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Veronika Koren, Sophie Denève
Computational Account of Spontaneous Activity as a Signature of Predictive Coding
www.ploscompbiol.org
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-17.3308 -2.7155 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 329.8393 439.37 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 331.8803 441.8645 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
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[()]TJ
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0.7313 0 Td
()Tj
0.669 0 Td
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1.0488 1.3549 Td
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5.9998 0 0 5.9998 366.6897 453.1464 cm
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(1)Tj
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-0.7937 -3.3638 Td
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Q
q
1 j
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10 0 0 10 374.8535 441.8645 cm
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1 TL
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[(\n)]TJ
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[()]TJ
ET
Q
q
1 j
1 J
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5.9998 0 0 5.9998 387.4393 446.0031 cm
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/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 389.5937 441.8645 cm
BT
/F15 1 Tf
1 TL
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()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 396.7937 439.37 cm
BT
/F12 1 Tf
1 TL
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[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 398.8346 441.8645 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.9524 0 Td
( )Tj
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0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 421.7385 446.0031 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 423.8362 441.8645 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
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[()14()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 436.2519 446.0031 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 563.4708 441.8645 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
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(3)Tj
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()Tj
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1.2188 0 Td
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2.4265 0 Td
[()]TJ
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(\()Tj
/F12 1 Tf
[()]TJ
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(\),)Tj
1.7971 0 Td
(we)Tj
1.3437 0 Td
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1.6554 0 Td
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5.8733 0 Td
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0 Tc
1.0318 0 Td
[()]TJ
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0.754 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
0.669 0 Td
()Tj
/F16 1 Tf
1.0488 0.7994 Td
(R)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 467.6598 350.4188 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(1)Tj
/F13 1 Tf
-0.2362 -1.4929 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 475.8803 344.7496 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[(\n)]TJ
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 488.4661 348.3779 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 490.5637 344.7496 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.6724 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 520.6677 348.3779 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 522.7653 344.7496 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()14()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 535.1244 348.3779 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 537.2787 344.7496 cm
BT
/F6 1 Tf
1 TL
0.005 Tc
0 0 Td
(,)Tj
0.4535 0 Td
(which)Tj
2.6929 0 Td
(is)Tj
-36.8728 -1.3946 Td
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4.4391 0 Td
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1.0431 0 Td
(the)Tj
1.4967 0 Td
(expression)Tj
4.5581 0 Td
(for)Tj
/F12 1 Tf
1.4116 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 334.2047 329.159 cm
BT
/F12 1 Tf
1 TL
0.0067 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 336.2456 330.8031 cm
BT
/F6 1 Tf
1 TL
0.005 Tc
0 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
/F6 1 Tf
(\).)Tj
1.4569 0 Td
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1.8085 0 Td
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2.4265 0 Td
[()]TJ
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(\()Tj
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[()]TJ
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1.5647 0 Td
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1.7575 0 Td
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1.4966 0 Td
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2.5966 0 Td
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(\()Tj
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[()]TJ
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1.7007 0 Td
(therefore)Tj
3.9345 0 Td
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3.0331 0 Td
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-35.3988 -1.2982 Td
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1.4967 0 Td
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2.2904 0 Td
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5.1023 0 Td
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1.5307 0 Td
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1.0204 0 Td
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1.474 0 Td
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3.3562 0 Td
(x)Tj
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(\()Tj
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[()]TJ
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(\))Tj
1.6725 0 Td
(=)Tj
0.7993 0 Td
([)Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 527.9811 303.137 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 531.5527 304.8377 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
/F6 1 Tf
(\),)Tj
/F12 1 Tf
-33.1538 -1.3039 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 204.6614 290.1543 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 208.233 291.7984 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
/F6 1 Tf
(\),)Tj
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0 Tc
1.4229 0 Td
(.)Tj
0.4309 0 Td
(.)Tj
0.4365 0 Td
(.)Tj
/F6 1 Tf
-0.004 Tc
(,)Tj
/F12 1 Tf
0.7144 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 242.9291 290.1543 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 245.2535 291.7984 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
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(\()Tj
/F12 1 Tf
[()]TJ
/F6 1 Tf
(\)])Tj
1.5363 0 Td
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1.0148 0 Td
(the)Tj
1.4684 0 Td
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1.4683 0 Td
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0 Tc
3.3846 0 Td
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[()]TJ
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0.55 0 Td
()Tj
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[()]TJ
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0.7313 0 Td
()Tj
0.6576 0 Td
()Tj
1.0488 0 Td
()Tj
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0.2892 0.017 Td
(^)Tj
/F12 1 Tf
-0.017 -0.017 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 424.6866 289.3039 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 428.2015 291.7984 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
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[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
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(;)Tj
/F13 1 Tf
0.8447 0.017 Td
(^)Tj
/F12 1 Tf
-0.017 -0.017 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 449.1212 289.3039 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 452.6362 291.7984 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
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(;)Tj
0.8221 0 Td
(:::)Tj
0.7993 0 Td
(;)Tj
/F13 1 Tf
0.4592 0.017 Td
(^)Tj
/F12 1 Tf
-0.0226 -0.017 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 485.8582 289.3039 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 488.6929 291.7984 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
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[()]TJ
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0.7256 0 Td
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0.8787 0 Td
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1.0148 0 Td
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1.4684 0 Td
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3.5433 0 Td
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1.474 0 Td
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3.8778 0 Td
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1.3039 0 Td
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3.4696 0 Td
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0.9071 0 Td
(=)Tj
0.7993 0 Td
([)Tj
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[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 323.7732 263.3385 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(11)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 330.8598 264.9826 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
/F12 1 Tf
0.4478 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 341.9716 263.3385 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(12)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 349.0582 264.9826 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
/F13 1 Tf
0 Tc
0.4478 0 Td
(.)Tj
0.4309 0 Td
(.)Tj
0.4365 0 Td
(.)Tj
/F12 1 Tf
-0.004 Tc
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 371.5653 263.3385 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(1)Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 380.296 264.9826 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
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(;)Tj
/F13 1 Tf
0 Tc
0.4478 0 Td
(.)Tj
0.4309 0 Td
(.)Tj
0.4365 0 Td
(.)Tj
/F6 1 Tf
-0.004 Tc
(;)Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 405.0141 263.3385 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
/F6 1 Tf
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 410.9102 264.9826 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
/F12 1 Tf
0.4422 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 421.9653 263.3385 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
/F6 1 Tf
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 427.8614 264.9826 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
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(,)Tj
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0 Tc
0.4422 0 Td
(.)Tj
0.4365 0 Td
(.)Tj
0.4308 0 Td
(.)Tj
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-0.004 Tc
[()]TJ
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Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 450.3118 263.3385 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 457.8519 264.9826 cm
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1 TL
-0.004 Tc
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(where)Tj
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2.6532 0 Td
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2.0806 0 Td
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2.0579 0 Td
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10 0 0 10 423.7228 496.97 cm
BT
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1 TL
0 0 Td
(2)Tj
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0.941 0.6859 Td
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13.0337 0 Td
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(6)Tj
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[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 339.8173 447.7606 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 341.9716 450.3118 cm
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1 TL
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[()]TJ
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0.7256 0 Td
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1 J
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[()]TJ
0 -1.1527 Td
[()]TJ
ET
Q
q
1 j
1 J
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BT
/F15 1 Tf
1 TL
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()Tj
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[()]TJ
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0.8844 0 Td
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[()]TJ
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0 Tc
9.4223 -1.9049 Td
(_)Tj
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-0.2607 -0.2154 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 299.622 326.0976 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 301.7196 328.5921 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6576 0 Td
()Tj
1.0432 0 Td
( )Tj
/F18 1 Tf
0.771 0 Td
(l)Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 345.4299 326.0976 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 347.5842 328.5921 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6009 0 Td
()Tj
/F16 1 Tf
0.9921 0.9467 Td
