Ординатура / Офтальмология / Английские материалы / Computational Maps in the Visual Cortex_Miikkulainen_2005
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418 |
A |
LISSOM Simulation Specifications |
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Parameter |
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Value |
Used in |
Description |
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Nd |
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Ndo |
Table A.2 |
Cortical density, i.e. width and height of a unit area of cortex |
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Ld |
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Ldo |
Table A.2 |
LGN density, i.e. width and height of a unit area of the LGN |
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(the area that projects to Nd) |
Rd |
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Rdo |
Table A.2 |
Retinal density, i.e. width and height of a unit area of retina |
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(the area that projects to Ld) |
sg |
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1.0 |
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Table A.2 |
Global size scale of the model in area units Nd, Ld, and Rd |
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nA |
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2 |
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Table A.2 |
Number of afferent RFs per cortical unit (e.g. 1 ON and 1 OFF) |
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L |
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rA |
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d |
+ 0.5 |
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Section 4.2.3 |
Maximum radius of the cortical afferent connections† |
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4 |
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rEi |
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Nd |
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Section 4.2.3 |
Initial value for rE, the maximum radius of the lateral excita- |
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tory connections, before shrinking† |
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Nd |
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rEf |
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max(2.5, |
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Table A.3 |
Minimum final value of the rE after shrinking† |
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rI |
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Nd |
− 1 |
Section 4.2.3 |
Maximum radius of the lateral inhibitory connections† |
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sw |
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rA |
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Table A.2 |
Scale of rA relative to the default |
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rAo |
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σA |
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rA |
Section 4.2.3 |
Radius of the initial Gaussian-shaped afferent connections† |
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1.3 |
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σE |
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0.78rEi |
Section 4.2.3 |
Radius of the initial Gaussian lateral excitatory connections† |
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σI |
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2.08rI |
Section 4.2.3 |
Radius of the initial Gaussian lateral inhibitory connections† |
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σc |
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0.5sw |
Rd |
Equation 4.1 |
Radius of LGN DoG center Gaussian† |
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Ld |
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σs |
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4σc |
Equation 4.1 |
Radius of LGN DoG surround Gaussian† |
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rL |
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4.7σs |
Section 4.2.2 |
Maximum radius of the LGN afferent connections† |
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N L
R
sr
sgNd
sgLd + 2(rA − 0.5)
sg Rd + 2 RLdd (rA − 0.5) +2(rL − 0.5)
2
L
Ld+2(rA−0.5)
Section 4.2.1
Section 4.2.1
Section 4.2.1
Table A.2
Width and height of the cortex, in number of units Width and height of the LGN, in number of units
Width and height of the retina, in number of units
LGN area scale relative to the reference simulation†
σu |
3sw |
Equation 4.2 |
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σa |
σaosw |
Equation 5.1 |
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σb |
σbosw |
Equation 5.1 |
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rd |
25.0sw |
Equation 9.2 |
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σd |
3.0sw |
Equation 9.2 |
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sb |
1.0 |
Section 5.4.1 |
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ob |
0.5 |
Section 4.2.1 |
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Rp |
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L |
Section 4.3.1 |
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dr |
2.2rA |
Section 4.3.1 |
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ss |
0.0 |
Section 5.4.4 |
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sd |
2.0 |
Table A.2 |
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st |
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1 |
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Tables A.2, A.3 |
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sd |
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np |
max(1, sdsr) |
Section 4.3.1 |
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Radius of unoriented Gaussian inputs
Radius of the major axis of ellipsoidal Gaussian inputs Radius of the minor axis of ellipsoidal Gaussian inputs Radius of the full-brightness portion of disk-shaped patterns† Radius of the Gaussian falloff in brightness at the disk edge Brightness scale of the retina (contrast of fully bright stimulus) Brightness value of the background of the retina
Width & height of the random scatter of discrete pattern centers Minimum separation between the centers of multiple inputs† Scale of the input pattern scatter from the calculated value Input density scale (ratio between average cortical activity for one oriented Gaussian to the average for the actual pattern) Iteration scaling factor; can be adjusted to use fewer iterations if input patterns are more dense at each iteration, or vice versa Number of discrete input patterns per iteration (e.g. Gaussians)
(Table continues on the next page)
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A.3 |
Choosing Parameters for New Simulations |
419 |
