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Ординатура / Офтальмология / Английские материалы / Computational Maps in the Visual Cortex_Miikkulainen_2005

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C

HLISSOM Simulation Specifications

The HLISSOM model extends basic LISSOM by including afferent normalization (Equation 8.1) that allows processing natural images, as well as a PGO sheet and a face-selective area. The PGO sheet is connected to the LGN and V1 is connected to the FSA so that each unit has a full receptive field (Figure A.1). The simulation parameters are based on the default LISSOM parameters of Appendix A, except as specified in the subsections that follow.

C.1 V1 Only

The V1-only simulations showing the effect of γn (Figures 8.2 and 8.3) were based on disk-shaped input patterns. The parameters were the same as in the “Disks” simulation in Section A.5.1, except the area was very large (sg = 8) to allow large, detailed retinal stimuli to be tested, the cortical density was very low (Nd = 24) to reduce memory and computational requirements, and the LGN radius was slightly smaller (σc = 0.4) for historical reasons. During self-organization, γn was zero; after self-organization, that parameter was adjusted as shown in the figure.

The small-scale V1-only HLISSOM simulations in Figure 8.5 and in Chapter 9 were based on the default LISSOM simulation parameters of Appendix A; in particular, afferent normalization was not used (γn = 0). In the input and weight stream simulations (Figure 8.5) the cortical density was lower (Nd = 64) so that the random initial weights would be visible.

The V1 simulation parameters were always evolved on a schedule for a 10,000iteration run; in the prenatal simulations of Figure 9.1, the training was simply interrupted at 1000. The prenatal “Disks” and “Noisy Disks” simulations were based on the same parameters as the full-length simulations described in Section A.5.1, up to iteration 1000. The prenatal “Noise” simulation was identical to “Noisy disks”, except no disks were drawn.

The “Nature” simulation in Figure 9.6 was based on the “Images” simulation described in Section A.5.2, except the inhibitory and excitatory strengths were fixed (γI = 2.0 and γE = 1.2), and the input density scale was higher (sd = 8) without

430 C HLISSOM Simulation Specifications

Parameter

Value

 

 

 

 

 

 

 

 

 

Nd

 

24

 

 

 

Rd

 

48

 

 

 

rA

Ld

 

 

+ 0.5

 

 

 

 

 

 

0.96

 

 

 

rEi

 

Nd

 

 

 

6

 

 

 

 

 

 

 

 

rI

 

Nd

 

 

 

2.4

 

 

 

 

 

 

 

 

θli

 

0.1

 

 

sw

 

9.5

 

 

 

 

rAo

 

 

 

 

σA

 

9.5

 

 

 

 

1.3

 

 

 

 

 

 

 

 

σc

0.75sw

 

 

 

 

 

 

 

 

Parameter

Value

 

 

 

 

 

 

σs

1.6σc

 

dr

20 6.5sw

 

 

9.5

 

 

 

sd

1.5

 

 

 

 

st

0.5

 

 

 

 

np

max(1, 0.5sdsr)

 

γA

1.07

 

 

 

nA

 

 

 

 

 

 

 

 

 

γL

10.2

 

 

 

sb

 

 

 

 

 

rI2o

 

wd

6wdo

 

 

 

 

 

 

r

2

 

 

 

 

 

I

 

 

 

 

 

Table C.1. Defaults for FSA simulations. FSA simulations were based on the defaults from Table A.2, modified as shown in this table. Some of these defaults are overridden in individual FSA simulations, as described in the text.

changing the time scale (st = 0.5). These same parameters were used in all postnatal simulations, but starting at 1000 after the prenatal training described above.

The simulations with prenatal training followed by natural images (Figures 9.3– 9.6) were based on the prenatal training parameters specified until 1000, followed by 9000 iterations with the “Nature” parameters.

C.2 Face-Selective Area Only

The FSA (like any cortical area) has its own independent set of parameters. To keep the notation simple, the same parameter names are used for the FSA as for V1, and each area is described separately in this appendix. For example, when discussing the FSA parameters, Nd refers to the size of the FSA network and Ld to the size of the V1 network.

The default FSA parameter values are listed in Table C.1, overriding those in Table A.2. The values for σA, sw, and dr were determined empirically in earlier work (Bednar and Miikkulainen 2000a). The afferent radius rA was significantly increased compared to the V1 simulations to allow whole faces and face outlines to be learned. The other parameters were adjusted accordingly; for example, to allow the initial activity bubbles to remain similar to those in networks with a smaller afferent radius, the afferent weights were initialized with a fixed-width Gaussian instead of random noise.