(X)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 377.2346 318.1039 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 386.8724 328.5921 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 393.5622 326.0976 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()12()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 397.1905 328.5921 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 400.5354 326.0976 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 402.5763 328.5921 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6009 0 Td
( )Tj
/F16 1 Tf
0.9978 0.9467 Td
(X)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 431.6031 318.1039 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[(\r)]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 441.8645 328.5921 cm
BT
/F20 1 Tf
1 TL
0 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 447.8173 326.0976 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()4(\r)]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 452.7496 328.5921 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 457.3984 326.0976 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[(\r)]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 460.7433 328.5921 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
9.547 0 Td
()Tj
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(8)Tj
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()Tj
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-0.004 Tc
-36.3455 -3.2711 Td
(where)Tj
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2.6476 0 Td
[()]TJ
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(=)Tj
/F11 1 Tf
0.7994 0 Td
(w)Tj
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 249.0519 300.359 cm
BT
/F11 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(T)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 256.0251 295.8803 cm
BT
/F11 1 Tf
1 TL
-0.004 Tc
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(w)Tj
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0.907 0 Td
(is)Tj
0.8447 0 Td
(the)Tj
1.4684 0 Td
(matrix)Tj
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(lateral)Tj
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(and)Tj
1.7348 0 Td
(recurrent)Tj
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(Note)Tj
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(eq)Tj
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1.8198 0 Td
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(\(autapse\))Tj
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(better)Tj
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(how)Tj
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(the)Tj
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(a)Tj
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(neuron)Tj
-33.6299 -1.2983 Td
(tracking)Tj
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(one)Tj
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(\(auto-encoder\).)Tj
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(neuron)Tj
3.1464 0 Td
(a)Tj
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[()]TJ
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(predictive)Tj
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(corresponds)Tj
5.1362 0 Td
(to)Tj
1.0262 0 Td
(a)Tj
0.6519 0 Td
(simple)Tj
2.8573 0 Td
(reset)Tj
2.0977 0 Td
(mechanism,)Tj
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(computationally)Tj
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1.0261 0 Td
(self)Tj
1.5364 0 Td
(connec-)Tj
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1.9956 0 Td
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(-1.)Tj
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(membrane)Tj
4.5637 0 Td
(potential)Tj
3.7588 0 Td
(is)Tj
0.8503 0 Td
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(to)Tj
1.0261 0 Td
(the)Tj
1.4683 0 Td
(prediction)Tj
4.3767 0 Td
(error,)Tj
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0 Tc
-30.1717 -1.4683 Td
[()]TJ
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0.7484 0 Td
()Tj
/F12 1 Tf
[()]TJ
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0.7256 0 Td
()Tj
0.6633 0 Td
()Tj
/F12 1 Tf
1.0488 0 Td
[()]TJ
/F15 1 Tf
0.4876 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.601 0 Td
( )Tj
/F13 1 Tf
1.0147 0.017 Td
(^)Tj
/F12 1 Tf
-0.017 -0.017 Td
[()]TJ
/F15 1 Tf
0.5385 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7257 0 Td
()Tj
/F6 1 Tf
-0.004 Tc
(.)Tj
0.8334 0 Td
(Whenever)Tj
4.3596 0 Td
(it)Tj
0.7824 0 Td
(crosses)Tj
3.0274 0 Td
(the)Tj
1.474 0 Td
(threshold)Tj
4.0082 0 Td
(\()Tj
/F14 1 Tf
0 Tc
[()]TJ
/F12 1 Tf
[()18( )20()8()]TJ
/F15 1 Tf
2.8289 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
456.3213 205.9654 m
463.2945 205.9654 l
463.2945 205.5685 l
456.3213 205.5685 l
f*
5.9998 0 0 5.9998 456.3212 206.8724 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 460.2897 209.0834 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 458.3055 199.8425 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 466.0724 203.244 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
476.5606 205.9654 m
479.5654 205.9654 l
479.5654 205.5685 l
476.5606 205.5685 l
f*
5.9998 0 0 5.9998 476.5606 206.8724 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
0 -1.1716 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 479.5653 203.244 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\),)Tj
0.788 0 Td
(the)Tj
1.4683 0 Td
(neuron)Tj
3.1465 0 Td
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1.9332 0 Td
(a)Tj
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(At)Tj
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(this)Tj
1.6724 0 Td
(time,)Tj
2.2564 0 Td
(the)Tj
1.4683 0 Td
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(of)Tj
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(the)Tj
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(and)Tj
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[()]TJ
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[()]TJ
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(=)Tj
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0.7994 0 Td
[()]TJ
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(1.)Tj
1.491 0 Td
(Notice)Tj
2.8743 0 Td
(that)Tj
1.7802 0 Td
(there)Tj
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(no)Tj
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4.5807 0 Td
(the)Tj
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(artificially.)Tj
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(one)Tj
1.6951 0 Td
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(example)Tj
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(to)Tj
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(the)Tj
1.474 0 Td
(firing)Tj
2.4264 0 Td
(rule)Tj
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0 Tc
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[()]TJ
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0.7483 0 Td
()Tj
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[()]TJ
/F15 1 Tf
0.7257 0 Td
()Tj
/F20 1 Tf
0.6689 0 Td
(>)Tj
ET
Q
q
1 j
1 J
0 w
231.9307 152.9008 m
234.9354 152.9008 l
234.9354 152.5039 l
231.9307 152.5039 l
f*
5.9998 0 0 5.9998 231.9307 153.8645 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
0 -1.1811 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 234.9354 150.2362 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
0.4422 0 Td
(where)Tj
2.6532 0 Td
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q
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[()]TJ
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BT
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q
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[()]TJ
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(\))Tj
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(=)Tj
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[()]TJ
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Q
q
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1 J
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7.4999 0 0 6.6659 317.8771 365.5559 cm
BT
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1 TL
-0.0053 Tc
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(1)Tj
ET
Q
q
1 j
1 J
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10 0 0 10 321.4488 367.2566 cm
BT
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(\()Tj
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[()]TJ
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(\),)Tj
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1.4229 0 Td
[()]TJ
ET
Q
q
1 j
1 J
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7.4999 0 0 6.6659 342.3118 365.5559 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
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(2)Tj
ET
Q
q
1 j
1 J
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10 0 0 10 345.8834 367.2566 cm
BT
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[()]TJ
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0.4365 0 Td
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[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 382.5637 365.5559 cm
BT
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1 TL
-0.0053 Tc
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[()]TJ
ET
Q
q
1 j
1 J
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10 0 0 10 387.7795 367.2566 cm
BT
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[()]TJ
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(=)Tj
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[( )]TJ
ET
Q
q
1 j
1 J
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7.4999 0 0 6.6659 489.9968 352.5732 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
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(1)Tj
ET
Q
q
1 j
1 J
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10 0 0 10 493.5118 354.274 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
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/F12 1 Tf
[()]TJ
/F6 1 Tf
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1.4229 0 Td
[( )]TJ
ET
Q
q
1 j
1 J
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7.4999 0 0 6.6659 511.37 352.5732 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
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(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 514.9417 354.274 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
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[()]TJ
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0.4309 0 Td
(.)Tj
0.4365 0 Td
(.)Tj
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[( )]TJ
ET
Q
q
1 j
1 J
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7.4999 0 0 6.6659 548.6173 352.5732 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
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[()]TJ
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Q
q
1 j
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10 0 0 10 553.833 354.274 cm
BT
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1 TL
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[()]TJ
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(\()Tj
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[()]TJ
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(\))Tj
1.7121 0 Td
(=)Tj
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0.8051 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 229.833 339.5905 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
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(1)Tj
ET
Q
q
1 j
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10 0 0 10 233.348 341.2346 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
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[()]TJ
/F6 1 Tf
(\),)Tj
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1.4229 0 Td
[()]TJ
ET
Q
q
1 j
1 J
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7.4999 0 0 6.6659 252.2834 339.5905 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
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(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 255.7984 341.2346 cm
BT
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[()]TJ
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[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 290.5511 339.5905 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
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1 j
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10 0 0 10 295.7669 341.2346 cm
BT
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()Tj
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[()]TJ
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1.44 0 Td
()Tj
0.6576 0 Td
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1.0488 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 353.7637 325.9275 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 358.2425 321.7889 cm
BT
/F15 1 Tf
1 TL
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[()]TJ
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0.8844 0 Td
()Tj
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[()]TJ
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0.7313 0 Td
()Tj
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( )Tj
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1.0205 0 Td
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[()]TJ
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0.5443 0 Td
()Tj
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[()]TJ
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()Tj
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(m)Tj