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(Table continued from the previous page) |
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Parameter |
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Value |
Used in |
Description |
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γA |
1.0 |
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Equation 4.5 |
Scaling factor for the afferent weights |
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γE |
0.9 |
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Equation 4.7 |
Scaling factor for the lateral excitatory weights |
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γI |
0.9 |
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Equation 4.7 |
Scaling factor for the lateral inhibitory weights |
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γL |
2.33 |
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Equation 4.3 |
Scaling factor for LGN’s afferent weights† |
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sb |
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γn |
0.0 |
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Equation 8.1 |
Strength of divisive gain control (only in HLISSOM) |
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tsi |
9 |
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Section 4.3.3 |
Initial value for ts, the number of settling iterations |
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θli |
0.083 |
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Equation 4.4 |
Initial value for θl, the lower threshold of the sigmoid activa- |
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tion function |
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θui |
θli + 0.55 |
Equation 4.4 |
Initial value for θu, the upper threshold of the sigmoid activa- |
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tion function† |
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tf |
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tf ost |
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Section 4.4 |
Number of training iterations |
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αAi |
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0.0070 |
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Equation 4.8 |
Initial value for αA, the afferent learning rate† |
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nAstsd |
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αEi |
0.002rE2 o |
Equation 4.8 |
Initial value for αE, the lateral excitatory learning rate† |
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stsdrE2 |
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0.00025r2 |
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αI |
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I o |
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Equation 4.8 |
Lateral inhibitory learning rate |
† |
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stsdrI2 |
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rI2o |
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wd |
2wdo |
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Section 4.4.2 |
Lateral inhibitory connection death threshold |
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r |
2 |
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I |
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td |
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tf |
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Section 4.4.2 |
Iteration at which inhibitory connections are first pruned |
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Table A.2. Defaults for constant parameters. This table specifies how the parameter values for the different simulations can be constructed, based on the reference values from Table A.1. These parameters have constant values in each simulation. Those with the subscript “i” represent the initial values for parameters that are changed over the simulation, as shown in Table A.3. The table is organized into sections including user-defined network size, connection radius, calculated network size, and input pattern parameters on the previous page, and connection strength, activation, and learning parameters on this page. The numerical values in formulas marked with a dagger (†) were determined empirically in earlier work (Bednar and Miikkulainen 2000b; Sirosh 1995). Parameters that are listed as being used in Table A.2 are temporary variables, introduced to make the notation easier to follow. Those listed as being used in various equations and sections are actual parameters of the LISSOM model itself.
A.3 Choosing Parameters for New Simulations
Despite the seemingly large number of parameters, few of them need to be adjusted when running a new simulation. The most commonly changed parameters are the cortical density Nd and area scale sg, because these parameters directly determine the time and memory requirements of the simulations. The default Nd of 142 representing a 5 mm × 5 mm area is a good match to the density of columns in V1 orientation maps (Bednar, Kelkar, and Miikkulainen 2004), but in practice much smaller
420 A LISSOM Simulation Specifications
Iteration |
rE |
ts |
θl |
θu |
αA |
αE |
0st |
max(rEf , rEi) |
200st |
max(rEf , 0.6000rEi |
500st |
max(rEf , 0.4200rEi |
1000st max(rEf , 0.3360rEi 2000st max(rEf , 0.2688rEi 3000st max(rEf , 0.2150rEi 4000st max(rEf , 0.1290rEi 5000st max(rEf , 0.0774rEi 6500st max(rEf , 0.0464rEi 8000st max(rEf , 0.0279rEi 20000st max(rEf , 0.0167rEi
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tsi tsi tsi tsi
tsi + 1 tsi + 1 tsi + 1 tsi + 2 tsi + 3 tsi + 4 tsi + 4
θli θli θli θli θli θli θli θli θli θli
θli
+0.01
+0.02
+0.05
+0.08
+0.10
+0.10
+0.11
+0.12
+0.13
+0.14
θui θui θui θui θui θui θui θui θui θui
θui
+0.01
+0.02
+0.03
+0.05
+0.08
+0.11
+0.14
+0.17
+0.20
+0.23
αAi |
αEi |
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αAi |
αEi |
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50 |
αAi |
0.5αEi |
70 |
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50 |
αAi |
0.5αEi |
70 |
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40 |
αAi |
0.5αEi |
70 |
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40 |
αAi |
0.5αEi |
70 |
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30 |
αAi |
0.5αEi |
70 |
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30 |
αAi |
0.5αEi |
70 |
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30 |
αAi |
0.5αEi |
70 |
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30 |
αAi |
0.5αEi |
70 |
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15 |
αAi |
0.5αEi |
70 |
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Table A.3. Default parameter change schedule. The values of these parameters at the beginning of simulation are given in Table A.2; this table describes how their values change at each subsequent iteration. The new values are calculated at the start of each listed iteration.
