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Generalized Linear Models (GLMs) II
Statistical modeling and analysis of neural dataNEU 560, Spring 2018
Lecture 10
Jonathan Pillow
1
Summary:
k̂ = (XTX)�1XTY
1. “Linear-Gaussian” GLM: Y |X,~k ⇠ N (X~k,�2I)
2. Bernoulli GLM:
3. Poisson GLM:
yt|~xt,~k ⇠ Ber(f(~xt · ~k))
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<latexit sha1_base64="miR0nT8tJiZuVKiaUfnxELw/TEo=">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</latexit>
2
stimulus filter Poissonspiking
stimulusk f λ(t)
conditional intensity(spike rate)
exponentialnonlinearity
Linear-Nonlinear-Poisson
3
Fitting the nonlinearity
<latexit sha1_base64="IqiPS5IzX5l+5dK33I8KUFXVZ9E=">AAAB7HicbVBNT8JAEJ3iF+IX6tHLRmLiibRe1BvRi0dMLJBAQ7bLFla222Z3akIa/oMXD2q8+oO8+W9coAcFXzLJy3szmZkXplIYdN1vp7S2vrG5Vd6u7Ozu7R9UD49aJsk04z5LZKI7ITVcCsV9FCh5J9WcxqHk7XB8O/PbT1wbkagHnKQ8iOlQiUgwilZq9UYUybhfrbl1dw6ySryC1KBAs1/96g0SlsVcIZPUmK7nphjkVKNgkk8rvczwlLIxHfKupYrG3AT5/NopObPKgESJtqWQzNXfEzmNjZnEoe2MKY7MsjcT//O6GUZXQS5UmiFXbLEoyiTBhMxeJwOhOUM5sYQyLeythI2opgxtQBUbgrf88irxL+rXde/erTVuijTKcAKncA4eXEID7qAJPjB4hGd4hTcncV6cd+dj0Vpyiplj+APn8weYqo6q</latexit>
Filter k specifies a directionin stimulus space (i.e. a 1D subspace)
4
1) Project onto subspace spanned by k
Fitting the nonlinearity
<latexit sha1_base64="flj8ApT0XXAd006blhKZPNAqW50=">AAACDHicbZC9TsMwFIUdfkv5CzCyWBQkpiphAQakChbGIhFaqYkix3VaK44T2U6lKskLsPAqLAyAWHkANt4Gt80ALUey9Once3V9T5AyKpVlfRtLyyura+u1jfrm1vbOrrm3/yCTTGDi4IQlohsgSRjlxFFUMdJNBUFxwEgniG4m9c6ICEkTfq/GKfFiNOA0pBgpbfnmsTsiGGZ+BK+gGwqEc3eIFIzKvChmVBSlbzaspjUVXAS7ggao1PbNL7ef4CwmXGGGpOzZVqq8HAlFMSNl3c0kSRGO0ID0NHIUE+nl02tKeKKdPgwToR9XcOr+nshRLOU4DnRnjNRQztcm5n+1XqbCCy+nPM0U4Xi2KMwYVAmcRAP7VBCs2FgDwoLqv0I8RDoTpQOs6xDs+ZMXwTlrXjbtO6vRuq7SqIFDcAROgQ3OQQvcgjZwAAaP4Bm8gjfjyXgx3o2PWeuSUc0cgD8yPn8AlRabeQ==</latexit>