The prenatal phase of the FSA-only simulations in Section 10.3 (Figure 10.12) was identical to the default simulation in Table C.1.

The simulations with the different face training pattern types (Figure 10.11) were otherwise identical to the default, except γA was varied as shown in Table C.2 to ensure that the average FSA activity was the same for each pattern. These values

C.2 Face-Selective Area Only

431

Figure

Pattern

γA

10.11a

 

1.070

10.11b

 

0.697

10.11c

 

0.697

10.11d

 

0.550

10.11e

 

0.577

 

 

 

Figure Pattern

γA

10.11f

0.490

10.11g

1.983

10.11h

1.416

10.11i

0.948

Table C.2. Parameters for different types of face training patterns. Each of the simulations in the subfigures of Figure 10.11 was based on the same parameters, except γA was adjusted by hand until the average activity resulting from each pattern was similar. The resulting γA is shown for each pattern in this table.

Iteration

θl (prenatally trained)

θl (na¨ıve)

αA

20000st

θli + 0.070

10

αAi

θli

 

 

70

 

 

20400st

θli + 0.070

10

αAi

θli

 

 

70

 

 

22000st

θli + 0.070

 

9

αAi

θli

70

 

 

 

 

24000st

θli + 0.070

 

8

αAi

θli

70

28000st

θli + 0.070

 

7

αAi

θli

70

 

 

 

 

32000st

θli + 0.070

 

7

αAi

θli

70

 

 

 

 

36000st

θli + 0.080

 

6

αAi

θli

70

 

 

 

 

40000st

θli + 0.090

 

6

αAi

θli

70

 

 

 

 

+0.120

+0.120

+0.120

+0.090

+0.070

+0.070

+0.050

+0.045

Table C.3. Parameter change schedule for postnatal FSA simulations. The FSA simulations in Section 10.3 continued past 20,000 iterations to model postnatal learning; the formulas above describe how the parameter values were obtained for these additional iterations. Together, Tables A.3 and C.3 specify the parameter change schedule for the entire FSA-only simulations.

were determined by presenting a set of random inputs while adjusting γA until the sum of the cortical response was the same as for the three-dot pattern (Figure 10.11a).

The postnatal phase of the FSA-only simulations in Section 10.3 (Figure 10.15) continued with the parameters that were in effect at the end of prenatal training, except αA was reduced further as shown in Table C.3, the sigmoid range θu − θl was reduced to 0.48, and the sigmoid’s lower threshold θl was set separately for each network as described in Section 10.3 and shown in Table C.3.

432

C

HLISSOM Simulation Specifications

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

V1 settings

 

 

 

 

FSA settings

 

 

 

Iteration

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

θl

γA

γE

γI

γn

 

θl

γA

γE

γI

γn

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

28000st

 

θli + 0.140

1.90

0.9

0.9

0

θli + 0.130

3.0

0.9

0.9

0

 

 

30000st

 

θli + 0.267

2.70

1.0

1.1

1

θli + 0.250

5.0

0.5

0.7

2

 

 

35000st

 

θli + 0.317

2.90

1.1

1.3

2

θli + 0.400

9.0

0.4

0.6

5

 

 

40000st

 

θli + 0.417

3.25

1.2

1.4

4

θli + 0.710

10.6

0.4

0.6

9

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Table C.4. Parameter change schedule for combined V1 and FSA simulations. During iterations 20,000–28,000st, the network with both V1 and FSA (Section 10.2) was trained using the schedule for iterations 0–8000st in Table A.3. Beyond 28000st, it was trained as shown in this table. Because each cortical area has an independent set of parameters, the values for V1 and the FSA are listed separately.

In the statistical significance tests discussed in Figures 10.16 and 10.17, each input image was presented at 16 different locations chosen from the nodes of a regular 4 × 4 grid centered on the retina, with each grid step four retinal units wide. The example images shown in these figures are located at the center of the retina.