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[()]TJ
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13.5722 0 Td
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(11)Tj
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()Tj
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()Tj
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( )Tj
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[()]TJ
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()Tj
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[()]TJ
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[()]TJ
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14.0314 0 Td
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(12)Tj
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()Tj
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()Tj
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()Tj
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[()]TJ
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[( )]TJ
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[()]TJ
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14.0654 0 Td
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(13)Tj
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7.8008 -1.7971 Td
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()Tj
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13.5552 0 Td
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(14)Tj
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[()]TJ
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()Tj
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1.7688 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 297.7511 142.4125 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 302.2299 138.7842 cm
BT
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1 TL
0 0 Td
()Tj
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[()]TJ
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()Tj
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1.1565 0.2607 Td
(_)Tj
-0.1417 -0.2551 Td
(^)Tj
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[()]TJ
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0.6009 0 Td
()Tj
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[()]TJ
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()Tj
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( )Tj
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(m)Tj
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0.6407 0.0114 Td
(_)Tj
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-0.0851 -0.017 Td
[()]TJ
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0.4422 0 Td
()Tj
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(n)Tj
/F13 1 Tf
-0.034 -1.3662 Td
(2)Tj
/F15 1 Tf
13.7989 0.6859 Td
()Tj
/F6 1 Tf
(17)Tj
/F15 1 Tf
()Tj
/F6 1 Tf
-0.004 Tc
-35.8693 -3.1974 Td
(The)Tj
1.7802 0 Td
(effect)Tj
2.364 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4741 0 Td
(linear)Tj
2.5058 0 Td
(cost)Tj
/F12 1 Tf
1.8141 0 Td
[()]TJ
/F6 1 Tf
0.6633 0 Td
(is)Tj
0.8447 0 Td
(to)Tj
1.0262 0 Td
(raise)Tj
2.0806 0 Td
(thresholds)Tj
4.371 0 Td
(of)Tj
1.0148 0 Td
(all)Tj
1.1452 0 Td
(neurons)Tj
3.5091 0 Td
(for)Tj
1.3833 0 Td
(the)Tj
1.4684 0 Td
(same)Tj
2.262 0 Td
(amount)Tj
3.3505 0 Td
(and)Tj
-34.0721 -1.2983 Td
(therefore)Tj
3.8551 0 Td
(to)Tj
1.0205 0 Td
(penalize)Tj
3.4979 0 Td
(high)Tj
2.0069 0 Td
(firing)Tj
2.4265 0 Td
(rates)Tj
2.1089 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(population)Tj
4.5921 0 Td
(\()Tj
0.83 0.64 0.02 0 k
(eq)Tj
1.4909 0 Td
(17)Tj
0 g
(\).)Tj
1.7348 0 Td
(The)Tj
1.7801 0 Td
(quadratic)Tj
4.0139 0 Td
(cost)Tj
/F12 1 Tf
1.8084 0 Td
[()]TJ
/F6 1 Tf
0.7257 0 Td
(has)Tj
1.542 0 Td
(the)Tj
-35.0869 -1.3039 Td
(same)Tj
2.2507 0 Td
(network-wide)Tj
5.794 0 Td
(effect)Tj
2.3584 0 Td
(on)Tj
1.2699 0 Td
(thresholds)Tj
4.3653 0 Td
(\()Tj
0.83 0.64 0.02 0 k
(eq)Tj
1.4854 0 Td
(17)Tj
0 g
(\))Tj
1.5023 0 Td
(and)Tj
1.7292 0 Td
(in)Tj
1.0261 0 Td
(addition)Tj
3.5942 0 Td
(adds)Tj
2.0636 0 Td
(a)Tj
0.6463 0 Td
(hyperpolarizing)Tj
6.5423 0 Td
(current)Tj
-34.6277 -1.2983 Td
(to)Tj
1.0262 0 Td
(the)Tj
1.4683 0 Td
(membrane)Tj
4.5637 0 Td
(potential)Tj
3.7644 0 Td
(\()Tj
0.83 0.64 0.02 0 k
(eq)Tj
1.491 0 Td
(16)Tj
0 g
(\),)Tj
1.7348 0 Td
(only)Tj
1.9786 0 Td
(to)Tj
1.0205 0 Td
(the)Tj
1.474 0 Td
(neuron)Tj
3.1464 0 Td
(that)Tj
1.7744 0 Td
(recently)Tj
3.4129 0 Td
(spiked.)Tj
3.0387 0 Td
(We)Tj
1.6101 0 Td
(interpret)Tj
3.736 0 Td
(the)Tj
-35.24 -1.2982 Td
(latter)Tj
2.2904 0 Td
(as)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(spike-triggered)Tj
6.2532 0 Td
(adaptation.)Tj
4.7112 0 Td
(Finally,)Tj
3.1691 0 Td
(a)Tj
0.652 0 Td
(noise)Tj
2.33 0 Td
(term)Tj
2.1316 0 Td
(is)Tj
0.8447 0 Td
(added,)Tj
2.8856 0 Td
(which)Tj
2.6532 0 Td
(is)Tj
0.8448 0 Td
(a)Tj
0.6576 0 Td
(white)Tj
2.4208 0 Td
(noise)Tj
-34.3273 -1.304 Td
(with)Tj
1.9956 0 Td
(zero)Tj
1.9332 0 Td
(mean)Tj
2.4491 0 Td
(and)Tj
1.7292 0 Td
(standard)Tj
3.7247 0 Td
(deviation)Tj
/F12 1 Tf
3.9458 0 Td
[()]TJ
/F6 1 Tf
(.)Tj
0.9127 0 Td
(Noise)Tj
2.5115 0 Td
(processes)Tj
3.9684 0 Td
(across)Tj
2.6815 0 Td
(neurons)Tj
3.5036 0 Td
(are)Tj
1.4514 0 Td
(uncorrelated,)Tj
5.5445 0 Td
(i.e.)Tj
/F20 1 Tf
0 Tc
-36.3512 -1.4626 Td
(<)Tj
/F17 1 Tf
-0.004 Tc
0.9921 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 214.8661 446.0598 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 216.8503 447.7039 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
/F6 1 Tf
(\))Tj
/F17 1 Tf
()Tj
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 231.6472 446.0598 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 233.5181 447.7039 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0 Tc
0.6406 0.3628 Td
(0)Tj
/F6 1 Tf
-0.004 Tc
0.3231 -0.3628 Td
(\))Tj
/F20 1 Tf
0 Tc
0.5556 0 Td
(>)Tj
/F6 1 Tf
-0.004 Tc
0.9921 0 Td
(=)Tj
/F17 1 Tf
0.7994 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 271.2755 446.0598 cm
BT
/F12 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()12()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 275.1874 447.7039 cm
BT
/F17 1 Tf
1 TL
-0.004 Tc
0 0 Td
()Tj
/F6 1 Tf
(\()Tj
/F12 1 Tf
[()]TJ
/F14 1 Tf
1.3266 0 Td
[()]TJ
/F12 1 Tf
0.788 0 Td
[()]TJ
/F15 1 Tf
0 Tc
0.3005 0.3628 Td
(0)Tj
/F6 1 Tf
-0.004 Tc
0.3174 -0.3628 Td
(\).)Tj
ET
Q
q
1 j
1 J
0 w
11.9999 0 0 11.9999 200.0125 411.9874 cm
BT
/F0 1 Tf
1 TL
-0.0033 Tc
0 0 Td
(2)Tj
0.7653 0 Td
(Spiking)Tj
3.4583 0 Td
(in)Tj
0.9827 0 Td
(active)Tj
2.8016 0 Td
(and)Tj
1.8614 0 Td
(quiescent)Tj
4.4504 0 Td
(state:)Tj
2.6457 0 Td
(A)Tj
0.8835 0 Td
(minimal)Tj
3.6189 0 Td
(model)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 200.0125 394.9795 cm
BT
/F6 1 Tf
1 TL
-0.012 Tc
0 0 Td
(In)Tj
1.0941 0 Td
(general,)Tj
3.2939 0 Td
(costs)Tj
2.1259 0 Td
(on)Tj
1.2586 0 Td
(spiking)Tj
3.0897 0 Td
(are)Tj
1.423 0 Td
(required)Tj
3.5773 0 Td
(to)Tj
1.0092 0 Td
(achieve)Tj
3.118 0 Td
(efficient)Tj
3.3733 0 Td
(coding)Tj
2.9139 0 Td
(in)Tj
1.0205 0 Td
(networks)Tj
3.8438 0 Td
(with)Tj
1.9672 0 Td
(delays)Tj
2.6192 0 Td
(in)Tj
-35.7276 -1.2982 Td
(synaptic)Tj
3.4582 0 Td
(transmission.)Tj
5.4878 0 Td
(In)Tj
1.0942 0 Td
(case)Tj
1.8255 0 Td
(costs)Tj
2.1316 0 Td
(on)Tj
1.2586 0 Td
(spiking)Tj
3.0898 0 Td
(are)Tj
1.4229 0 Td
(not)Tj
1.5534 0 Td
(properly)Tj
3.5603 0 Td
(controlling)Tj
4.5638 0 Td
(the)Tj
1.4456 0 Td
(activity,)Tj
3.2712 0 Td
(we)Tj
-34.1629 -1.2983 Td
(observe)Tj
3.2087 0 Td
(a)Tj
0.6463 0 Td
(peculiar)Tj
3.3676 0 Td
(phenomenon)Tj
5.5105 0 Td
(which)Tj
2.6135 0 Td
(consists)Tj
3.2825 0 Td
(in)Tj
1.0205 0 Td
(fast)Tj
1.5647 0 Td
(oscillation)Tj
4.2406 0 Td
(of)Tj
0.9978 0 Td
(the)Tj
1.4457 0 Td
(read-out)Tj
3.5943 0 Td
(of)Tj
1.0034 0 Td
(neural)Tj
2.7213 0 Td
(activ-)Tj
-35.2174 -1.3039 Td
(ity.)Tj
1.4343 0 Td
(The)Tj
1.7574 0 Td
(fast)Tj
1.5704 0 Td
(oscillation)Tj
4.235 0 Td
(is)Tj
0.8333 0 Td
(caused)Tj
2.8687 0 Td
(by)Tj
1.1678 0 Td
(alternative)Tj
4.3314 0 Td
(spiking)Tj
3.084 0 Td
(of)Tj
0.9978 0 Td
(neurons)Tj
3.4526 0 Td
(with)Tj
1.9672 0 Td
(positive)Tj
3.2485 0 Td
(and)Tj
1.7065 0 Td
(negative)Tj
-32.6549 -1.2983 Td
(selectivity,)Tj
4.2689 0 Td
(which)Tj
2.6079 0 Td
(we)Tj
1.3096 0 Td
(call)Tj
1.542 0 Td
(the)Tj
1.8255 0 Td
(ping-pong)Tj
4.337 0 Td
(effect.)Tj
2.9196 0 Td
(In)Tj
1.0885 0 Td
(case)Tj
1.8312 0 Td
(of)Tj
0.9978 0 Td
(the)Tj
1.4457 0 Td
(network)Tj
3.4979 0 Td
(with)Tj
1.9615 0 Td
(delays,)Tj
2.8403 0 Td
(the)Tj
1.4457 0 Td
(ping-)Tj
-33.9191 -1.2982 Td
(pong)Tj
2.2223 0 Td
(appears)Tj
3.2201 0 Td
(even)Tj
2.041 0 Td
(without)Tj
3.2711 0 Td
(any)Tj
1.6328 0 Td
(noise)Tj
2.2847 0 Td
(and)Tj
1.7064 0 Td
(is)Tj
0.8334 0 Td
(therefore)Tj
3.7814 0 Td
(due)Tj
1.6724 0 Td
(to)Tj
1.0092 0 Td
(transmission)Tj
5.2667 0 Td
(delays.)Tj
2.8346 0 Td
(In)Tj
1.0942 0 Td
(the)Tj
1.4457 0 Td
(network)Tj
-34.316 -1.2983 Td
(without)Tj
3.2711 0 Td
(transmission)Tj
5.2724 0 Td
(delays)Tj
2.6192 0 Td
(and)Tj
1.7065 0 Td
(without)Tj
3.2711 0 Td
(noise,)Tj
2.5058 0 Td
(there)Tj
2.2224 0 Td
(is)Tj
0.8334 0 Td
(no)Tj
1.2585 0 Td
(such)Tj
2.0296 0 Td
(effect,)Tj
2.5342 0 Td
(but)Tj
1.5307 0 Td
(it)Tj
0.7653 0 Td
(appears)Tj
3.2258 0 Td
(if)Tj
0.7597 0 Td
(the)Tj
1.4457 0 Td
(net-)Tj
-35.2514 -1.3039 Td
(work)Tj
2.245 0 Td
(gets)Tj
1.7348 0 Td
(perturbed)Tj
4.1158 0 Td
(by)Tj
1.1679 0 Td
(the)Tj
1.4457 0 Td
(noise.)Tj
2.5058 0 Td
(The)Tj
1.7575 0 Td
(network)Tj
3.4922 0 Td
(that)Tj
1.7461 0 Td
(is)Tj
0.8334 0 Td
(not)Tj
1.5477 0 Td
(regulated)Tj
3.8665 0 Td
(by)Tj
1.1678 0 Td
(costs)Tj
2.1317 0 Td
(on)Tj
1.2585 0 Td
(spiking)Tj
3.0841 0 Td
(is)Tj
0.8334 0 Td
(highly)Tj
-34.9339 -1.2983 Td
(unstable)Tj
3.5035 0 Td
(and)Tj
1.7065 0 Td
(quickly)Tj
3.0841 0 Td
(enters)Tj
2.5965 0 Td
(inefficient)Tj
4.1782 0 Td
(coding)Tj
2.9084 0 Td
(regimes)Tj
3.2995 0 Td
(with)Tj
1.9615 0 Td
(excessively)Tj
4.4391 0 Td
(high)Tj
1.9729 0 Td
(firing)Tj
2.381 0 Td
(rates.)Tj
2.2848 0 Td
(How-)Tj
-34.316 -1.2982 Td
(ever,)Tj
2.0749 0 Td
(in)Tj
1.0205 0 Td
(the)Tj
1.4456 0 Td
(presence)Tj
3.651 0 Td
(of)Tj
0.9978 0 Td
(noise,)Tj
2.5058 0 Td
(not)Tj
1.5477 0 Td
(all)Tj
1.1225 0 Td
(spikes)Tj
2.5966 0 Td
(are)Tj
1.4286 0 Td
(inefficient.)Tj
4.3937 0 Td
(If)Tj
0.8277 0 Td
(there)Tj
2.228 0 Td
(is)Tj
0.8277 0 Td
(a)Tj
0.6463 0 Td
(spike)Tj
2.245 0 Td
(that)Tj
1.7462 0 Td
(is)Tj
0.8334 0 Td
(only)Tj
1.9445 0 Td
(due)Tj
1.6781 0 Td
(to)Tj
-35.7616 -1.304 Td
(the)Tj
1.4456 0 Td
(noise,)Tj
2.5058 0 Td
(another)Tj
3.2712 0 Td
(spike)Tj
2.245 0 Td
(of)Tj
0.9978 0 Td
(a)Tj
0.6463 0 Td
(neuron)Tj
3.1011 0 Td
(with)Tj
1.9672 0 Td
(opposite)Tj
3.549 0 Td
(selectivity)Tj
4.0535 0 Td
(is)Tj
0.8334 0 Td
(in)Tj
1.0204 0 Td
(fact)Tj
1.6271 0 Td
(best)Tj
1.7801 0 Td
(suited)Tj
2.5796 0 Td
(to)Tj
1.0091 0 Td
(quickly)Tj
3.0841 0 Td
(cor-)Tj
-35.7163 -1.2982 Td
(rect)Tj
1.7007 0 Td
(the)Tj
1.4457 0 Td
(coding)Tj
2.914 0 Td
(error.)Tj
2.4434 0 Td
(In)Tj
1.0885 0 Td
(the)Tj
1.4457 0 Td
(present)Tj
3.1124 0 Td
(section)Tj
2.9877 0 Td
(we)Tj
1.3153 0 Td
(illustrate)Tj
3.6113 0 Td
(efficient)Tj
3.3789 0 Td
(and)Tj
1.7064 0 Td
(inefficient)Tj
4.1726 0 Td
(spikes)Tj
2.5965 0 Td
(with)Tj
1.9672 0 Td
(a)Tj
-35.8863 -1.2983 Td
(minimal)Tj
3.5659 0 Td
(model)Tj
2.7042 0 Td
(with)Tj
1.9616 0 Td
(2)Tj
0.6917 0 Td
(neurons)Tj
3.4525 0 Td
(and)Tj
1.7065 0 Td
(the)Tj
1.4456 0 Td
(linear)Tj
2.4605 0 Td
(cost,)Tj
1.9956 0 Td
(where)Tj
2.6078 0 Td
(spikes)Tj
2.6022 0 Td
(can)Tj
1.5988 0 Td
(be)Tj
1.1338 0 Td
(followed)Tj
3.5886 0 Td
(step-by-step.)Tj
-0.004 Tc
-30.3191 -1.3039 Td
(In)Tj
1.1111 0 Td
(the)Tj
1.4684 0 Td
(case)Tj
1.8652 0 Td
(of)Tj
1.0148 0 Td
(a)Tj
0.6519 0 Td
(single)Tj
2.5342 0 Td
(neuron,)Tj
3.3675 0 Td
(Up)Tj
1.474 0 Td
(states)Tj
2.3981 0 Td
(are)Tj
1.4457 0 Td
(not)Tj
1.576 0 Td
(observed.)Tj
4.0195 0 Td
(However,)Tj
4.0762 0 Td
(encoding)Tj
3.9345 0 Td
(a)Tj
0.652 0 Td
(signal)Tj
2.5455 0 Td
(that)Tj
-35.3308 -1.2983 Td
(can)Tj
1.6214 0 Td
(be)Tj
1.1451 0 Td
(both)Tj
2.0523 0 Td
(positive)Tj
3.3109 0 Td
(and)Tj
1.7234 0 Td
(negative)Tj
3.532 0 Td
(with)Tj
1.9899 0 Td
(only)Tj
1.9729 0 Td
(one)Tj
1.6951 0 Td
(neuron)Tj
3.1407 0 Td
(results)Tj
2.8007 0 Td
(in)Tj
1.0318 0 Td
(poor)Tj
2.1203 0 Td
(estimation)Tj
4.439 0 Td
(of)Tj
1.0091 0 Td
(the)Tj
1.4627 0 Td
(signal.)Tj
-35.0473 -1.2982 Td
(A)Tj
0.9014 0 Td
(single)Tj
2.5341 0 Td
(neuron)Tj
3.1465 0 Td
(is)Tj
0.8447 0 Td
(only)Tj
1.9785 0 Td
(able)Tj
1.8312 0 Td
(to)Tj
1.0261 0 Td
(correct)Tj
3.0388 0 Td
(the)Tj
1.4683 0 Td
(estimate)Tj
3.5433 0 Td
(in)Tj
1.0375 0 Td
(the)Tj
1.4683 0 Td
(direction)Tj
3.8551 0 Td
(of)Tj
1.0148 0 Td
(its)Tj
1.1452 0 Td
(weight.)Tj
3.1124 0 Td
(Consider)Tj
3.8891 0 Td
(the)Tj
-35.8353 -1.3039 Td
(case)Tj
1.8595 0 Td
(where)Tj
2.6532 0 Td
(the)Tj
1.4683 0 Td
(single)Tj
2.5285 0 Td
(neuron)Tj
3.1464 0 Td
(has)Tj
1.5477 0 Td
(a)Tj
0.652 0 Td
(positive)Tj
3.3165 0 Td
(weight.)Tj
3.1124 0 Td
(When)Tj
2.6816 0 Td
(the)Tj
1.474 0 Td
(estimate)Tj
3.5433 0 Td
(is)Tj
0.8504 0 Td
(too)Tj
1.5307 0 Td
(small)Tj
2.3244 0 Td
(with)Tj
2.0012 0 Td
(respect)Tj
-34.6901 -1.2983 Td
(to)Tj
1.0261 0 Td
(the)Tj
1.4683 0 Td
(signal,)Tj
2.7666 0 Td
(this)Tj
1.6724 0 Td
(neuron)Tj
3.1465 0 Td
(fires)Tj
1.9332 0 Td
(spikes,)Tj
2.8686 0 Td
(which)Tj
2.6533 0 Td
(brings)Tj
2.7382 0 Td
(the)Tj
1.4683 0 Td
(estimate)Tj
3.549 0 Td
(closer)Tj
2.5512 0 Td
(to)Tj
1.0261 0 Td
(the)Tj
1.4683 0 Td
(signal.)Tj
2.7666 0 Td
(However,)Tj
-33.1027 -1.2983 Td
(when)Tj
2.4037 0 Td
(the)Tj
1.4684 0 Td
(estimate)Tj
3.5432 0 Td
(is)Tj
0.8504 0 Td
(too)Tj
1.5307 0 Td
(large)Tj
2.16 0 Td
(with)Tj
2.0013 0 Td
(respect)Tj
3.0387 0 Td
(to)Tj
1.0204 0 Td
(the)Tj
1.4741 0 Td
(signal,)Tj
2.7665 0 Td
(this)Tj
1.6725 0 Td
(neuron)Tj
3.1464 0 Td
(cannot)Tj
2.982 0 Td
(correct)Tj
3.0388 0 Td
(for)Tj
