{}{x})`$, and signal to noise correlations $`S(\stackrel{}{x})/N`$, respectively.

The task is to find the function $`F(\stackrel{}{k})`$ which maximizes the information $``$. Clearly, we need to impose certain costs on this function, since otherwise the information gain can become enormous for $`F\mathrm{}`$. This cost ultimately originates from the difficulties of creating and maintaining neural connections that gather and transmit information over some distance, and is hard to quantify. Following Ref. BRZ , we shall assume that the overall cost (in appropriate ‘information’ units) has the form

$$𝒞=d^2xC\left(\stackrel{}{x}\right)F\left(\stackrel{}{x}\right)^2d^2x\frac{\lambda +\mu x^2}{2}F\left(\stackrel{}{x}\right)^2=\frac{A}{2}\frac{d^2k}{(2\pi )^2}\left[\lambda \left|F(\stackrel{}{k})\right|^2+\mu \left|\stackrel{}{}_kF(\stackrel{}{k})\right|^2\right].$$
(8)
This expression can be regarded as an expansion in powers of $`F`$ and $`x`$, with the assumption that the cost is invariant under changing the sign of $`F`$, and independent of the direction of the vector $`\stackrel{}{x}`$. It imposes a penalty for creating connections which increases quadratically with the length of the connection. Our central conclusion is in fact insensitive to the form of $`C(\stackrel{}{x})`$.

If the costs are prohibitive, there will be no filtering of signals. To avoid such cases, we compare only filters that are constrained such that $`d^2xF\left(\stackrel{}{x}\right)^2=1`$ (or any other constant). In the optimization process, this constraint can be implemented via a Lagrange multiplier, resulting in an effective cost similiar to the term proportional to $`\lambda `$ in Eq. (8). Thus, this term and the constraint can be used interchangably. In Ref. BRZ  it was shown that optimizing Eq. (7) subject to the cost of Eq. (8), in the limit of low signal to noise, is equivalent to solving a Schrödinger equation with $`F(\stackrel{}{k})`$ playing the role of the wave function in a potential $`𝒮(k)`$, and the Lagrange multiplier taking the value of the ground state energy. A potential of the form $`𝒮(k)k^2`$ was there used to obtain receptive fields with on center/off surround character. In the next section we generalize this approach by considering correlated visual and aural inputs.

### II.2 Coupled Audio–Visual Inputs

In our idealized model, a neuron in the optic tectum of the owl filters input signals coming from both the visual and auditory systems, and its output is given by the generalization of Eq. (5) to

$$O\left(\stackrel{}{x}\right)=d^2yF_\alpha \left(\stackrel{}{x}\stackrel{}{y}\right)s_\alpha \left(\stackrel{}{y}\right)+\eta \left(\stackrel{}{x}\right),$$
(9)
where $`\alpha `$ is summed over A and V for audio and visual signals, respectively. Assuming as before that the signals $`s_\alpha `$ and the noise $`\eta `$ are independent, correlations of the output are obtained as

$$O(\stackrel{}{x}_1)O(\stackrel{}{x}_2)=d^2y_1d^2y_2F_\alpha \left(\stackrel{}{x}_1\stackrel{}{y}_1\right)F_\beta \left(\stackrel{}{x}_2\stackrel{}{y}_2\right)S_{\alpha \beta }\left(\stackrel{}{y}_1\stackrel{}{y}_2\right)+N\delta ^2\left(\stackrel{}{x}_1\stackrel{}{x}_2\right).$$
(10)
For translationally invariant signals, the output information is given by the generalization of Eq. (7) to

$$=\frac{A}{2}\frac{d^2k}{(2\pi )^2}\mathrm{ln}\left[1+F_\alpha (\stackrel{}{k})𝒮_{\alpha \beta }(\stackrel{}{k})F_\beta (\stackrel{}{k})\right],$$
(11)
where $`𝒮_{\alpha \beta }(\stackrel{}{k})`$ is a $`2\times 2`$ matrix of (Fourier transformed) signal to noise correlations.