values also often work well (as shown in Section 15.2.3). The default sg of 1.0 covers an area large enough to include several orientation patches in each direction, but more area is useful when processing larger images.
Apart from the simulation size parameters, most simulations differ primarily by the choice of input patterns. Starting from an existing simulation, usually only a few parameters need to be changed to obtain a similar simulation with a new set of patterns. If the new pattern is similar in overall shape, often all that is needed is to set the afferent input scale γA or the sigmoid threshold θli to a value that, on average, produces a similar level of cortical activity. Usually a quantitatively similar map results, as shown in the simulations with and without ON/OFF channels in Section 6.2.3.
For a large change in pattern shape or size, such as using natural images instead of Gaussian patterns, two parameters need to be adjusted. First, the input scale or threshold needs to be changed to get results as similar to the original working simulation as possible. Second, the input density scale sd needs to be adjusted to compensate for the remaining differences in the amount of input per iteration. Of course, because the system is nonlinear, it is not always possible to compensate completely.
As an example, if Gaussian input patterns are replaced with large, sharp-edged squares, each input will produce multiple activity bubbles in V1 instead of one bubble. The input scale γA should be set to a value that results in bubbles about the same size as in the Gaussian simulation, and sd should be set to the average number of bubbles per iteration in the new simulation. For input patterns with large, spread-out areas of activity, the lateral interaction strengths γE and γI can also be increased to ensure that distinct activity bubbles form. Other parameters do not usually need to be changed when changing the input patterns.
A.3 Choosing Parameters for New Simulations |
421 |
rA |
sg L d |
sg L d + 2(rA − 0.5) |
(a) Computing the LGN size L
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rL |
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Rd r |
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L d A |
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sg Rd |
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Ld |
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(a) Computing the retina size R
Fig. A.1. Mapping between neural sheets in LISSOM. In computing the LGN size L and the retina size R (Table A.2), a buffer area is added around the lower level sheet so that all neurons at the higher level have complete receptive fields. (a) In the mapping from LGN to V1, the outer square represents the LGN sheet, the dashed area maps point-for-point to V1, and the circle represents the receptive field of the top left V1 neuron. For instance, if sg = 8 and Ld = 24, the dashed line encloses an area of 192 × 192 LGN units (8 × 24 = 192). This area is extended on all sides by rA −0.5 units to make sure that all V1 neurons have complete receptive fields. Thus, the LGN contains 204 × 204 neurons in total (192 + 2 × 6 = 204). (b) The mapping from retina to V1 is formed analogously, by extending the buffering down one more level. The outer square represents the retina, the dotted area maps point-for-point to the LGN, and the dashed area maps point-for-point to V1. The circle on the right shows the receptive field of the top right LGN neuron and the circle on the left represents the receptive field of the top left V1 neuron, with its radius expressed in retinal units (hence the factor Rd/Ld). For example, if Rd = 48, the dashed area is sgRd = 8×48 = 384 retinal units wide and the dotted area 384+2×4824 ×6 = 408 retinal units wide. For an LGN radius of rL = 16.5, the full retina therefore consists of 440 × 440 neurons (384 + 2 × 4824 × 6 + 2 × 16 = 440).
The connection weights in each RF are usually initialized to uniformly distributed positive random values; sometimes a Gaussian pattern is used instead to speed up the self-organization. The range [0..1] was used for these values in all simulations; note however that the range does not matter because the weights in each group of RFs (afferent, lateral excitatory, and lateral inhibitory) are immediately normalized to sum to 1.0.