1.3833 0 Td
(it)Tj
0.788 0 Td
(and)Tj
-35.2684 -1.2982 Td
(the)Tj
1.4683 0 Td
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/F6 1 Tf
(\))Tj
1.4116 0 Td
(and)Tj
1.7292 0 Td
(the)Tj
1.474 0 Td
(signal)Tj
2.5398 0 Td
(remains)Tj
3.4469 0 Td
(silent)Tj
2.3641 0 Td
(at)Tj
0.9524 0 Td
(all)Tj
1.1452 0 Td
(times)Tj
-32.2636 -1.2983 Td
(\()Tj
/F13 1 Tf
0 Tc
0.2778 0.0171 Td
()Tj
/F12 1 Tf
0.0623 -0.0171 Td
[()]TJ
/F15 1 Tf
0.3799 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6576 0 Td
()Tj
/F13 1 Tf
1.0489 0 Td
(0)Tj
/F15 1 Tf
0.9014 0 Td
(8)Tj
/F12 1 Tf
[()]TJ
/F6 1 Tf
-0.004 Tc
0.8956 0 Td
(\).)Tj
0.788 0 Td
(Since)Tj
2.3584 0 Td
(we)Tj
1.3267 0 Td
(are)Tj
1.4456 0 Td
(most)Tj
2.2054 0 Td
(interested)Tj
4.1669 0 Td
(in)Tj
1.0318 0 Td
(activity)Tj
3.1181 0 Td
(that)Tj
1.7801 0 Td
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2.4378 0 Td
(from)Tj
2.2053 0 Td
(the)Tj
1.4684 0 Td
(network)Tj
3.5489 0 Td
(alone,)Tj
2.6079 0 Td
(we)Tj
-35.4441 -1.3039 Td
(will)Tj
1.6611 0 Td
(assume)Tj
3.1464 0 Td
(that)Tj
1.7801 0 Td
(neuron)Tj
3.1464 0 Td
(1)Tj
0.6973 0 Td
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0.8447 0 Td
(perturbed)Tj
4.1896 0 Td
(by)Tj
1.1792 0 Td
(the)Tj
1.474 0 Td
(noise,)Tj
2.5512 0 Td
(sufficiently)Tj
4.6148 0 Td
(strong)Tj
2.7722 0 Td
(to)Tj
1.0205 0 Td
(provoke)Tj
3.4979 0 Td
(one)Tj
1.7008 0 Td
(single)Tj
-34.2762 -1.2982 Td
(spike,)Tj
2.5115 0 Td
(and)Tj
1.7291 0 Td
(that)Tj
1.78 0 Td
(after)Tj
2.0353 0 Td
(the)Tj
1.4683 0 Td
(first)Tj
1.8029 0 Td
(spike)Tj
2.2847 0 Td
(there)Tj
2.2677 0 Td
(is)Tj
0.8447 0 Td
(no)Tj
1.2756 0 Td
(more)Tj
2.3414 0 Td
(noise)Tj
2.33 0 Td
(in)Tj
1.0318 0 Td
(the)Tj
1.474 0 Td
(system.)Tj
3.1635 0 Td
(At)Tj
1.2019 0 Td
(the)Tj
1.4683 0 Td
(moment)Tj
3.6397 0 Td
(of)Tj
1.0148 0 Td
(per-)Tj
-35.6652 -1.2983 Td
(turbation,)Tj
4.2009 0 Td
(we)Tj
1.3322 0 Td
(assume)Tj
3.1464 0 Td
(that)Tj
/F12 1 Tf
1.7745 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 311.1874 488.296 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 316.97 489.9968 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(is)Tj
0.8447 0 Td
(at)Tj
0.9524 0 Td
(the)Tj
1.474 0 Td
(resting)Tj
2.9537 0 Td
(potential,)Tj
/F12 1 Tf
3.9855 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 425.707 488.296 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 431.4897 489.9968 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(=)Tj
0.7993 0 Td
(0.)Tj
0.9184 0 Td
(Before)Tj
2.812 0 Td
(the)Tj
1.4683 0 Td
(first)Tj
1.8085 0 Td
(spike,)Tj
2.5115 0 Td
(firing)Tj
-33.4655 -1.2982 Td
(rates)Tj
2.1089 0 Td
(are)Tj
1.4457 0 Td
(at)Tj
0.9524 0 Td
(zero)Tj
1.9333 0 Td
(and)Tj
1.7347 0 Td
(the)Tj
1.4684 0 Td
(estimated)Tj
4.0705 0 Td
(signal)Tj
2.5455 0 Td
(is)Tj
0.8447 0 Td
(silent,)Tj
/F13 1 Tf
0 Tc
2.5342 0 Td
(^)Tj
/F12 1 Tf
0.0567 0 Td
[()]TJ
/F15 1 Tf
0.3855 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
0.6634 0 Td
()Tj
/F13 1 Tf
1.0488 0 Td
(0)Tj
/F6 1 Tf
-0.004 Tc
(.)Tj
0.9466 0 Td
(Firing)Tj
2.6646 0 Td
(rule)Tj
1.7915 0 Td
(reduces)Tj
3.2768 0 Td
(to)Tj
1.0261 0 Td
(the)Tj
-32.2239 -1.6838 Td
(following:)Tj
/F15 1 Tf
0 Tc
( )Tj
/F12 1 Tf
4.7565 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 254.211 463.8047 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
0 -1.0677 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 258.1228 460.3464 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(^)Tj
/F12 1 Tf
0.0623 -0.017 Td
[()]TJ
/F15 1 Tf
0.3799 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.9864 0 Td
()Tj
/F18 1 Tf
0.9978 0 Td
(s)Tj
/F16 1 Tf
0.5499 0.7993 Td
(R)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 299.9055 456.8881 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 301.9464 458.5889 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 304.3275 460.1763 cm
BT
/F18 1 Tf
1 TL
0 0 Td
(Z)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 309.3165 457.6818 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 311.4708 460.1763 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
/F12 1 Tf
[()14()]TJ
/F20 1 Tf
1.5024 0 Td
(>)Tj
ET
Q
q
1 j
1 J
0 w
344.2394 462.8976 m
351.2693 462.8976 l
351.2693 462.5008 l
344.2394 462.5008 l
f*
5.9998 0 0 5.9998 344.2393 465.959 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 348.2645 468.17 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
/F12 1 Tf
0 -0.9637 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 346.2803 456.7748 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 353.4803 460.1763 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
363.4016 462.8976 m
366.4063 462.8976 l
366.4063 462.5008 l
363.4016 462.5008 l
f*
5.9998 0 0 5.9998 363.6283 463.8047 cm
BT
/F18 1 Tf
1 TL
0 0 Td
(n)Tj
/F13 1 Tf
-0.0377 -1.1716 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 368.107 460.1763 cm
BT
/F20 1 Tf
1 TL
0 0 Td
(:)Tj
/F6 1 Tf
-0.004 Tc
0.4875 0 Td
(To)Tj
1.338 0 Td
(initiate)Tj
3.0273 0 Td
(spiking,)Tj
3.3676 0 Td
(integration)Tj
4.6488 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(noise)Tj
2.3301 0 Td
(alone)Tj
-34.4917 -1.3889 Td
(has)Tj
1.542 0 Td
(to)Tj
1.0262 0 Td
(lead)Tj
1.8482 0 Td
(to)Tj
1.0261 0 Td
(crossing)Tj
3.5263 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(threshold:)Tj
/F18 1 Tf
0 Tc
2.8913 -2.7609 Td
(s)Tj
/F16 1 Tf
0.72 1.3549 Td
(Z)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 360.9637 429.9023 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 363.0047 432.1133 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 364.8755 429.9023 cm
BT
/F15 1 Tf
1 TL
0 0 Td
( )Tj
/F12 1 Tf
0.7653 0 Td
[()14()]TJ
/F13 1 Tf
-2.2204 -3.3544 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 376.6677 418.6771 cm
BT
/F18 1 Tf
1 TL
0 0 Td
(Z)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 381.6566 416.1826 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 385.1716 418.6771 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
/F12 1 Tf
[()14()]TJ
/F20 1 Tf
1.5023 0 Td
(>)Tj
ET
Q
q
1 j
1 J
0 w
417.9969 421.3417 m
428.1449 421.3417 l
428.1449 420.9449 l
417.9969 420.9449 l
f*
10 0 0 10 417.9968 425.4803 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 424.6866 429.1086 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
0 -1.0677 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 420.6047 411.8173 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
/F20 1 Tf
0.924 0.6859 Td
(:)Tj
/F6 1 Tf
-0.004 Tc
-21.7868 -2.9422 Td
(Let's)Tj
2.0579 0 Td
(assume)Tj
3.1465 0 Td
(that)Tj
1.7801 0 Td
(neuron)Tj
3.1464 0 Td
(1)Tj
0.6974 0 Td
(satisfied)Tj
3.4469 0 Td
(this)Tj
1.6724 0 Td
(condition)Tj
4.1159 0 Td
(at)Tj
0.9523 0 Td
(time)Tj
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2.0296 0 Td
[()]TJ
/F6 1 Tf
0.5159 0 Td
(=)Tj
/F12 1 Tf
0.7994 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
6.9998 0 0 6.9998 458.5889 392.8818 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 463.1811 389.2535 cm
BT
/F14 1 Tf
1 TL
-0.004 Tc
0 0 Td
[()]TJ
/F12 1 Tf
0.788 0 Td
[()14()]TJ
/F6 1 Tf
1.0148 0 Td
(and)Tj
1.7291 0 Td
(fired)Tj
2.0976 0 Td
(a)Tj
0.6577 0 Td
(spike.)Tj
2.5058 0 Td
(The)Tj
-35.1096 -2.2903 Td
(firing)Tj
2.4264 0 Td
(rate)Tj
1.7461 0 Td
(of)Tj
1.0148 0 Td
(neuron)Tj
3.1465 0 Td
(1)Tj
0.6973 0 Td
(jumps)Tj
2.7042 0 Td
(to)Tj
1.0205 0 Td
(1)Tj
0.6973 0 Td
(and)Tj
1.7291 0 Td
(decays)Tj
2.846 0 Td
(thereafter,)Tj
/F12 1 Tf
0 Tc
5.2554 0.8447 Td
[( )]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 436.4787 372.3023 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 432.8503 357.8456 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[( )]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 436.4787 355.3511 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 423.3259 383.4708 cm
BT
/F16 1 Tf
1 TL
0 0 Td
( )Tj
1.8311 0 Td
(!)Tj
/F15 1 Tf
1.0602 -1.7121 Td
(!)Tj
/F14 1 Tf
2.3867 0.8448 Td
[()2( )]TJ
/F15 1 Tf
1.5534 0 Td
( )Tj
/F18 1 Tf
1.1509 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 531.0425 378.4251 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 535.4078 374.7968 cm
BT
/F15 1 Tf
1 TL
0 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()14()]TJ
/F15 1 Tf
0.839 0 Td
()Tj
/F13 1 Tf
-3.8266 -1.6951 Td
(0)Tj
/F16 1 Tf
-5.04 2.5624 Td
( )Tj
9.802 0 Td
(!)Tj
/F6 1 Tf
-0.004 Tc
0.7824 -1.712 Td
(.)Tj
-37.0882 -2.296 Td
(At)Tj
1.2019 0 Td
(time)Tj
/F12 1 Tf
2.0296 0 Td
[()]TJ
/F6 1 Tf
0.5159 0 Td
(=)Tj
/F12 1 Tf
0.7994 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 248.485 347.0173 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 250.5826 343.3889 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
0.4478 0 Td
(the)Tj
1.4684 0 Td
(predicted)Tj
4.0025 0 Td
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2.5398 0 Td
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2.7042 0 Td
(in)Tj
1.0318 0 Td
(the)Tj
1.474 0 Td
(direction)Tj
3.8494 0 Td
(of)Tj
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1.0205 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 442.6015 341.7448 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 446.1165 343.3889 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
/F13 1 Tf
0 Tc
0.3855 0.017 Td
(^)Tj
/F12 1 Tf
0.0623 -0.017 Td
[()]TJ
/F15 1 Tf
0.3799 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6576 0 Td
(!)Tj
/F12 1 Tf
1.2813 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 487.7858 340.8944 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 492.8314 343.3889 cm
BT
/F14 1 Tf
1 TL
0 0 Td
[()2( )]TJ
/F15 1 Tf
1.5533 0 Td
( )Tj
/F18 1 Tf
1.1509 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9978 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 547.7669 347.0173 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 549.9212 343.3889 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F6 1 Tf
-0.004 Tc
(.)Tj
-34.9906 -1.2982 Td
(Since)Tj
2.3584 0 Td
(the)Tj
1.4684 0 Td
(signal)Tj
2.5398 0 Td
(is)Tj
0.8504 0 Td
(at)Tj
0.9524 0 Td
(zero,)Tj
2.16 0 Td
(this)Tj
1.6724 0 Td
(creates)Tj
2.9537 0 Td
(prediction)Tj
4.3767 0 Td
(error,)Tj
/F12 1 Tf
2.4888 0 Td
[()]TJ
/F6 1 Tf
(\()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 430.0724 334.0346 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 432.2267 330.4062 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\))Tj
0.5555 0 Td
(=)Tj
/F12 1 Tf
0.7994 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 450.6519 334.8283 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 454.2236 330.4062 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(.)Tj
0.4478 0 Td
(Weight)Tj
3.1635 0 Td
(of)Tj
1.0148 0 Td
(neuron)Tj
3.1464 0 Td
(2)Tj
0.6973 0 Td
(points)Tj
2.7213 0 Td
(in)Tj
-36.612 -1.2982 Td
(the)Tj
1.4683 0 Td
(opposite)Tj
3.617 0 Td
(direction)Tj
3.8551 0 Td
(than)Tj
2.0296 0 Td
(the)Tj
1.4684 0 Td
(weight)Tj
2.8856 0 Td
(of)Tj
1.0148 0 Td
(neuron)Tj
3.1464 0 Td
(1)Tj
0.6974 0 Td
(and)Tj
1.7291 0 Td
(neuron)Tj
3.1464 0 Td
(2)Tj
0.6972 0 Td
(should)Tj
2.914 0 Td
(now)Tj
1.9616 0 Td
(spike)Tj
2.2847 0 Td
(in)Tj
1.0375 0 Td
(order)Tj
2.4208 0 Td
(to)Tj
-36.3739 -1.3039 Td
(correct)Tj
3.0387 0 Td
(for)Tj
1.3833 0 Td
(the)Tj
1.4684 0 Td
(prediction)Tj
4.3766 0 Td
(error.)Tj
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1.1055 0 Td
(fact,)Tj
1.8936 0 Td
(this)Tj
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(is)Tj
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(since)Tj
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(neuron)Tj
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(2)Tj
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(receives)Tj
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(an)Tj
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(excitatory)Tj
4.1612 0 Td
(PSP)Tj
1.8142 0 Td
(while)Tj
2.3698 0 Td
(neuron)Tj
3.1464 0 Td
(1)Tj
0.6973 0 Td
(is)Tj
0.8447 0 Td
(inhibited)Tj
3.8551 0 Td
(through)Tj
3.4469 0 Td
(the)Tj
1.4684 0 Td
(negative)Tj
3.5319 0 Td
(self-connection:)Tj
/F12 1 Tf
0 Tc
-12.2626 -1.8935 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 337.3795 269.9716 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 340.8944 272.4661 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6576 0 Td
()Tj
/F12 1 Tf
1.0488 0 Td
[()22()]TJ
/F14 1 Tf
1.5194 0 Td
[()2( )]TJ
/F15 1 Tf
1.5534 0 Td
( )Tj
/F18 1 Tf
1.1565 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 435.4582 276.6047 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 437.5559 272.4661 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
-11.4462 -3.4695 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 329.7259 235.2755 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 333.2409 237.77 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6576 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
357.6189 240.4346 m
366.0094 240.4346 l
366.0094 240.0378 l
357.6189 240.0378 l
f*
10 0 0 10 357.6188 244.5732 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 362.4944 248.2015 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 359.3196 230.9102 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
/F15 1 Tf
0.89 0.6859 Td
( )Tj
/F12 1 Tf
0.9978 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 383.074 241.9086 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 388.1196 237.77 cm
BT
/F14 1 Tf
1 TL
0 0 Td
[()2( )]TJ
/F15 1 Tf
1.5533 0 Td
( )Tj
/F18 1 Tf
1.1566 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 443.1118 241.9086 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 445.2094 237.77 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F6 1 Tf
-0.004 Tc
-23.3233 -2.8913 Td
(At)Tj
1.2076 0 Td
(time)Tj
/F12 1 Tf
2.0239 0 Td
[()]TJ