Once more, we have to impose some constraints in order to make the maximization of the information in Eq. (11) with respect to the functions $`F_\mathrm{V}`$ and $`F_\mathrm{A}`$ biologically meaningful. In principle, there could be different costs for connections processing aural and visual signals. In the absence of concrete data, we make the simple choice of using the same form as Eq. (8) for both sets of filters, so that the overall cost is

$$𝒞=\frac{A}{2}\frac{d^2k}{(2\pi )^2}\left[\lambda F_\alpha (\stackrel{}{k})F_\alpha (\stackrel{}{k})+\mu \stackrel{}{}_kF_\alpha (\stackrel{}{k})\stackrel{}{}_kF_\alpha (\stackrel{}{k})\right].$$
(12)
The first term in the above cost function can again be interpreted as a Lagrange multiplier $`\lambda `$ imposing a normalization constraint

$$d^2xF_\alpha (\stackrel{}{x})^2=A\frac{d^2k}{(2\pi )^2}F_\alpha (\stackrel{}{k})F_\alpha (\stackrel{}{k})=1.$$
(13)
### II.3 Signal Correlations

To proceed further, we need the matrix of signal to noise correlations, which has the form

$`𝒮(\stackrel{}{k})=\left(\begin{array}{cc}𝒮_\mathrm{V}(k)& (k)e^{i\stackrel{}{k}\stackrel{}{c}}\\ (k)e^{i\stackrel{}{k}\stackrel{}{c}}& 𝒮_\mathrm{A}(k)\end{array}\right).`$
The diagonal terms represent the self correlations of each signal. Since many sources generate both sight and sound, the audio and visual signals will be correlated. These correlations are captured by the off-diagonal term $`(k)`$. In the experiments on owlsBK , the visual signal is artificially displaced by a fixed angle in the horizontal direction. If we indicate this angle by the vector $`\stackrel{}{c}`$, an aural signal at location $`\stackrel{}{x}`$ becomes correlated with a visual signal at $`(\stackrel{}{x}+\stackrel{}{c})`$. After Fourier transformation, this shift appears as the exponential factor $`\mathrm{exp}(i\stackrel{}{k}\stackrel{}{c})`$ in the off-diagonal terms of the correlation matrix.

So far, we have treated sight and sound on the same footing. It is reasonable to assume that under most (well lit) conditions the quality of visual information is much higher than the aural one. For ease of computation, we shall further assume that the actual signal to noise ratio is quite small, resulting in the set of inequalities

$$𝒮_\mathrm{A}(\stackrel{}{k})(\stackrel{}{k})𝒮_\mathrm{V}(\stackrel{}{k})1.$$
(14)
In this limit of small signal to noise, the logarithm in Eq. (11) can be approximated by its argument (without the one), resulting in a quadratic form in the filter functions. Our task then comes down to maximizing the function

$`𝒲\left[F_\alpha (\stackrel{}{k})\right]𝒞`$ $`=`$ $`{\displaystyle \frac{A}{2}}{\displaystyle }{\displaystyle \frac{d^2k}{(2\pi )^2}}\left[𝒮_\mathrm{V}(\stackrel{}{k})\right|F_\mathrm{V}(\stackrel{}{k})|^2+𝒮_\mathrm{A}(\stackrel{}{k})|F_\mathrm{A}(\stackrel{}{k})|^2`$ (15)
$`+(\stackrel{}{k})\left(F_\mathrm{V}(\stackrel{}{k})F_\mathrm{A}^{}(\stackrel{}{k})e^{i\stackrel{}{k}\stackrel{}{c}}+F_\mathrm{V}^{}(\stackrel{}{k})F_\mathrm{A}(\stackrel{}{k})e^{i\stackrel{}{k}\stackrel{}{c}}\right)`$
$`\lambda \left(\right|F_\mathrm{V}(\stackrel{}{k})|^2+\left|F_\mathrm{A}(\stackrel{}{k})|^2\right)\mu \left(\right|\stackrel{}{}_kF_\mathrm{V}(\stackrel{}{k})|^2+\left|\stackrel{}{}_kF_\mathrm{A}(\stackrel{}{k})|^2\right)],`$
with respect to $`F_\mathrm{V}`$ and $`F_\mathrm{A}`$.