Sections A.4–A.9 describe the specific differences from the default parameters for each of the LISSOM simulations in this book. To determine the actual parameter values, one can begin with a copy of Tables A.2 and A.3, make the specified changes, and then calculate the new values for each parameter starting at the top of each table. For instance, for a simulation that changes θli to 0.05, parameter θui would become 0.6 instead of 0.65 (Table A.2). The same method can be used to determine a consistent set of parameter values for any new simulations.
422 A LISSOM Simulation Specifications
A.4 Retinotopic Maps
The retinotopic map simulations in Chapter 4 were based on the default values, except the training Gaussians were unoriented (σa = σb = σu = 3), the sigmoid threshold was lower (θli = 0.035) so that the smaller patterns would produce as much cortical activity as oriented Gaussians do, the RF centers were randomly scattered by 5% in both the x and y directions, and the lateral connections were uniformly random instead of initialized to a Gaussian profile.
The default parameter values result in two Gaussian inputs per iteration. While in principle it would be possible to self-organize the network with patterns that consist of more than two, given that the Gaussians are relatively large compared with the retina, it would be difficult to distribute them uniformly on the retina while enforcing a minimum distance dr between them. After two Gaussians have been placed randomly, the only remaining possible locations are often near the corners. The corners would therefore be trained more often than the other areas of the map, resulting in a distorted organization. More Gaussians could be used if a larger retinal and cortical area was simulated (i.e. with a larger sg), or the Gaussians were narrower.
A.5 Orientation Maps
The LISSOM simulations with oriented Gaussians in Sections 5.3 and 6.2 (Figures 5.5–5.12, “Gaussians” in 5.13, “ON/OFF” in 6.4 and 6.5) were based on the default parameter values, except the inhibitory connection death threshold wd was increased to 6wdorI2o/rI2 for historical reasons. These values were also used in the “Plus/Minus” simulation in Figure 5.13, except the sign of the input scale sb was chosen randomly for each pattern. The other types of orientation map simulations presented in that figure are described in the following subsections.
A.5.1 Disks, Noisy Disks, and Noise
The simulations with circular disks (“Disks” in Figure 5.13) were based on the default LISSOM parameters, except only one disk was drawn per iteration (np = 1), the lateral interactions were stronger to allow long contours to be separated into distinct activity bubbles (γI = 2.0, γE = 1.2), and the LGN afferent scale was slightly stronger (γL = 3/sb) because each activity bubble was slightly weaker than in the oriented Gaussian simulation.
The disk input patterns were fully activated in the circular region (rd = 25sw) around their centers, and the activation then fell off according to a Gaussian pattern (σd = 3). Each input center was separated far enough so that input patterns would never overlap (dr = 1.5rd). Because the disk stimuli are large compared with Ld, the area in which disk centers are chosen was increased so that even the neurons at the edges of the map are equally likely to receive input from all parts of the disks. Thus, the number of inputs per eye was corrected to reflect this larger area (np =
max 1, |
2rd+Ldsg |
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2rd+Ld |
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A.6 Ocular Dominance Maps |
423 |
The simulations with noisy circular disks (“Noisy disks” in Figure 5.13) were identical to the simulations with noiseless disks, except uniform random noise in the range [−0.5..0.5) was added to the input pattern. The simulations with uniform random noise alone (“Noise” in Figure 5.13) were based on the noisy disk simulations, except no disks were drawn, the LGN parameters were adjusted to produce stronger LGN activations (σc = 0.75, σs = 3σc, and γL = 5/sb), and the V1 parameters were changed to produced stronger V1 activations (γE = 0.9, γI = 0.9, γA = 3).
A.5.2 Natural Images
The orientation map simulations trained on natural images (“Nature” in Figure 5.13) were based on the default LISSOM parameters, except the retinal density was doubled to provide more image resolution (Rd = 48), only one image was drawn per iteration (np = 1), the input density scale was higher (sd = 4) because each input resulted in about four activity bubbles on average, the number of iterations was fixed to the default value (st = 0.5) instead of being adjusted automatically for the input density scale, the LGN afferent scale was increased to produce activity for lowcontrast inputs (γL = 4.7/sb), and the sigmoid threshold was lower (θli = 0.076) to allow responses to low-contrast stimuli.