/F6 1 Tf
0.5216 0 Td
(=)Tj
/F12 1 Tf
0.7993 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
6.9998 0 0 6.9998 260.5039 212.485 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 262.885 208.8566 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
0.4422 0 Td
(we)Tj
1.3322 0 Td
(have)Tj
2.058 0 Td
(that)Tj
/F12 1 Tf
1.7801 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 325.644 207.1559 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 329.159 208.8566 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
6.9998 0 0 6.9998 335.5653 212.485 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 337.9464 208.8566 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\))Tj
0.5612 0 Td
(=)Tj
/F12 1 Tf
0.7994 0 Td
[()22()]TJ
/F6 1 Tf
1.1848 0 Td
(and)Tj
/F12 1 Tf
0 Tc
1.7292 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 387.3259 206.3622 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 390.8409 208.8566 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 398.1543 212.485 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 400.2519 208.8566 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
0.6633 0 Td
()Tj
1.0431 0 Td
( )Tj
ET
Q
q
1 j
1 J
0 w
426.6709 211.5213 m
432.6236 211.5213 l
432.6236 211.1244 l
426.6709 211.1244 l
f*
5.9998 0 0 5.9998 426.6708 212.485 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 429.6188 214.6393 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 428.1448 205.4551 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 432.6236 208.8566 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(.)Tj
0.4478 0 Td
(Following)Tj
4.2179 0 Td
(the)Tj
1.4684 0 Td
(firing)Tj
2.4264 0 Td
(rule,)Tj
2.0183 0 Td
(there)Tj
2.262 0 Td
(will)Tj
-36.1017 -1.559 Td
(be)Tj
1.1509 0 Td
(a)Tj
0.6519 0 Td
(spike)Tj
2.2847 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(neuron)Tj
3.1464 0 Td
(2)Tj
0.6973 0 Td
(if)Tj
/F12 1 Tf
0 Tc
0.7767 0 Td
[()22()]TJ
/F20 1 Tf
1.2416 0 Td
(>)Tj
ET
Q
q
1 j
1 J
0 w
334.8283 195.9874 m
340.7244 195.9874 l
340.7244 195.5906 l
334.8283 195.5906 l
f*
5.9998 0 0 5.9998 334.8283 196.8944 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 337.7196 199.1055 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 336.3023 189.8645 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 342.3685 193.2661 cm
BT
/F20 1 Tf
1 TL
0 0 Td
(;)Tj
/F6 1 Tf
-0.004 Tc
0.4932 0 Td
(which)Tj
2.6475 0 Td
(reduces)Tj
3.2825 0 Td
(to)Tj
/F12 1 Tf
0 Tc
1.0262 0 Td
[()]TJ
/F20 1 Tf
0.7653 0 Td
(>)Tj
ET
Q
q
1 j
1 J
0 w
435.0047 195.9874 m
438.0094 195.9874 l
438.0094 195.5906 l
435.0047 195.5906 l
f*
5.9998 0 0 5.9998 435.0614 196.8944 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
/F13 1 Tf
-0.0094 -1.1716 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 438.0094 193.2661 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(.)Tj
0.4478 0 Td
(If)Tj
0.8447 0 Td
(for)Tj
1.389 0 Td
(example)Tj
/F12 1 Tf
3.549 0 Td
[()]TJ
/F6 1 Tf
0.7029 0 Td
(=)Tj
/F12 1 Tf
0.7994 0 Td
[()]TJ
/F6 1 Tf
(,)Tj
0.9241 0 Td
(neuron)Tj
3.1464 0 Td
(2)Tj
-35.6028 -1.5533 Td
(will)Tj
1.6611 0 Td
(spike.)Tj
2.5115 0 Td
(In)Tj
1.1112 0 Td
(the)Tj
1.4683 0 Td
(meantime,)Tj
4.4787 0 Td
(condition)Tj
4.1159 0 Td
(for)Tj
1.3833 0 Td
(spiking)Tj
3.1464 0 Td
(in)Tj
1.0318 0 Td
(neuron)Tj
3.1464 0 Td
(1)Tj
0.6973 0 Td
(is)Tj
/F15 1 Tf
0 Tc
0.8504 0 Td
( )Tj
ET
Q
q
1 j
1 J
0 w
465.3921 180.3969 m
471.3449 180.3969 l
471.3449 180 l
465.3921 180 l
f*
5.9998 0 0 5.9998 465.3921 181.3606 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 468.3401 183.5149 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 466.8661 174.3307 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 474.1228 177.7322 cm
BT
/F20 1 Tf
1 TL
0 0 Td
(>)Tj
ET
Q
q
1 j
1 J
0 w
484.611 180.3969 m
490.5638 180.3969 l
490.5638 180 l
484.611 180 l
f*
5.9998 0 0 5.9998 484.611 181.3606 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 487.559 183.5149 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 486.085 174.3307 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 490.5637 177.7322 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(.)Tj
0.4422 0 Td
(Neuron)Tj
3.3391 0 Td
(1)Tj
0.6917 0 Td
(will)Tj
1.6668 0 Td
(there-)Tj
-35.1947 -1.3889 Td
(fore)Tj
1.8085 0 Td
(remain)Tj
3.0841 0 Td
(silent.)Tj
2.5908 0 Td
(Spike)Tj
2.3981 0 Td
(of)Tj
1.0148 0 Td
(neuron)Tj
3.1465 0 Td
(2)Tj
0.6973 0 Td
(sends)Tj
2.4434 0 Td
(the)Tj
1.4684 0 Td
(predicted)Tj
3.9968 0 Td
(signal)Tj
2.5455 0 Td
(in)Tj
1.0375 0 Td
(the)Tj
1.4683 0 Td
(direction)Tj
3.855 0 Td
(of)Tj
/F12 1 Tf
1.0148 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 532.3464 162.1417 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 535.8614 163.8425 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(.)Tj
/F13 1 Tf
0 Tc
-24.4571 -1.8765 Td
(^)Tj
/F12 1 Tf
0.0567 -0.017 Td
[()]TJ
/F15 1 Tf
0.3855 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
0.6634 0 Td
()Tj
/F12 1 Tf
1.0488 0 Td
[()]TJ
/F14 1 Tf
1.0431 0 Td
[()2( )]TJ
/F15 1 Tf
1.5534 0 Td
( )Tj
/F18 1 Tf
1.1508 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4514 0 Td
( )Tj
/F12 1 Tf
0.9978 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 385.4551 149.0456 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 387.6094 144.907 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
0.9864 0 Td
( )Tj
/F12 1 Tf
0.9978 0 Td
[()]TJ
/F14 1 Tf
1.0318 0 Td
[()2( )]TJ
/F15 1 Tf
1.5534 0 Td
( )Tj
/F18 1 Tf
1.1508 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4514 0 Td
( )Tj
/F12 1 Tf
0.9977 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 472.7055 149.0456 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(00)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 476.4472 144.907 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F6 1 Tf
-0.004 Tc
-26.447 -2.2903 Td
(At)Tj
1.2075 0 Td
(time)Tj
/F12 1 Tf
2.0239 0 Td
[()]TJ
/F6 1 Tf
0.5216 0 Td
(=)Tj
/F12 1 Tf
0.7994 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
6.9998 0 0 6.9998 260.5039 125.6314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 265.096 122.0031 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(+)Tj
/F12 1 Tf
0.7993 0 Td
[()14()]TJ
/F6 1 Tf
1.0148 0 Td
(=)Tj
/F12 1 Tf
0.7994 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
6.9998 0 0 6.9998 294.1795 125.6314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
0.3482 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 298.9984 122.0031 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
0.4422 0 Td
(predicted)Tj
4.0024 0 Td
(signal)Tj
2.5399 0 Td
(is)Tj
0.8504 0 Td
(the)Tj
1.4683 0 Td
(following:)Tj
/F13 1 Tf
0 Tc
4.1499 0.017 Td
(^)Tj
/F12 1 Tf
0.0567 -0.017 Td
[()]TJ
/F15 1 Tf
0.3855 0 Td
()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 445.2094 125.6314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(00)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 448.9511 122.0031 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
0.6576 0 Td
()Tj
/F12 1 Tf
1.0488 0 Td
[()]TJ
/F14 1 Tf
1.0431 0 Td
[()2( )]TJ
/F15 1 Tf
1.5534 0 Td
( )Tj
/F18 1 Tf
1.1565 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4514 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 531.4393 125.6314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 533.537 122.0031 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
0.9921 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
/F6 1 Tf
-0.004 Tc
(.)Tj
0.9241 0 Td
(If)Tj
-36.2605 -1.2982 Td
(weights)Tj
3.2485 0 Td
(have)Tj
2.0579 0 Td
(the)Tj
1.4684 0 Td
(same)Tj
2.2563 0 Td
(strength,)Tj
/F12 1 Tf
3.7418 0 Td
[()]TJ
/F6 1 Tf
0.7086 0 Td
(=)Tj
/F12 1 Tf
0.7994 0 Td
[()]TJ
/F6 1 Tf
(,)Tj
0.9184 0 Td
(predicted)Tj
4.0025 0 Td
(signal)Tj
2.5398 0 Td
(is)Tj
0.8504 0 Td
(now)Tj
1.9559 0 Td
(close)Tj
2.1827 0 Td
(to)Tj
1.0261 0 Td
(zero)Tj
1.9332 0 Td
(and)Tj
1.7291 0 Td
(therefore)Tj
3.855 0 Td
(close)Tj
-35.274 -1.3039 Td
(to)Tj
1.0262 0 Td
(the)Tj
1.4683 0 Td
(desired)Tj
3.1238 0 Td
(signal.)Tj
2.7665 0 Td
(Prediction)Tj
4.4107 0 Td
(error)Tj
2.2677 0 Td
(is)Tj
0.8448 0 Td
(now)Tj
/F12 1 Tf
1.9615 0 Td
[()]TJ
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1.0035 0 Td
(neuron)Tj
3.0954 0 Td
(2,)Tj
0.9071 0 Td
(a)Tj
0.6463 0 Td
(depolarizing)Tj
5.1079 0 Td
(PSP)Tj
1.7914 0 Td
(is)Tj
0.8334 0 Td
(sent)Tj
-35.8693 -1.2983 Td
(back)Tj
2.0466 0 Td
(to)Tj
1.0092 0 Td
(neuron)Tj
3.1011 0 Td
(1.)Tj
0.9014 0 Td
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0.8334 0 Td
(this)Tj
1.644 0 Td
(PSP)Tj
1.7915 0 Td
(alone)Tj
2.3358 0 Td
(is)Tj
0.8333 0 Td
(strong)Tj
2.7213 0 Td
(enough)Tj
3.1748 0 Td
(to)Tj
1.0091 0 Td
(make)Tj
2.3584 0 Td
(the)Tj
1.4457 0 Td
(neuron)Tj
3.101 0 Td
(1)Tj
0.686 0 Td
(fire)Tj
1.5363 0 Td
(again,)Tj
2.5512 0 Td
(neuron)Tj
3.0954 0 Td
(1)Tj
-36.1755 -1.2982 Td
(will)Tj
1.6328 0 Td
(do)Tj
1.2359 0 Td
(so,)Tj
1.2869 0 Td
(even)Tj
2.0409 0 Td
(though)Tj
3.0274 0 Td
(there)Tj
2.2224 0 Td
(is)Tj
0.8334 0 Td
(no)Tj
1.2585 0 Td
(prediction)Tj
4.2917 0 Td
(error)Tj
2.2223 0 Td
(to)Tj
1.0091 0 Td
(correct)Tj
2.9877 0 Td
(for.)Tj
1.5761 0 Td
(In)Tj
1.0941 0 Td
(such)Tj
2.0296 0 Td
(a)Tj
0.6463 0 Td
(regime,)Tj
3.1577 0 Td
(the)Tj
1.4457 0 Td
(two)Tj
1.6837 0 Td
(neu-)Tj
-35.6822 -1.304 Td
(rons)Tj
1.9729 0 Td
(continue)Tj
3.7021 0 Td
(to)Tj
1.0091 0 Td
(spike,)Tj
2.4605 0 Td
(correcting)Tj
4.2406 0 Td
(and)Tj
1.7064 0 Td
(again)Tj
2.3301 0 Td
(re-creating)Tj
4.507 0 Td
(the)Tj
1.4457 0 Td
(prediction)Tj
4.2916 0 Td
(error.)Tj
2.4435 0 Td
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1.7573 0 Td
(time)Tj
1.9956 0 Td
(it)Tj
0.771 0 Td
(takes)Tj
-34.6334 -1.2982 Td
(to)Tj
1.0092 0 Td
(the)Tj
1.4456 0 Td
(network)Tj
3.4923 0 Td
(to)Tj
1.0091 0 Td
(integrate)Tj
4.4447 0 Td
(the)Tj
1.4457 0 Td
(information)Tj
4.9889 0 Td
(about)Tj
2.4548 0 Td
(a)Tj
0.6463 0 Td
(spike)Tj
2.245 0 Td
(and)Tj
1.7065 0 Td
(react)Tj
2.1316 0 Td
(accordingly,)Tj
5.0172 0 Td
(there)Tj
2.2224 0 Td
(is)Tj
-34.2593 -1.2983 Td
(already)Tj
3.0558 0 Td
(a)Tj
0.6463 0 Td
(new)Tj
1.8481 0 Td
(spurious)Tj
3.6 0 Td
(spike)Tj
2.2451 0 Td
(that)Tj
1.7461 0 Td
(has)Tj
1.5193 0 Td
(been)Tj
2.0863 0 Td
(generated.)Tj
4.286 0 Td
(Spiking)Tj
3.1974 0 Td
(is)Tj
0.8334 0 Td
(mechanically)Tj
5.3745 0 Td
(driven)Tj
2.7608 0 Td
(by)Tj
1.1679 0 Td
(lateral)Tj
-34.367 -1.3039 Td
(connectivity)Tj
5.04 0 Td
(and)Tj
1.7065 0 Td
(is)Tj
0.8277 0 Td
(due)Tj
1.6781 0 Td
(to)Tj
1.0091 0 Td
(the)Tj
1.4457 0 Td
(delay)Tj
2.2677 0 Td
(in)Tj
1.0148 0 Td
(synaptic)Tj
3.4639 0 Td
(transmission,)Tj
5.4821 0 Td
(which)Tj
2.6136 0 Td
(in)Tj
1.0204 0 Td
(our)Tj
1.5988 0 Td
(case)Tj
1.831 0 Td
(is)Tj
0.8334 0 Td
(also)Tj
1.7348 0 Td
(a)Tj
0.6463 0 Td
(delay)Tj
2.2677 0 Td
(in)Tj
-36.4816 -1.2983 Td
(sharing)Tj
3.1351 0 Td
(the)Tj
1.4457 0 Td
(information)Tj
4.989 0 Td
(about)Tj
2.4547 0 Td
(the)Tj
1.4457 0 Td
(prediction)Tj
4.2916 0 Td
(error.)Tj
2.4435 0 Td
(The)Tj
1.7574 0 Td
(requirement)Tj
5.1137 0 Td
(of)Tj
0.9978 0 Td
(a)Tj
0.6463 0 Td
(single)Tj
2.483 0 Td
(PSP)Tj
1.7915 0 Td
(being)Tj
-32.995 -1.2982 Td
(strong)Tj
2.7213 0 Td
(enough)Tj
3.1748 0 Td
(to)Tj
1.0091 0 Td
(drive)Tj
2.2223 0 Td
(the)Tj
1.4457 0 Td
(membrane)Tj
4.5014 0 Td
(potential)Tj
3.6907 0 Td
(across)Tj
2.6305 0 Td
(threshold)Tj
3.9345 0 Td
(might)Tj
2.5625 0 Td
(seem)Tj
2.2167 0 Td
(unrealistic.)Tj
4.5126 0 Td
(How-)Tj
-34.6221 -1.2983 Td
(ever,)Tj
2.075 0 Td
(in)Tj
1.0205 0 Td
(bigger)Tj
2.6702 0 Td
(network)Tj
3.4979 0 Td
(where)Tj
2.6079 0 Td
(neurons)Tj
3.4525 0 Td
(are)Tj
1.423 0 Td
(allowed)Tj
3.2315 0 Td
(to)Tj
1.0091 0 Td
(synchronize,)Tj
5.1987 0 Td
(it)Tj
0.7711 0 Td
(is)Tj
0.8277 0 Td
(enough)Tj
3.1747 0 Td
(that)Tj
1.7461 0 Td
(the)Tj
1.4456 0 Td
(sum)Tj
1.9049 0 Td
(of)Tj
-36.0564 -1.3039 Td
(PSPs)Tj
2.143 0 Td
(of)Tj
0.9978 0 Td
(all)Tj
1.1225 0 Td
(neurons)Tj
3.4526 0 Td
(with)Tj
1.9616 0 Td
(opposite)Tj
3.5546 0 Td
(selectivity)Tj
4.0535 0 Td
(makes)Tj
2.7156 0 Td
(the)Tj
1.4456 0 Td
(neuron)Tj
3.0955 0 Td
(fire,)Tj
1.7574 0 Td
(which)Tj
2.6079 0 Td
(is)Tj
0.8334 0 Td
(highly)Tj
2.6587 0 Td
(plausible.)Tj
-0.004 Tc
-31.2034 -1.2983 Td
(The)Tj
1.7858 0 Td
(occurrence)Tj
4.6601 0 Td
(of)Tj
1.0148 0 Td
(inefficient)Tj
4.2633 0 Td
(spiking)Tj
3.1464 0 Td
(is)Tj
0.8447 0 Td
(illustrated)Tj
4.2236 0 Td
(with)Tj
1.9956 0 Td
(a)Tj
0.6576 0 Td
(toy)Tj
1.4797 0 Td
(example.)Tj
3.7757 0 Td
(We)Tj
1.6043 0 Td
(will)Tj
1.6668 0 Td
(consider)Tj
3.651 0 Td
(the)Tj
-35.9657 -1.2983 Td
(special)Tj
2.8914 0 Td
(case)Tj
1.8595 0 Td
(where)Tj
2.6532 0 Td
(the)Tj
1.4683 0 Td
(two)Tj
1.7121 0 Td
(neurons)Tj
3.5036 0 Td
(have)Tj
2.0637 0 Td
(antisymmetric)Tj
6.0037 0 Td
(weights,)Tj
/F11 1 Tf
3.4753 0 Td
(w)Tj
/F6 1 Tf
0.9014 0 Td
(=)Tj
0.7993 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 483.3637 301.5496 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 486.9354 303.2503 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
/F12 1 Tf
0.4478 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 498.0472 301.5496 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 501.5622 303.2503 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\))Tj
0.5612 0 Td
(=)Tj
0.7994 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
/F6 1 Tf
(,)Tj
/F14 1 Tf
1.2699 0 Td
[()]TJ
/F12 1 Tf
[()]TJ
/F6 1 Tf
(\),)Tj
1.8425 0 Td
(which)Tj
-34.6277 -1.4683 Td
(results)Tj
2.8063 0 Td
(in)Tj
1.0374 0 Td
(the)Tj
1.4684 0 Td
(same)Tj
2.2563 0 Td
(thresholds)Tj
4.3767 0 Td
(for)Tj
1.3833 0 Td
(the)Tj
1.4683 0 Td
(two)Tj
1.7122 0 Td
(neurons,)Tj
/F12 1 Tf
0 Tc
3.7303 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 409.096 286.0724 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
/F20 1 Tf
(;)Tj
/F13 1 Tf
