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An extensive dataset of spiking ac2vity to reveal the syntax of the
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1. Department of Vision & Cognition, Netherlands Institute for Neuroscience (KNAW), 1105 BA
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Amsterdam, Netherlands.
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2. Department of Integrative Neurophysiology, VU University, De Boelelaan 1085, 1081 HV
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Amsterdam, Netherlands.
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3. Department of Neurosurgery, Academic Medical Centre, Postbus 22660, 1100 DD Amsterdam,
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Netherlands.
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4. Laboratory of Visual Brain Therapy, Sorbonne Université, INSERM, CNRS, Institut de la Vision, 17
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Monkey N
V1/V1
V4/V4
IT/IT
V1/V4
V1/IT
V4/IT
Spontanous noise correlation
0.19
0.16
0.09
0.16
0.17
0.14
Peak noise correlation
0.11
0.10
0.09
0.11
0.11
0.09
Signal correlation
0.35
0.12
0.14
0.27
0.27
0.12
Monkey F
Spontanous noise correlation
0.11
0.17
0.13
0.11
0.11
0.12
Peak noise correlation
0.07
0.07
0.08
0.07
0.07
0.07
Signal correlation
0.19
0.28
0.21
0.17
0.16
0.20
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Stimulus presentation
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"
/*"):*<*-9*7"##+#>B"-&9G:&("8=&.*<"6:'="92*"UTVWX5"8=&.*"7&9&H&<*12"F'-<8<98-."'6"
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"
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"""
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"""
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"
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𝑓(𝑡)=*𝑑∙exp*(𝜇𝛼+0.5𝜎1𝛼1−𝛼𝑡)∙(𝐺
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