C.3 Combined V1 and Face-Selective Area

In the simulations of Section 10.2, V1 and FSA were combined into a single model. The retina, LGN, and V1 parameters were identical to the “Disks” simulation in Section A.5.1, except V1 density was very low (Nd = 24) to reduce memory and computational requirements, the V1 area was very large (sg = 8) to allow large retinal stimuli to be tested, the LGN radius was slightly smaller (σc = 0.4) to match earlier simulations, and the simulation continued past 20,000st. The FSA parameters were identical to the default FSA-only parameters (Section C.2), except the FSA was slightly more dense (Nd = 36/0.94), the input region was significantly more dense (Ld = 170) because the FSA connects to V1 and not to LGN, the area scale corresponded to the full area of V1 (sg = 0.94), the afferent radius was smaller (rA = 0.375Ld + 0.5) and the weights initially random because only small training patterns were used, the afferent scale was larger (γA = 3) to compensate for the patchy activity in the input region (V1), and the parameters were changed on the schedule listed in Table C.4. The FSA also had only one input (V1) instead of two (the ON and OFF LGN channels), i.e. nA = 1 for the FSA.

Because FSA training followed that of V1, V1 training iteration 20,000st was treated as if it was iteration zero in FSA training, and thus the parameters e.g. for iteration 28000st were determined like parameters for iteration 8000st of the default FSA simulation. After iteration 28000st, the V1 and FSA parameters followed the schedule shown in Table C.4. The V1 parameters were chosen to ensure that the network responded well to large natural images. First, the value of γn was gradually

C.3 Combined V1 and Face-Selective Area

433

increased from zero to make the responses less dependent on image contrast. As a result, similar shapes that have different contrasts in different areas of an image would lead to comparable V1 responses. The other parameters were adjusted to compensate for the effect of γn, ensuring that V1 responses to the highest-contrast patterns were near 1.0, and lower contrasts resulted in lower V1 responses. The parameters of the FSA were chosen similarly, except the FSA sigmoid threshold θl was also gradually increased so that the response would be nearly binary. In this way, the FSA response was used to decide whether a facelike pattern had occurred in the input, as described in Section 10.2.

D

PGLISSOM Simulation Specifications

The PGLISSOM simulations focused on demonstrating valid self-organization of the orientation map, and on grouping visual features. These simulations were based on the reduced LISSOM specifications of Appendix B, with slightly different parameter values to reduce the required computational resources.

D.1 Self-Organization

The self-organization simulations in Section 11.5 form the baseline for all PGLISSOM simulations. The parameter values for this simulation are specified in Table D.1 and the schedule for parameter adaptation over the course of self-organization in Table D.2. The parameter values were found to be effective by running several experiments, and small changes to them did not affect the global behavior of the model.

All weights were initialized with uniformly random values distributed within [0..1]; the afferent connections were mapped to the retina so that each neuron had a complete receptive field (Figure A.1). GMAP was smaller than SMAP to make simulations faster and more compact. The γC was lower in SMAP than in GMAP so that activity caused by high excitation in GMAP would not interfere with selforganization in SMAP, and to allow the self-organized global order of SMAP to be transferred to GMAP. The value γE was lower and γI higher in GMAP than in SMAP to prevent the map from becoming too active.

The inhibitory connections in GMAP did not adapt (αIG = 0); the initial broad connectivity remains to provide background inhibition, as explained in Section 11.2. The afferent and intracolumnar learning rates αA and αC in both maps were decreased over time, so that the order in the map could gradually start stabilizing. The long-range lateral inhibitory connections in SMAP and long-range excitatory lateral connections in GMAP were pruned on the same schedule, specified by wd and td.

The base thresholds θb in both maps were set to θb0 when the input was first presented, the activation-based values in Table D.1 were then computed, and the base thresholds were fixed to those values for the remaining settling iterations. As a result, the network would not become too active or completely silent for any of the training

436 D PGLISSOM Simulation Specifications

Parameter

Value

 

 

NS

136

NG

54

L

46

rA

6

rESi

7

rEG

40

rIS

10

rIG

54

σai

3.9

σbi

0.8

θl

0.01

θu

1.3

γA

1.1

 

 

Parameter

Value

 

 

γES

0.8

γEG

0.2

γIS

0.9

γIG

2.5

tf

40000

np

1

αAi

0.012

αE

0.008

αIS

0.008

αIG

0.0

wd

0.001

td

40000

Parameter

Value

Used in

Description

rC

2

Section 11.2

θb0

0.05

Equation 11.5

γbSi

0.5

Table D.2

γbGi

0.5

Table D.2

θbS

γbS maxij vij (t)