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 417.2031 288.5669 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
/F20 1 Tf
0.6633 0 Td
(>)Tj
ET
Q
q
1 j
1 J
0 w
441.6378 291.2882 m
447.5906 291.2882 l
447.5906 290.8913 l
441.6378 290.8913 l
f*
5.9998 0 0 5.9998 441.6377 292.2519 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 444.5858 294.4062 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 443.1118 285.1653 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 449.8015 288.5669 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
459.7228 291.2882 m
462.7276 291.2882 l
462.7276 290.8913 l
459.7228 290.8913 l
f*
5.9998 0 0 5.9998 459.9496 292.2519 cm
BT
/F18 1 Tf
1 TL
0 0 Td
(n)Tj
/F13 1 Tf
-0.0377 -1.1811 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 462.7275 288.5669 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(.)Tj
0.4478 0 Td
(The)Tj
1.7802 0 Td
(membrane)Tj
4.5694 0 Td
(potentials)Tj
-33.0687 -1.4116 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(two)Tj
1.7121 0 Td
(neurons)Tj
3.5036 0 Td
(are)Tj
1.4513 0 Td
(the)Tj
1.4683 0 Td
(following:)Tj
/F12 1 Tf
0 Tc
-5.5672 -2.7269 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 257.2157 244.6866 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 260.7307 247.1811 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
0.6633 0 Td
()Tj
/F12 1 Tf
1.0488 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 291.7417 244.6866 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 295.2566 247.1811 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F13 1 Tf
(0)Tj
/F15 1 Tf
()Tj
1.491 0 Td
()Tj
/F16 1 Tf
0.9921 1.3606 Td
(Z)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 330.4062 258.4629 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 332.4472 260.6173 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 325.644 238.2803 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 334.8283 247.1811 cm
BT
/F15 1 Tf
1 TL
0 0 Td
( )Tj
/F18 1 Tf
1.0374 0 Td
(l)Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 356.8818 244.6866 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 360.3968 247.1811 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
0.6066 0 Td
()Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 388.5732 251.3196 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 392.0314 247.1811 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 400.5921 244.6866 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 404.107 247.1811 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6009 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 432.0566 244.6866 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 435.5716 247.1811 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
/F12 1 Tf
[()14()]TJ
/F15 1 Tf
2.109 0 Td
()Tj
/F18 1 Tf
0.9921 0 Td
(s)Tj
/F16 1 Tf
0.72 1.3606 Td
(Z)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 491.3574 258.4629 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 493.3984 260.6173 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 486.5385 238.2803 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 497.4236 247.1811 cm
BT
/F18 1 Tf
1 TL
0 0 Td
(Z)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 502.4125 244.6866 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 505.9275 247.1811 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7256 0 Td
()Tj
/F12 1 Tf
[()14()]TJ
-22.881 -4.2349 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 291.0047 202.337 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 294.5196 204.8314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6576 0 Td
()Tj
/F12 1 Tf
1.0488 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 325.5874 202.337 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 329.1023 204.8314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F13 1 Tf
(0)Tj
/F15 1 Tf
()Tj
1.491 0 Td
()Tj
/F16 1 Tf
0.9921 1.3549 Td
(Z)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 364.2519 216.0566 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 366.2929 218.2677 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 359.433 195.9307 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 368.6173 204.8314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
( )Tj
/F18 1 Tf
1.0431 0 Td
(l)Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 390.7275 202.337 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 394.1858 204.8314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6009 0 Td
()Tj
/F12 1 Tf
0.9978 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 422.3622 208.97 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 425.8771 204.8314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 434.4377 202.337 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 437.896 204.8314 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6066 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
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321.789 631.6157 l
321.789 631.2189 l
313.3984 631.2189 l
f*
10 0 0 10 313.3984 635.7543 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 318.274 639.3826 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 315.0992 622.0913 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
/F15 1 Tf
0.89 0.6859 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 338.8535 633.0897 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 343.8992 628.9511 cm
BT
/F14 1 Tf
1 TL
0 0 Td
[()2( )]TJ
/F15 1 Tf
1.5533 0 Td
( )Tj
/F18 1 Tf
1.1566 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 398.8913 633.0897 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 400.9889 628.9511 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
0.9921 0 Td
()Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 425.707 633.0897 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 430.8094 628.9511 cm
BT
/F14 1 Tf
1 TL
0 0 Td
[()2( )]TJ
/F15 1 Tf
1.5533 0 Td
( )Tj
/F18 1 Tf
1.1509 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 485.7448 633.0897 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(00)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 489.4299 628.9511 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
-20.0294 -3.1974 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 295.7669 594.4818 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 299.2818 596.9763 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
0.6576 0 Td
()Tj
/F12 1 Tf
1.0488 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 328.5921 601.1149 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 333.6377 596.9763 cm
BT
/F14 1 Tf
1 TL
0 0 Td
[()2( )]TJ
/F15 1 Tf
1.5533 0 Td
( )Tj
/F18 1 Tf
1.1509 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9978 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 388.5732 601.1149 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 390.7275 596.9763 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
0.9864 0 Td
( )Tj
/F12 1 Tf
0.9978 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 415.4456 601.1149 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 420.4913 596.9763 cm
BT
/F14 1 Tf
1 TL
0 0 Td
[()2( )]TJ
/F15 1 Tf
1.5533 0 Td
( )Tj
/F18 1 Tf
1.1566 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 475.4834 601.1149 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(00)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 479.1685 596.9763 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F6 1 Tf
0.005 Tc
-26.7192 -2.262 Td
(At)Tj
1.2246 0 Td
(time)Tj
/F12 1 Tf
2.0636 0 Td
[()]TJ
/F6 1 Tf
0.5273 0 Td
(=)Tj
/F12 1 Tf
0.8107 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
6.9998 0 0 6.9998 261.2976 577.9842 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
0.3401 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 266.0598 574.3559 cm
BT
/F6 1 Tf
1 TL
0.005 Tc
0 0 Td
(,)Tj
0.4535 0 Td
(the)Tj
1.4967 0 Td
(firing)Tj
2.4831 0 Td
(rule)Tj
1.8255 0 Td
(for)Tj
1.4117 0 Td
(neuron)Tj
3.2031 0 Td
(1)Tj
0.703 0 Td
(is)Tj
/F12 1 Tf
0 Tc
0.8674 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 395.3763 577.9842 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 398.8913 574.3559 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F13 1 Tf
(1)Tj
/F15 1 Tf
1.1111 0 Td
( )Tj
/F14 1 Tf
1.1452 0 Td
[()2()]TJ
0.9014 0 Td
[( )]TJ
/F15 1 Tf
0.6747 0 Td
( )Tj
/F18 1 Tf
1.1622 0 Td
(l)Tj
/F15 1 Tf
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
1.4513 0 Td
( )Tj
/F12 1 Tf
0.9921 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 476.674 577.9842 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 478.8283 574.3559 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F20 1 Tf
1.4343 0 Td
(>)Tj
ET
Q
q
1 j
1 J
0 w
503.6598 577.0205 m
506.6646 577.0205 l
506.6646 576.6236 l
503.6598 576.6236 l
f*
5.9998 0 0 5.9998 503.8299 577.9842 cm
BT
/F18 1 Tf
1 TL
0 0 Td
(n)Tj
/F13 1 Tf
-0.0283 -1.1811 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 506.6645 574.3559 cm
BT
/F6 1 Tf
1 TL
0.005 Tc
0 0 Td
(.)Tj
0.4535 0 Td
(With)Tj
2.3187 0 Td
(the)Tj
1.4967 0 Td
(cost)Tj
1.8425 0 Td
(at)Tj
-36.7764 -1.3946 Td
(zero,)Tj
2.2054 0 Td
(the)Tj
1.4967 0 Td
(condition)Tj
4.1952 0 Td
(for)Tj
1.4117 0 Td
(spiking)Tj
3.2031 0 Td
(in)Tj
1.0545 0 Td
(neuron)Tj
3.2031 0 Td
(1)Tj
0.703 0 Td
(is)Tj
0.8674 0 Td
(again)Tj
2.4151 0 Td
(satisfied)Tj
3.5263 0 Td
(and)Tj
1.7574 0 Td
(neuron)Tj
3.2032 0 Td
(1)Tj
0.703 0 Td
(will)Tj
1.7006 0 Td
(fire)Tj
1.6044 0 Td
(another)Tj
-33.2501 -1.2982 Td
(spike.)Tj
2.5625 0 Td
(In)Tj
1.1226 0 Td
(an)Tj
1.2189 0 Td
(efficient)Tj
3.5262 0 Td
(working)Tj
3.617 0 Td
(regime,)Tj
3.2768 0 Td
(however,)Tj
3.9062 0 Td
(spike)Tj
2.33 0 Td
(at)Tj
/F12 1 Tf
0.9695 0 Td
[()]TJ
/F6 1 Tf
0.5215 0 Td
(=)Tj
/F12 1 Tf
0.8051 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 441.6944 551.0551 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
0.3496 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 448.1007 547.4267 cm
BT
/F6 1 Tf
1 TL
0.005 Tc
0 0 Td
(should)Tj
2.9706 0 Td
(not)Tj
1.5988 0 Td
(occur,)Tj
2.7496 0 Td
(since)Tj
2.2847 0 Td
(the)Tj
1.491 0 Td
(pre-)Tj
-35.9033 -1.2982 Td
(diction)Tj
3.1181 0 Td
(error)Tj
2.313 0 Td
(has)Tj
1.5704 0 Td
(already)Tj
3.1748 0 Td
(been)Tj
2.1543 0 Td
(corrected)Tj
4.0705 0 Td
(with)Tj
2.0353 0 Td
(the)Tj
1.4967 0 Td
(spike)Tj
2.3301 0 Td
(of)Tj
1.0318 0 Td
(neuron)Tj
3.203 0 Td
(2.)Tj
0.9354 0 Td
(In)Tj
1.1282 0 Td
(order)Tj
2.4661 0 Td
(to)Tj
1.0432 0 Td
(keep)Tj
2.1146 0 Td
(spiking)Tj
-34.1855 -1.3039 Td
(efficient,)Tj
3.7587 0 Td
(it)Tj
0.805 0 Td
(is)Tj
0.8674 0 Td
(necessary)Tj
4.0989 0 Td
(to)Tj
1.0431 0 Td
(increase)Tj
3.549 0 Td
(the)Tj
1.4967 0 Td
(linear)Tj
2.5625 0 Td
(cost.)Tj
2.0806 0 Td
(The)Tj
1.8085 0 Td
(following)Tj
4.0591 0 Td
(has)Tj
1.5704 0 Td
(to)Tj
1.0431 0 Td
(be)Tj
1.1679 0 Td
(satisfied:)Tj
/F12 1 Tf
3.7644 0 Td
[()]TJ
/F20 1 Tf
0 Tc
0.6746 0 Td
(>)Tj
/F6 1 Tf
0.005 Tc
0.9921 0 Td
(2)Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 563.1874 525.8834 cm
BT
/F6 1 Tf
1 TL
0.0067 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 569.0267 521.4047 cm
BT
/F20 1 Tf
1 TL
0 0 Td
()Tj
/F6 1 Tf
0.005 Tc
-36.9011 -1.2982 Td
(with)Tj
/F20 1 Tf
0 Tc
2.0296 0 Td
()Tj
/F6 1 Tf
0.005 Tc
0.6179 0 Td
(=)Tj
0.8107 0 Td
(1)Tj
/F14 1 Tf
0.703 0 Td
[()]TJ
/F6 1 Tf
0.7994 0 Td
(exp\()Tj
/F14 1 Tf
[()]TJ
/F17 1 Tf
()Tj
/F6 1 Tf
(\()Tj
/F12 1 Tf
[()]TJ
/F14 1 Tf
3.7417 0 Td
[()]TJ
/F12 1 Tf
0.7993 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 298.0913 512.0503 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 300.2456 508.422 cm
BT
/F6 1 Tf
1 TL
0.005 Tc
0 0 Td
(\)\).)Tj
1.1508 0 Td
(Efficient)Tj
3.6737 0 Td
(working)Tj
3.6226 0 Td
(regime)Tj
3.0444 0 Td
(allows)Tj
2.7496 0 Td
(for)Tj
1.4173 0 Td
(the)Tj
1.4967 0 Td
(error-correcting)Tj
6.8541 0 Td
(spike,)Tj
-34.0324 -1.2982 Td
(but)Tj
1.5761 0 Td
(prevents)Tj
3.6906 0 Td
(spikes)Tj
2.7043 0 Td
(that)Tj
1.8141 0 Td
(are)Tj
1.4739 0 Td
(not)Tj
1.5988 0 Td
(error-correcting,)Tj
7.0922 0 Td
(which)Tj
2.6986 0 Td
(implies)Tj
3.1861 0 Td
(the)Tj
1.4967 0 Td
(following:)Tj
/F13 1 Tf
0 Tc
4.2916 0 Td
(2)Tj
/F20 1 Tf
()Tj
1.1735 0 Td
(<)Tj
ET
Q
q
1 j
1 J
0 w
538.4693 498.1039 m
544.422 498.1039 l
544.422 497.7071 l
538.4693 497.7071 l
f*
5.9998 0 0 5.9998 540.17 499.0677 cm
BT
/F18 1 Tf
1 TL
0 0 Td
(n)Tj
/F12 1 Tf
-0.2834 -1.1716 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 541.4173 493.7385 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 547.2 495.4393 cm
BT
/F20 1 Tf
1 TL
0 0 Td
(<)Tj
/F13 1 Tf
1.0488 0 Td
(1)Tj
/F6 1 Tf
0.005 Tc
(.)Tj
-35.7673 -1.4003 Td
(With)Tj
2.3131 0 Td
(such)Tj
2.0976 0 Td
(relation)Tj
3.3846 0 Td
(between)Tj
3.5943 0 Td
(linear)Tj
2.5625 0 Td
(cost,)Tj
2.0806 0 Td
(weights)Tj
3.3108 0 Td
(and)Tj
1.7575 0 Td
(the)Tj
1.4967 0 Td
(strength)Tj
3.583 0 Td
(of)Tj
1.0374 0 Td
(the)Tj
1.4967 0 Td
(leak)Tj
1.8539 0 Td
(current,)Tj
3.4809 0 Td
(there)Tj
2.3073 0 Td
(is)Tj
-36.3569 -1.2982 Td
(one)Tj
1.7235 0 Td
(noisy)Tj
2.4037 0 Td
(spike)Tj
2.3301 0 Td
(followed)Tj
3.7247 0 Td
(by)Tj
1.1962 0 Td
(the)Tj
1.4967 0 Td
(error-correcting)Tj
6.8598 0 Td
(spike,)Tj
2.5625 0 Td
(after)Tj
2.0806 0 Td
(which)Tj
2.6986 0 Td
(there)Tj
2.3074 0 Td
(are)Tj
1.474 0 Td
(no)Tj
1.2926 0 Td
(further)Tj
-32.1504 -1.2983 Td
(spikes.)Tj
-0.004 Tc
1.1962 -1.3039 Td
(If)Tj
0.8504 0 Td
(linear)Tj
2.5115 0 Td
(cost)Tj
1.8085 0 Td
(is)Tj
0.8504 0 Td
(too)Tj
1.5307 0 Td
(small,)Tj
2.5512 0 Td
(i.e.)Tj
/F12 1 Tf
1.3606 0 Td
[()]TJ
/F20 1 Tf
0 Tc
0.6633 0 Td
(<)Tj
/F6 1 Tf
-0.004 Tc
0.9921 0 Td
(2)Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 352.8 446.9102 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 358.5826 442.4314 cm