The lateral interaction strengths and learning rates were also adjusted during training rather than being fixed. In the default simulation, the lateral inhibitory weights self-organize into only a few small regions, because Gaussian patterns have no long-range correlations. In contrast, natural images have significant long-range correlations, and inhibitory weights spread over a larger area. To keep the balance between excitatory and inhibitory lateral weights approximately constant, the lateral inhibitory strength was set to γI = 1.75 at first, increased to 2.2 at iteration 1000st and 2.6 at iteration 2000st. The lateral inhibitory learning rate αI was
set to α |
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0.00005rI2o |
at first, increased to α |
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0.00010rI2o |
at iteration 1000s |
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stsdrI2 |
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stsdrI2 |
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α |
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0.00015rI2o |
at iteration 2000s |
, and α |
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0.00025rI2o |
at iteration 5000s |
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I |
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stsdrI2 |
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stsdrI2 |
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these modifications, the orientation map self-organizes robustly with natural image input.
A.6 Ocular Dominance Maps
The ocular dominance simulations in Section 5.4 (Figures 5.15–5.17, “Dimming” in Figure 5.19) were based on the default LISSOM parameters, except four LGN regions (nA = 4) and two eyes were included, the sigmoid threshold was lower (θli = 0.035) so that the smaller patterns would produce as much cortical activity as oriented Gaussians do, the training Gaussians were unoriented (σu = 4.25), and the input brightness scale sb was randomly chosen from the range [0..1] for the left eye and calculated as 1 − sb for the right eye for each input.
The strabismic ocular dominance simulations (Figure 5.18, “Strabismic” in Figure 5.19) were otherwise similar, except the inputs were scattered randomly between
424 A LISSOM Simulation Specifications
the eyes (ss = 1.0). The “Mild” ocular dominance simulations in Figure 5.19 were identical to “Strabismic”, except the input scale was constant (sb = 1.0), and the inputs were scattered slightly between the eyes (ss = 0.2). The “Moderate” ocular dominance simulations were identical to “Mild”, except the inputs were scattered more between the eyes (ss = 0.4).
A.7 Direction Maps
The direction map simulations in Section 5.5 (Figures 5.21–5.24, “Speed 1” in Figure 5.25) were based on the default LISSOM parameters, except eight LGN regions were included (nA = 8), only a single pattern was presented per iteration (sd = 1) to avoid overlapping patterns that are moving in different directions, and the LGN afferent scale was slightly stronger (γL = 2.38/sb) because the response is weaker when inputs are not perfectly aligned in each eye. Because sd = 1, st = 0.5, which means that these simulations were run for 20,000 iterations with one pattern apiece rather than the default 10,000 with two patterns apiece. As long as input patterns can be placed randomly on the retina without overlap, and the responses are localized in V1 (as they usually are), using more patterns in fewer iterations is equivalent to using fewer patterns in more iterations.
The simulations with different speeds (Figure 5.25) were otherwise identical except the LGN afferent scale was adjusted for each speed to keep the average response comparable: γL = 2.33/sb for speed 0, γL = 2.38/sb for speed 1, γL = 2.53/sb for speed 2, and γL = 2.80/sb for speed 3.
A.8 Combined Orientation / Ocular Dominance Maps
The combined orientation and ocular dominance simulations in Section 5.6.2 (Figures 5.27 and 5.28) were based on the “Gaussians” orientation-only simulation, except four LGN regions (nA = 4) and two eyes were included. For each input, the input scale sb was chosen randomly for the left eye and calculated as 1 − sb for the right eye.
A.9 Combined Orientation / Ocular Dominance / Direction Maps
In Section 5.6, the Gaussian-trained combined orientation, ocular dominance, and direction simulations (Figure 5.29, “Gaussians” in Figure 5.32) were based on the “Gaussians” orientation-only simulation, except 16 LGN regions (nA = 16) and two eyes were included, only a single pattern was presented per iteration (sd = 1) to avoid overlapping patterns that are moving in different directions, and the LGN afferent scale was adjusted to match the value computed for the direction-only map (γL = 2.38/sb). For each input, the input scale sb was chosen randomly for the left
A.9 Combined Orientation / Ocular Dominance / Direction Maps |
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eye and calculated as 1 − sb for the right eye. The speed of the moving patterns was 1, i.e. subsequent frames differed by 1.0 retinal units.