Equation 11.5

θbG

γbG maxij vij (t)

Equation 11.5

γCS

0.5

Equation 11.3

γCG

0.9

Equation 11.3

γθ

0.4

Equation 11.5

λE

3.0

Section 11.3.1

λI

0.5

Section 11.3.1

λC

1.0

Section 11.3.1

λθ

0.5

Equation 11.6

λr

0.92

Equation 11.7

tr

0

Section 11.3.3

tw

15

Section 11.4

αCi

0.012

Equation 11.8

Maximum radius of the intracolumnar connections Base threshold at the beginning of settling

Initial value for γbS, SMAP base threshold scaling factor Initial value for γbG, GMAP base threshold scaling factor SMAP base threshold

GMAP base threshold

Scaling factor for the SMAP intracolumnar weights Scaling factor for the GMAP intracolumnar weights Scaling factor for the relative refractory period Excitatory synaptic decay rate

Inhibitory synaptic decay rate

Intracolumnar synaptic decay rate

Refractory period decay rate

Spiking rate decay rate

Length of the absolute refractory period

Length of time window for computing average spiking rate Initial value for γC, the intracolumnar learning rate

Table D.1. Defaults for PGLISSOM simulations. The subscript “S” identifies parameters of the SMAP, and “G” those of the GMAP; parameters without these subscripts had the same values for both maps. Although many of the parameters are the same as in firing-rate LISSOM models (top half), their default values are slightly different to take into account the two-map architecture and the spiking units. A number of new parameters related to intracolumnar connections and spiking are also introduced (bottom half). Some of these defaults are overridden in the synchronization and grouping simulations, as described in the text.

 

 

 

 

 

D.2

Grouping

437

 

 

 

 

 

 

 

 

Iteration

rES

γbS

γbG

αA

αC

σa

σb

 

 

 

 

 

 

 

 

0st

rESi

γbSi

500st

0.57rESi

γbSi

1000st 0.429rESi

γbSi

5000st

0.429rESi

γbSi

15000st 0.429rESi 1.15γbSi 40000st 0.429rESi 1.15γbSi

γbGi

γbGi

γbGi 1.3γbGi 1.3γbGi 1.3γbGi

αAi αAi αAi

0.667αAi

0.667αAi

0.667αAi

αCi

σai

σbi

αCi

σai

σbi

αCi

1.718σai

0.875σbi

0.667αCi

1.718σai

0.875σbi

0.667αCi

1.718σai

0.875σbi

0.667αCi

1.718σai

0.875σbi

Table D.2. Parameter change schedule for PGLISSOM simulations. Starting with the initial values given in Table D.1, these parameters were adapted at each iteration as shown in the table.

inputs. The scaling factors γbS and γbS were later adjusted as shown in Table D.2 so that the activities could gradually become sparser. This method has a similar effect as adapting the sigmoid activation function (described in Section 4.4.3). To speed up self-organization, tr was set to zero so that the neurons could fire as rapidly as possible.

The input consisted of single randomly located and oriented elongated Gaussians. Over time, these Gaussians were made longer than in previous LISSOM simulations, so that sharper orientation tuning and longer lateral connections could develop, improving contour integration in the model (self-organizing very long-range lateral connections is computationally expensive and was therefore avoided in previous simulations). Continuous input values were used to approximate spiking input, making the simulations more efficient without discernible effect on how the model behaves. The training took about 30 hours and 178 MB of memory on a 1 GHz AMD Athlon Linux machine.

D.2 Grouping

After self-organization, several parameters were adjusted slightly in order to make grouping more robust in Chapter 13. First, the excitatory learning rate αE in GMAP was set to 0.1. Although not strictly necessary for grouping, such fast adaptation of lateral excitatory connections allows the network to quickly adjust the weights remaining after connection death to a level that allows robust synchronization (von der Malsburg 1981, 2003; Wang 1996). It does not affect the patchy structure of the lateral connections nor the organization of the map. Second, in GMAP γE was increased to 0.8, γI increased to 5.0, and 4% noise was added to the membrane potential of all GMAP neurons, as described in Sections 12.3.2 and 12.4.2. Third, the absolute refractory period tr was increased to 4.0. After self-organization, fast simulation is not critical, but it is important to have a high enough temporal resolution of activity so that multiple groups of neurons can desynchronize at the same time. Higher tr results

438 D PGLISSOM Simulation Specifications

in such higher resolution; it also makes synchronization more robust, as discussed in Section 12.4.3.