BT
/F20 1 Tf
1 TL
0 0 Td
()Tj
/F6 1 Tf
-0.004 Tc
(,)Tj
0.839 0 Td
(the)Tj
1.4683 0 Td
(two)Tj
1.7122 0 Td
(neurons)Tj
3.5036 0 Td
(continue)Tj
3.7643 0 Td
(to)Tj
1.0262 0 Td
(mutually)Tj
3.7757 0 Td
(excite)Tj
2.5228 0 Td
(each)Tj
-34.469 -1.2982 Td
(other.)Tj
2.5739 0 Td
(The)Tj
1.7801 0 Td
(scenario)Tj
3.5547 0 Td
(with)Tj
2.0012 0 Td
(antisymmetric)Tj
6.0036 0 Td
(weights)Tj
3.2485 0 Td
(\()Tj
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 401.6692 427.8047 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 407.3952 429.4488 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(=)Tj
/F14 1 Tf
0.7993 0 Td
[()]TJ
/F12 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 427.748 427.8047 cm
BT
/F6 1 Tf
1 TL
-0.0053 Tc
0 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 431.3196 429.4488 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(\))Tj
0.5555 0 Td
(is)Tj
0.8504 0 Td
(particularly)Tj
4.8132 0 Td
(prone)Tj
2.5852 0 Td
(to)Tj
1.0261 0 Td
(drive)Tj
2.2677 0 Td
(long-)Tj
-35.2286 -1.2982 Td
(lasting)Tj
2.8346 0 Td
(Up)Tj
1.4684 0 Td
(states.)Tj
2.6191 0 Td
(With)Tj
2.2734 0 Td
(antisymmetric)Tj
6.0038 0 Td
(weights,)Tj
3.4695 0 Td
(each)Tj
2.024 0 Td
(spike)Tj
2.2847 0 Td
(produces)Tj
3.872 0 Td
(the)Tj
1.4683 0 Td
(same)Tj
2.2507 0 Td
(amount)Tj
3.3505 0 Td
(of)Tj
1.0148 0 Td
(excita-)Tj
-34.9338 -1.3039 Td
(tion)Tj
1.8425 0 Td
(\(to)Tj
1.3606 0 Td
(the)Tj
1.474 0 Td
(pair)Tj
1.8085 0 Td
(neuron\))Tj
3.4866 0 Td
(and)Tj
1.7291 0 Td
(inhibition)Tj
4.2236 0 Td
(\(to)Tj
1.3663 0 Td
(itself\).)Tj
2.6702 0 Td
(The)Tj
1.7802 0 Td
(amount)Tj
3.3561 0 Td
(of)Tj
1.0148 0 Td
(self-inhibition)Tj
5.896 0 Td
(and)Tj
1.7291 0 Td
(lateral)Tj
-33.7376 -1.2983 Td
(excitation)Tj
4.1499 0 Td
(to)Tj
1.0261 0 Td
(a)Tj
0.652 0 Td
(given)Tj
2.3811 0 Td
(neuron)Tj
3.1464 0 Td
(therefore)Tj
3.8551 0 Td
(perfectly)Tj
3.668 0 Td
(compensates,)Tj
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1.355 0 Td
(subtract)Tj
3.5433 0 Td
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/F12 1 Tf
0 Tc
2.2563 0 Td
[()]TJ
0.4706 0 Td
[()]TJ
1.593 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 485.348 711.0425 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[( )]TJ
0.6141 0 Td
[()22()]TJ
-0.6141 -1.0677 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 496.5732 707.4141 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F18 1 Tf
(t)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 504.7937 711.0425 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 506.948 707.4141 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
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0.009 Tc
0.6009 0 Td
(the)Tj
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-31.2942 -1.5023 Td
[()10()]TJ
1.3493 0 Td
[()60()]TJ
ET
Q
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1 j
1 J
0 w
5.9998 0 0 5.9998 227.1685 697.3228 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()5()]TJ
1.0016 0 Td
[(\n)12(")]TJ
0.8315 0 Td
[(")-36(#)]TJ
0.5386 0 Td
[()]TJ
-2.3717 -1.1622 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 244.2897 692.3905 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F18 1 Tf
(t)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 252.5102 696.0188 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 254.6645 692.3905 cm
BT
/F15 1 Tf
1 TL
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()Tj
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0.009 Tc
(,)Tj
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[(!)]TJ
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[()]TJ
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[()]TJ
1.593 0 Td
[()]TJ
/F15 1 Tf
()Tj
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(t)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 268.2141 638.9291 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 270.3118 634.7905 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
0.6746 0 Td
()Tj
1.0488 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
14 0 0 14 291.4015 648.0566 cm
BT
/F16 1 Tf
1 TL
0 0 Td
(X)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 299.1685 650.0409 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
-0.3874 -4.5828 Td
[()]TJ
/F15 1 Tf
()Tj
/F13 1 Tf
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 313.1716 648.3968 cm
BT
/F16 1 Tf
1 TL
0 0 Td
(Z)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 318.7275 650.3811 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
/F13 1 Tf
-0.7181 -5.0269 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 325.1338 634.7905 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 329.8393 632.296 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 331.937 634.7905 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
/F15 1 Tf
0.7313 0 Td
()Tj
/F12 1 Tf
[()14()]TJ
/F15 1 Tf
1.2359 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 355.4645 639.9496 cm
BT
/F15 1 Tf
1 TL
0 0 Td
( )Tj
/F13 1 Tf
0.7748 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
365.2724 637.5118 m
372.9827 637.5118 l
372.9827 637.115 l
365.2724 637.115 l
f*
10 0 0 10 366.633 641.5937 cm
BT
/F13 1 Tf
1 TL
0 0 Td
(1)Tj
/F12 1 Tf
-0.136 -1.3662 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
14 0 0 14 374.6267 648.0566 cm
BT
/F16 1 Tf
1 TL
0 0 Td
(X)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 382.3937 650.0409 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
-0.3874 -4.5828 Td
[()]TJ
/F15 1 Tf
()Tj
/F13 1 Tf
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 394.7527 634.7905 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
[()]TJ
0.856 0 Td
[()]TJ
1.5874 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 425.7637 638.9291 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[( )]TJ
0.6141 0 Td
[()22()]TJ
-0.6141 -1.1527 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 436.9889 634.7905 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F18 1 Tf
(t)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 445.2094 638.9291 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 447.307 634.7905 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
0.6179 0 Td
( )Tj
/F12 1 Tf
0.9978 0 Td
[()10()]TJ
1.3493 0 Td
[()60()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 490.6204 639.7228 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()5()]TJ
1.0016 0 Td
[(\n)12(")]TJ
0.8315 0 Td
[(")-36(#)]TJ
0.5386 0 Td
[()]TJ
-2.3717 -1.1622 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 507.7417 634.7905 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F18 1 Tf
(t)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 515.9622 638.9291 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 518.0598 634.7905 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
4.0535 0 Td
()Tj
/F6 1 Tf
(19)Tj
/F15 1 Tf
()Tj
/F6 1 Tf
-0.004 Tc
-34.6618 -3.5716 Td
(The)Tj
1.7859 0 Td
(peak)Tj
2.0919 0 Td
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4.2803 0 Td
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3.3562 0 Td
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(the)Tj
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3.7077 0 Td
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3.5886 0 Td
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-33.3295 -1.2982 Td
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2.914 0 Td
(out)Tj
1.5534 0 Td
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1.0261 0 Td
(be)Tj
1.1509 0 Td
(at)Tj
0.9524 0 Td
(zero)Tj
1.9332 0 Td
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(lag.)Tj
1.5931 0 Td
(For)Tj
1.6271 0 Td
(estimating)Tj
4.3993 0 Td
(the)Tj
1.4684 0 Td
(S-MUA,)Tj
3.5886 0 Td
(we)Tj
1.3266 0 Td
(used)Tj
2.0579 0 Td
(50)Tj
1.1736 0 Td
(trials,)Tj
2.4207 0 Td
(each)Tj
2.0295 0 Td
(200)Tj
1.6498 0 Td
(sec-)Tj
-34.8942 -1.2983 Td
(onds)Tj
2.1657 0 Td
(long.)Tj
/F11 1 Tf
-0.9695 -1.3039 Td
(3.2)Tj
1.4627 0 Td
(Duration)Tj
4.0989 0 Td
(of)Tj
1.0658 0 Td
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4.8699 0 Td
(intervals)Tj
3.8324 0 Td
(and)Tj
1.7915 0 Td
(duration)Tj
3.8891 0 Td
(of)Tj
1.0658 0 Td
(Up)Tj
1.4967 0 Td
(states.)Tj
/F6 1 Tf
3.3902 0 Td
(Inter-burst)Tj
4.6147 0 Td
(intervals,)Tj
-32.7739 -1.2983 Td
(similarly)Tj
3.685 0 Td
(to)Tj
1.0262 0 Td
(Inter-spike)Tj
4.6204 0 Td
(intervals,)Tj
3.8494 0 Td
(measure)Tj
3.5773 0 Td
(how)Tj
1.9389 0 Td
(much)Tj
2.5228 0 Td
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2.024 0 Td
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2.9877 0 Td
(between)Tj
3.5262 0 Td
(subsequent)Tj
-29.7579 -1.2982 Td
(bursts/Up)Tj
4.2293 0 Td
(states.)Tj
2.6192 0 Td
(Mean)Tj
2.5171 0 Td
(Inter-burst)Tj
4.6205 0 Td
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3.2655 0 Td
(is)Tj
0.8447 0 Td
(the)Tj
1.474 0 Td
(mean)Tj
2.4434 0 Td
(across)Tj
2.6759 0 Td
(all)Tj
1.1452 0 Td
(Inter-burst)Tj
4.6205 0 Td
(intervals)Tj
3.6225 0 Td
(that)Tj
-34.0778 -1.304 Td
(occurred)Tj
3.7928 0 Td
(during)Tj
2.931 0 Td
(a)Tj
0.6519 0 Td
(simulation)Tj
4.4957 0 Td
(trial.)Tj
2.0637 0 Td
(Distribution)Tj
5.176 0 Td
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1.0148 0 Td
(Inter-burst)Tj
4.6204 0 Td
(intervals)Tj
3.6227 0 Td
(was)Tj
1.7008 0 Td
(fitted)Tj
2.3186 0 Td
(with)Tj
2.0012 0 Td
(the)Tj
-34.3896 -1.2982 Td
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3.4413 0 Td
(distribution)Tj
4.9662 0 Td
(\(function)Tj
3.9685 0 Td
(gamfit)Tj
3.5773 0 Td
(by)Tj
1.1849 0 Td
(Matlab\).)Tj
3.5943 0 Td
(The)Tj
1.7858 0 Td
(criterion)Tj
3.6964 0 Td
(for)Tj
1.3833 0 Td
(an)Tj
1.2075 0 Td
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1.4684 0 Td
(state)Tj
2.0351 0 Td
(is)Tj
0.8504 0 Td
(at)Tj
0.9524 0 Td
(least)Tj
1.9843 0 Td
(20)Tj
-36.0961 -1.2983 Td
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3.2258 0 Td
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3.1465 0 Td
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2.4321 0 Td
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6.2418 0 Td
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1.99 0 Td
(criterion)Tj
3.6963 0 Td
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1.2983 0 Td
(by)Tj
1.1848 0 Td
(hand,)Tj
2.4832 0 Td
(however,)Tj
3.838 0 Td
(its)Tj
1.1451 0 Td
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-35.291 -1.2983 Td
(cise)Tj
1.6951 0 Td
(value)Tj
2.3074 0 Td
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2.0353 0 Td
(not)Tj
1.576 0 Td
(qualitatively)Tj
5.0797 0 Td
(change)Tj
3.0387 0 Td
(the)Tj
1.474 0 Td
(results.)Tj
-16.01 -1.3039 Td
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3.9118 0 Td
(of)Tj
1.0148 0 Td
(Up)Tj
1.474 0 Td
(states)Tj
2.3981 0 Td
(measures)Tj
3.9402 0 Td
(the)Tj
1.4683 0 Td
(length)Tj
2.7326 0 Td
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1.0148 0 Td
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1.474 0 Td
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2.6192 0 Td
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2.5171 0 Td
(duration)Tj
3.702 0 Td
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1.4684 0 Td
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2.4433 0 Td
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(dura-)Tj
-35.24 -1.2982 Td
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2.4265 0 Td
(collected)Tj
3.7417 0 Td
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2.9253 0 Td
(the)Tj
1.4683 0 Td
(simulation)Tj
4.4958 0 Td
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2.0636 0 Td
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1.0148 0 Td
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3.702 0 Td
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1.7008 0 Td
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1.5704 0 Td
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-35.0189 -1.2983 Td
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1.627 0 Td
(this)Tj
1.6725 0 Td
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3.0897 0 Td
(we)Tj
1.3266 0 Td
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2.3244 0 Td
(the)Tj
1.4683 0 Td
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4.972 0 Td
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1.0148 0 Td
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4.0591 0 Td
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2.0012 0 Td
(a)Tj
0.652 0 Td
(non-)Tj
-35.155 -1.3039 Td
(parametric)Tj
4.5808 0 Td
(kernel-smoothing)Tj
7.3814 0 Td
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4.9662 0 Td
(\(function)Tj
3.9685 0 Td
(fitdist,)Tj
3.5376 0 Td
(specification)Tj
5.2441 0 Td
(`Kernel',`epanech-)Tj
-29.6786 -1.2983 Td
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2.931 0 Td
(Matlab\).)Tj
3.6 0 Td
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1.6271 0 Td
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4.6148 0 Td
(intervals)Tj
3.6283 0 Td
(as)Tj
1.0148 0 Td
(well)Tj
1.8198 0 Td
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(the)Tj
1.4683 0 Td
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3.7021 0 Td
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2.6247 0 Td
(a)Tj
0.652 0 Td
(simulation)Tj
-32.5698 -1.2983 Td
(trial)Tj
1.8369 0 Td
(corresponding)Tj
6.0604 0 Td
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1.0205 0 Td
(1000)Tj
2.1259 0 Td
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3.3732 0 Td
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1.6952 0 Td
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4.8359 0 Td
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1.5421 0 Td
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4.6147 0 Td