### II.4 Results

The optimal filters are obtained from functional derivatives of Eq. (15). Setting the variations with respect to $`F_\mathrm{V}^{}(\stackrel{}{k})`$ to zero gives

$$\left[𝒮_\mathrm{V}(\stackrel{}{k})+\mu \stackrel{}{}_k^2\right]F_\mathrm{V}(\stackrel{}{k})=\lambda F_\mathrm{V}(\stackrel{}{k})\left\{(\stackrel{}{k})F_\mathrm{A}(\stackrel{}{k})e^{i\stackrel{}{k}\stackrel{}{c}}\right\},$$
(16)
while $`\delta W/\delta F_\mathrm{A}^{}=0`$, leads to

$$\left[\lambda \mu \stackrel{}{}_k^2\right]F_\mathrm{A}(\stackrel{}{k})=(\stackrel{}{k})F_\mathrm{V}(\stackrel{}{k})e^{i\stackrel{}{k}\stackrel{}{c}}+\left\{𝒮_\mathrm{A}(\stackrel{}{k})F_\mathrm{A}(\stackrel{}{k})\right\}.$$
(17)
In arranging the above equations, we have placed within curly brackets terms that are much smaller according to the hierarchy of inequalities in Eq. (14). Note that in the absence of any correlations between the two signals ($`=0`$), $`F_\mathrm{A}=0`$, since the aural signal is assumed to be much weaker than the visual one. Any non-zero $`F_\mathrm{A}`$ reduces $`F_\mathrm{V}`$ due to the normalization condition, resulting in a smaller value of $`𝒲`$. It is indeed the correlations between the two signals that lead to a finite value of $`F_\mathrm{A}`$, of the order of $`(/𝒮_\mathrm{V})`$. (Since $`\lambda 𝒪(𝒮_\mathrm{V})`$ as shown below.)

To leading order, Eq. (16) is the Schrödinger equation obtained in Ref. BRZ  for the visual receptive field. Without further discussion, we shall indicate its solution by

$$F_\mathrm{V}(\stackrel{}{x})=F_\mathrm{V}^0(\stackrel{}{x}),\mathrm{and}\lambda =E_\mathrm{V}=𝒪\left(𝒮_\mathrm{V}\right).$$
(18)
Note that we don’t imply that cells in the optic tectum should have a receptive field for visual signals identical to that in the visual cortex. The quality of signals, the costs of neural connections, and the response of the cells may well vary from one cortical area to another. The eigenvalue $`E_\mathrm{V}`$ is controlled by the strength of the visual correlations and is of the order of $`𝒮_\mathrm{V}`$.

To simplify the solution to Eq. (17), we first assume that $`(\stackrel{}{k})=R`$, a constant independent of $`\stackrel{}{k}`$. This is quite a reasonable assumption, corresponding to visual and aural signals that are correlated only if coming from the same direction, i.e. with $`s_\mathrm{V}(\stackrel{}{x}_1)s_\mathrm{A}(\stackrel{}{x}_2)=R\delta ^2(\stackrel{}{x}_1\stackrel{}{x}_2)`$. We can then Fourier transform the two sides of this equation to obtain

$$\left(E_\mathrm{V}+\mu x^2\right)F_\mathrm{A}(\stackrel{}{x})=RF_\mathrm{V}^0(\stackrel{}{x}\stackrel{}{c}),$$
(19)
and quite generally, for an arbitrary form of the cost function in Eq. (8), the solution is

$$F_\mathrm{A}(\stackrel{}{x})=\frac{R}{E_\mathrm{V}+C_A(x)}F_\mathrm{V}^0(\stackrel{}{x}\stackrel{}{c}).$$
(20)
Due to the quadratic form of Eq. (15), the above result is the linear response of the system to the correlations between signals.

The significance of our result is that the aural receptive field $`F_A(\stackrel{}{x})`$ is not simply the visual receptive field shifted by $`\stackrel{}{c}`$, as one might have guessed. Rather, the shape of $`F_A(\stackrel{}{x})`$ could be significantly distorted by the cost function $`C_A(x)`$. At the moment, the data may be too crude to determine the shape of $`F_A(\stackrel{}{x})`$, but it is still worthwhile to contemplate what sort of shape distortion may result in our simple model. For illustrative purposes, let us take $`F_V(\stackrel{}{x})\mathrm{exp}\left[(x/l)^2\right]`$ to be a Gaussian with $`l`$ a length scale characteristic of the visual receptive field and $`C_A(x)=\mu x^2`$. Then we predict (with $`\stackrel{}{c}=(c,0)`$ and $`\stackrel{}{x}=(x,y))`$

$$F_A(x,y)\frac{1}{1+\left(x^2+y^2\right)/L^2}\mathrm{exp}\left[\frac{(xc)^2+y^2}{l^2}\right],$$
(21)
where $`L\sqrt{E_V/\mu }`$ defines a length scale characteristic of the relative cost of connecting distant neurons. While there are three length scales $`L`$, $`l`$, and $`c`$ inovlved, the shape of $`F_A(x,y)`$ depends only on the two ratios $`L/l`$ and $`c/l`$.