Each contour integration test pattern was generated to approximate the patterns used with human subjects (such as that in Figure 13.1) as well as possible within the small model retina and cortex. The input patterns consisted of oriented Gaussians with σa = 1.87 and σb = 1.22. The elements of each contour were placed on the retina at approximately collinear locations (cocircular in the curvature experiment of Section 13.4) with a minimum separation of 0.5σa2 and an orientation that corresponded to the desired degree of jitter. In addition, the neurons in SMAP and GMAP had to be highly selective for that input in that location. The background elements were then placed on the retina at random locations and orientations with the same distance and selectivity constraints until no more possible such locations existed.

In the input distribution experiments in Section 13.4, slightly broader Gaussians were used to train the networks, and the Gaussians became elongated slightly slower. Initially, σa = 3.9 and σb = 1.1, and they were increased to σa = 7.1 and σb = 0.9 at iteration 10,000. As a result, self-organization was slower; on the other hand, the connectivity patterns remained broader, allowing the networks to perform robustly on the wider variety of inputs.

D.3 Synchronization

The synchronization and desynchronization simulations in Chapter 12 were all based on a simplified PGLISSOM network where synchronization behavior could be clearly demonstrated. The network consisted of a one-dimensional array of neurons, connected one-to-one to input and output. The input neurons spiked at every time step, and the membrane potential of each neuron was initialized to uniformly random distribution within [0..1]. The afferent weights were fixed at 1.0, the lateral excitatory weights at 1.0/nE (where nE is the number of incoming excitatory lateral connections of the neuron), and the lateral inhibitory weights at 1.0/nI (where nI is the number of incoming inhibitory lateral connections). The afferent contribution was set to γA = 0.63 and decay rate to λA = 1.0, spike generator decay rate was λθ = 0.05, sigmoid threshold and ceiling θl = 0.0, θu = 3.0, relative refractory threshold proportion γθ = 0.65, and base threshold θb = 0.1.

The remaining parameters were systematically varied in the experiments: (1) lateral excitatory and inhibitory connection patterns and radii rE and rI, (2) their contributions to the neuron activation γE and γI, (3) their synaptic decay rates λE and λI,

(4) the size and pattern of the afferent input, (5) the degree of noise in the membrane potential, and (6) the duration tr of the absolute refractory period.

E

SOM Simulation Specifications

In order to establish a baseline for comparison with LISSOM, the simulations in Chapter 3 were based on the SOM version of self-organizing maps, where the response is based on Euclidean distance similarity measure, and the weights are changed to minimize Euclidean difference between the input and the weight vector (Equation 3.15).

The default parameters, used in the basic self-organization simulation of Figure 3.6, are listed in Table E.1. The receptor surface was fully connected to each map unit. The connections were initialized to uniformly random values within [0..1]. The input consisted of single unoriented Gaussians whose centers were chosen from a uniformly random distribution so that they were evenly scattered over the receptor surface.

The magnification simulation (Figure 3.7) was run with the same network and learning parameters; the only difference was the distribution of the input Gaussians. The network was trained up to 10,000 iterations as before, at which point the input distribution was changed. Instead of a uniformly random distribution, in Figure 3.7a a Gaussian high-density area with center (0.5, 0.5) and radius 0.01 was added to a uniformly random distribution of range [0..0.2]. In Figure 3.7b, the distribution

consisted of a uniform distribution and two Gaussian high-density areas, one with center (0.5, 0.25) and the other (0.5, 0.75), and the major and minor axes of 0.05 and 0.0125 for both areas.

The three-dimensional model of ocular dominance (Figure 3.10) was abstracted further by representing the input location as a two-dimensional variable (x, y), instead of an activation pattern on a two-dimensional surface as in the two previous simulations. These values were drawn from a uniformly random distribution within [0..1]. The third variable, representing ocular dominance, was also uniformly randomly distributed, but within [0.15..0.15]. The inputs were thus three-dimensional vectors of (x, y, z) values, fully connected to the map units. The network was

self-organized in tf = 120, 000 input presentations, with the adaptation rate = 0.1 exp(4.0 t/tf ) and neighborhood width = max[13.3 exp(5.0 t/tf ), 0.8].

The specifications for the SOM in handwritten digit recognition simulations will be described in Appendix F.2.