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1.0148 0 Td
(variation)Tj
3.8041 0 Td
(2)Tj
0.6973 0 Td
(\(CV2\))Tj
2.7326 0 Td
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0.8447 0 Td
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1.474 0 Td
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1.0148 0 Td
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4.0252 0 Td
(computed)Tj
4.2633 0 Td
(from)Tj
2.211 0 Td
(sequences)Tj
4.2179 0 Td
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1.0148 0 Td
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4.6148 0 Td
(Intervals)Tj
3.702 0 Td
([)Tj
0.83 0.64 0.02 0 k
(14)Tj
0 g
(].)Tj
2.075 0 Td
(CV2)Tj
2.0522 0 Td
(of)Tj
1.0148 0 Td
(neuron)Tj
/F12 1 Tf
3.1464 0 Td
[()]TJ
/F6 1 Tf
0.4875 0 Td
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0.8504 0 Td
(com-)Tj
-34.6901 -1.2983 Td
(puted)Tj
2.5115 0 Td
(as)Tj
1.0148 0 Td
(follows:)Tj
/F12 1 Tf
0 Tc
9.7001 -2.6419 Td
[($)]TJ
/F13 1 Tf
1.372 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 350.9858 336.1889 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 355.9181 338.6834 cm
BT
/F15 1 Tf
1 TL
0 0 Td
()Tj
/F13 1 Tf
1.0488 0 Td
(2)Tj
ET
Q
q
1 j
1 J
0 w
373.0961 341.4047 m
424.7433 341.4047 l
424.7433 341.0079 l
373.0961 341.0079 l
f*
10 0 0 10 373.096 346.6204 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(j)Tj
/F12 1 Tf
[(%)8(%)]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 386.9858 344.1259 cm
BT
/F12 1 Tf
1 TL
0 0 Td
[()]TJ
/F15 1 Tf
()Tj
/F13 1 Tf
(1)Tj
ET
Q
q
1 j
1 J
0 w
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Q
q
1 j
1 J
0 w
496.8 648.5102 m
503.8299 648.5102 l
503.8299 648.1134 l
496.8 648.1134 l
f*
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/F12 1 Tf
1 TL
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[()]TJ
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Q
q
1 j
1 J
0 w
5 0 0 5 500.8251 651.6283 cm
BT
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Q
q
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1 J
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Q
q
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Q
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1 J
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Q
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[()]TJ
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Q
q
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1 J
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Q
q
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BT
/F15 1 Tf
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Q
q
1 j
1 J
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BT
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ET
Q
q
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BT
/F15 1 Tf
1 TL
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ET
Q
q
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1 J
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Q
q
1 j
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503.6031 617.726 m
509.0457 617.726 l
509.0457 617.3291 l
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[()]TJ
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Q
q
1 j
1 J
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BT
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ET
Q
q
1 j
1 J
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ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 411.9307 577.0204 cm
BT
/F15 1 Tf
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ET
Q
q
1 j
1 J
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BT
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1 TL
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[()]TJ
/F15 1 Tf
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/F12 1 Tf
[()]TJ
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ET
Q
q
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1 J
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BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
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1 J
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1 TL
0 0 Td
()Tj
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0.7257 0 Td
[(\n)]TJ
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Q
q
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1 J
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5.9998 0 0 5.9998 475.7669 574.9228 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 477.9212 571.2944 cm
BT
/F15 1 Tf
1 TL
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[()14()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 490.2803 574.9228 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
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1 TL
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1.8085 0 Td
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1.7008 0 Td
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0.8107 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 359.2062 555.7039 cm
BT
/F12 1 Tf
1 TL
0.0067 Tc
0 0 Td
[(\r)]TJ
ET
Q
q
1 j
1 J
0 w
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BT
/F17 1 Tf
1 TL
0.005 Tc
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()Tj
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(\()Tj
/F12 1 Tf
[()]TJ
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1.3492 0 Td
[()]TJ
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0.7994 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 389.537 561.8267 cm
BT
/F12 1 Tf
1 TL
0.0067 Tc
0 0 Td
[(\r)]TJ
ET
Q
q
1 j
1 J
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BT
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1 TL
0.005 Tc
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(\))Tj
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(with)Tj
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[()]TJ
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Q
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1 J
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BT
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1 TL
0.0067 Tc
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[(\r)]TJ
ET
Q
q
1 j
1 J
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Q
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1 J
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BT
/F13 1 Tf
1 TL
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(2)Tj
ET
Q
q
1 j
1 J
0 w
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Q
q
1 j
1 J
0 w
251.5465 478.7717 m
258.5764 478.7717 l
258.5764 478.3748 l
251.5465 478.3748 l
f*
5.9998 0 0 5.9998 251.5464 479.6787 cm
BT
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1 TL
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[()]TJ
ET
Q
q
1 j
1 J
0 w
5 0 0 5 255.5716 481.8897 cm
BT
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(2)Tj
ET
Q
q
1 j
1 J
0 w
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BT
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1 TL
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(2)Tj
ET
Q
q
1 j
1 J
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1 J
0 w
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BT
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[(&)]TJ
ET
Q
q
1 j
1 J
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BT
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1.8538 0 Td
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Q
q
1 j
1 J
0 w
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BT
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1 TL
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0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
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[()]TJ
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Q
q
1 j
1 J
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[()]TJ
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Q
q
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1 J
0 w
5.9998 0 0 5.9998 458.1354 394.6393 cm
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[()]TJ
ET
Q
q
1 j
1 J
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[( )]TJ
ET
Q
q
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1 J
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5.9998 0 0 5.9998 463.9181 394.6393 cm
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[()]TJ
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[( )]TJ
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[()]TJ
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q
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1 J
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[()]TJ
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Q
q
1 j
1 J
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[()]TJ
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[()]TJ
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Q
q
1 j
1 J
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[()]TJ
ET
Q
q
1 j
1 J
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[()]TJ
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Q
q
1 j
1 J
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5.9998 0 0 5.9998 424.3464 342.3118 cm
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[()]TJ
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1 j
1 J
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[()]TJ
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[()]TJ
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[()]TJ
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Q
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1 J
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[()]TJ
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[()]TJ
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[()]TJ
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[()]TJ
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q
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1 J
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[()]TJ
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1 j
1 J
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1 TL
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[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 515.8488 342.3118 cm
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/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 517.8897 344.8062 cm
BT
/F12 1 Tf
1 TL
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[( )]TJ
ET
Q
q
1 j
1 J
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5.9998 0 0 5.9998 521.5181 342.3118 cm
BT
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[()]TJ
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Q
q
1 j
1 J
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[()]TJ
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[()]TJ
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1 J
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/F12 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
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443.7921 328.8756 m
450.7654 328.8756 l
450.7654 328.4787 l
443.7921 328.4787 l
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5.9998 0 0 5.9998 443.7921 331.9937 cm
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1 TL
0 0 Td
[()]TJ
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Q
q
1 j
1 J
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5 0 0 5 447.7606 334.2047 cm
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(2)Tj
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[()]TJ
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1 J
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5.9998 0 0 5.9998 445.7763 322.8094 cm
BT
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(2)Tj
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1 J
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7.4999 0 0 6.6659 221.6692 310.6204 cm
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[()]TJ
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1 J
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7.4999 0 0 6.6659 490.3937 310.6204 cm
BT
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1 TL
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[()]TJ
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Q
q
1 j
1 J
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10 0 0 10 494.5889 312.2645 cm
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341.0646 298.2047 m
348.0945 298.2047 l
348.0945 297.8079 l
341.0646 297.8079 l
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5.9998 0 0 5.9998 341.0645 301.2661 cm
BT
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[()]TJ
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1 j
1 J
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5 0 0 5 345.0897 303.4771 cm
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[()]TJ
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(2)Tj
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1 j
1 J
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7.4999 0 0 6.6659 210.0472 253.9275 cm
BT
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1 TL
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(1,2)Tj
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Q
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1 j
1 J
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(10\))Tj
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7.4999 0 0 6.6659 376.3842 253.9275 cm
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