We now qualitatively describe the change in the shape of the aural receptive filed in Eq. (21), as the imposed shift $`c`$ is varied as in the experiments of Knudsen et al. (The exact analysis of the extremal points of Eq. (21) involves the solution of a cubic equation which will not be given here.) Two types of behavior are possible depending on the ratio $`l/L`$. For $`lL`$, where the cost of rewiring is negligible, the function $`F_A(x,y)`$ has a single maximum located at $`xc`$ (and $`y=0`$), i.e. simply following the imposed shift. When $`lL`$, however, there is an intermediate range of values of $`c`$, where the aural receptive field has two peaks, one close to the origin, $`\overline{x}_{}cL/lc`$, and another close to $`\overline{x}_+c`$. A typical profile with two peaks is depicted in Fig. 1.

## III Discussions

Equation (20) is the central result of our study. It provides the optimal linear filter for a weak signal correlated to a stronger one. Some specific features of this result in connection with the coupled visual and aural maps are:

* The shape of the aural receptive field is very much controlled by the visual information, modulated by the costs associated with neural connectivities.
* Artificially displacing the two signal sources, as in the case of the prismatic spectacles used on the barn owlsBK , modifies the aural receptive field. However, the resulting receptive field is not simply shifted (unless the costs of neural wirings are negligible), but also changes its shape.
* Equation (20) is the product of two functions, one peaked at the origin and the other at $`\stackrel{}{x}=\stackrel{}{c}`$. Depending of the relative strengths and widths of these two peaks, the receptive field may be more sensitive to signals at the original, or in the shifted location.
* The experiments find, not surprisingly, that adaptation to the prismatic glasses depends strongly on the age of the individual owl. This feature can be incorporated in our model with the reasonable assumption that the cost of neural connections increases with age of the individual.

This work is small step towards providing a quantitative framework for deducing the workings of the brain, starting from the tasks that it has to perform for the organism to function in its natural habitat. In this framework, the tasks of the sensory systems are more apparent: to extract the relevant signals from the background of natural inputs, and as a first step to localize the source of the signal in the external world. It is possible to experimentally gather information about the correlations of various signals in the natural world, and there are indeed several studies of the statistics of various aspects of visual imagesnatural . Of course, such statistics are also specific to the instrument (e.g. camera) used to obtain the image. More relevant are psycho-physical studies that probe how individuals parse the visual informationMalik . We are not aware of similar studies on the statistics of natural sounds in different directions, and their correlations with visual signals. Such studies may provide part of the material needed for a more detailed study.

The outcome of the procedure outlined in this paper is a set of filter functions, which are hopefully related to the actual connections between neurons. The shape and range of such connections can be studied directly by injection of biocytin dyedye , and indirectly by mapping the receptive field of a neuron via a microelectrode probe. Detailed studies of this kind for the owls reared with prismatic spectacles, and their comparison with Eq. (20) may provide insights about the cost of making neural connections, another necessary input to our general formalism.

The analytical formalism itself can be extended in several directions. Already, in Ref. BRZ  it was proposed that colored images can be studied by considering a vector signal $`\stackrel{}{s}`$ ranging over the color wheel. In regards to different sensory inputs, we may can also ask if and when it is advantageous to segregate outputs to distinct cortical areas, allowing for distinct maps $`\{O_\nu \}`$. A more ambitious goal is to extend the formalism to time dependent signals, allowing for filters with appropriate time delays that attempt to take advantage of temporal patterns in the signals.

###### Acknowledgements.

This work was supported in part by the NSF under grant numbers DMR-01-18213 (MK), PHY89-04035 and PHY95-07065 (AZ).