The combined simulation with noisy disks (“Noisy disks” in Figure 5.32), used the inputs from the “Noisy Disks” orientation-only simulation, drawn at speed 2. The parameters were the same as for the “Gaussians” combined OR/OD/DR simulation, except a higher input density scaling was used (sd = 2) because the inputs more regularly activated the cortex, γI and αI were adjusted as for the orientation-only simulation with natural images (Section A.5.2) to keep excitation and inhibition balanced, and the input scaling for the LGN was increased (γL = 3.0/sb) to keep the LGN responses comparable.
The combined model trained with natural images (Figure 5.31, “Nature in Figure 5.32) was identical to the orientation-only natural image simulation in Section A.5.2 except 16 LGN regions (nA = 16) and two eyes were included, the input density scale was increased without changing the number of iterations (sd = 2 and st = 1), and the LGN afferent scale was adjusted to match the value computed for the direction-only map (γL = 5.6/sb). For each input, the input scale sb was chosen randomly for the left eye and calculated as 1 − sb for the right eye. The speed of the moving patterns was 2.0.
B
Reduced LISSOM Simulation Specifications
As described in Section 6.2.3, as long as the inputs to the model consist of abstract patterns instead of natural images, simulations that utilize only the ON channel (or bypass the LGN entirely) lead to the same results as simulations that include both the ON and OFF channels of the LGN explicitly. Such reduced LISSOM networks allow demonstrating many phenomena efficiently and clearly, as was seen in Chapters 6, 7 and 11–15.
The reduced LISSOM simulations were based on the default LISSOM parameter values, except there was only one input sheet (instead of ON and OFF; thus nA = 1). This sheet was mapped directly to the cortex like the LGN sheets (Figure A.1), and sized like the LGN sheets (R = L). The most important adjustment is to set the sigmoid thresholds so that the total cortical response is the same (on average) as with ON and OFF channels, to compensate for the higher average value of retinal activity compared with the LGN activity. In the reduced LISSOM simulations, the input threshold was therefore set to θli = 0.1. These values are taken as the default reduced LISSOM parameters, occasionally overridden in individual simulations as specified in the following sections and in Appendices D and F.
B.1 Plasticity
The retinal lesion experiments in Section 6.3 were run with the default reduced LISSOM parameters, except the inhibitory connection death threshold wd was increased to 6wdorI2o/rI2 for historical reasons, and the afferent learning rate αA was increased to 0.003 after iteration 10,000; all other parameters remained the same as at the end of the self-organization. Faster learning makes the changes more visible, and also models the increased plasticity that might result during recovery from injury (Emery, Royo, Fischer, Saatman, and McIntosh 2003; Kaas 2001a,b). The cortical lesion experiments in Section 6.4 were based on the same parameters as the retinal lesion experiments, except the afferent learning rate remained at its default value even after the end of self-organization.
428 B Reduced LISSOM Simulation Specifications
B.2 Tilt Aftereffect
The tilt aftereffect experiments in Chapter 7 were based on the default reduced LISSOM parameters, except for historical reasons the cortical density was slightly larger (Nd = 192), the final excitatory radius rEf was reduced to 1.5, the inhibitory connection death threshold wd was decreased to 0.00005, and only a single pattern was presented per iteration (sd = 1). Also for historical reasons, the input patterns were blurred with a uniform 3 × 3 convolution kernel before they were presented to the network.
For the average tilt aftereffect plots in Figures 7.5 and 7.6, each input line was presented at nine different locations chosen from the nodes of a regular 3 × 3 grid centered in the retina, with each grid step four retinal units wide.
B.3 Scaling
The scaling equation and GLISSOM simulations in Chapter 15 were based on the tilt aftereffect parameters (Section B.2), scaled as described in each section of Chapter 15. For instance, the area scaling simulation (Figure 15.1) compares two sim-
ulations with Ld = 24 and Nd = 44, differing by their area scales (sg = 1 and sg = 4).
The scaling simulations were run on a 600 MHz Intel Pentium III Linux machine with 1024 MB of RAM. The timing results are user CPU times reported by the GNU “time” command; the CPU time is essentially the same as the elapsed wallclock time because the CPU utilization was always over 99%.