<latexit sha1_base64="O9GuRVuYx2v+9x/1eESUfYxnZko=">AAAB/XicbVC7TsMwFHXKq5RXADGxWFRITFXCAmwVLIxFIrRSE0WO47RWHTuynYoqqsSvsDAAYuU/2Pgb3DQDtBzpSsfn3Cvfe6KMUaUd59uqrayurW/UNxtb2zu7e/b+wYMSucTEw4IJ2YuQIoxy4mmqGellkqA0YqQbjW5mfndMpKKC3+tJRoIUDThNKEbaSKF95I8Jho/Qx7HQsHzk4Si0m07LKQGXiVuRJqjQCe0vPxY4TwnXmCGl+q6T6aBAUlPMyLTh54pkCI/QgPQN5SglKijK9afw1CgxTIQ0xTUs1d8TBUqVmqSR6UyRHqpFbyb+5/VznVwGBeVZrgnH84+SnEEt4CwLGFNJsGYTQxCW1OwK8RBJhLVJrGFCcBdPXibeeeuq5d45zfZ1lUYdHIMTcAZccAHa4BZ0gAcwKMAzeAVv1pP1Yr1bH/PWmlXNHII/sD5/AIQHlL4=</latexit>
5
projection onto uk
1) Project onto subspace spanned by k
Fitting the nonlinearity
<latexit sha1_base64="flj8ApT0XXAd006blhKZPNAqW50=">AAACDHicbZC9TsMwFIUdfkv5CzCyWBQkpiphAQakChbGIhFaqYkix3VaK44T2U6lKskLsPAqLAyAWHkANt4Gt80ALUey9Once3V9T5AyKpVlfRtLyyura+u1jfrm1vbOrrm3/yCTTGDi4IQlohsgSRjlxFFUMdJNBUFxwEgniG4m9c6ICEkTfq/GKfFiNOA0pBgpbfnmsTsiGGZ+BK+gGwqEc3eIFIzKvChmVBSlbzaspjUVXAS7ggao1PbNL7ef4CwmXGGGpOzZVqq8HAlFMSNl3c0kSRGO0ID0NHIUE+nl02tKeKKdPgwToR9XcOr+nshRLOU4DnRnjNRQztcm5n+1XqbCCy+nPM0U4Xi2KMwYVAmcRAP7VBCs2FgDwoLqv0I8RDoTpQOs6xDs+ZMXwTlrXjbtO6vRuq7SqIFDcAROgQ3OQQvcgjZwAAaP4Bm8gjfjyXgx3o2PWeuSUc0cgD8yPn8AlRabeQ==</latexit>
<latexit sha1_base64="O9GuRVuYx2v+9x/1eESUfYxnZko=">AAAB/XicbVC7TsMwFHXKq5RXADGxWFRITFXCAmwVLIxFIrRSE0WO47RWHTuynYoqqsSvsDAAYuU/2Pgb3DQDtBzpSsfn3Cvfe6KMUaUd59uqrayurW/UNxtb2zu7e/b+wYMSucTEw4IJ2YuQIoxy4mmqGellkqA0YqQbjW5mfndMpKKC3+tJRoIUDThNKEbaSKF95I8Jho/Qx7HQsHzk4Si0m07LKQGXiVuRJqjQCe0vPxY4TwnXmCGl+q6T6aBAUlPMyLTh54pkCI/QgPQN5SglKijK9afw1CgxTIQ0xTUs1d8TBUqVmqSR6UyRHqpFbyb+5/VznVwGBeVZrgnH84+SnEEt4CwLGFNJsGYTQxCW1OwK8RBJhLVJrGFCcBdPXibeeeuq5d45zfZ1lUYdHIMTcAZccAHa4BZ0gAcwKMAzeAVv1pP1Yr1bH/PWmlXNHII/sD5/AIQHlL4=</latexit>
2) take histogram of projected stimuli
6
STA response
1) Project onto subspace spanned by k
Fitting the nonlinearity
<latexit sha1_base64="flj8ApT0XXAd006blhKZPNAqW50=">AAACDHicbZC9TsMwFIUdfkv5CzCyWBQkpiphAQakChbGIhFaqYkix3VaK44T2U6lKskLsPAqLAyAWHkANt4Gt80ALUey9Once3V9T5AyKpVlfRtLyyura+u1jfrm1vbOrrm3/yCTTGDi4IQlohsgSRjlxFFUMdJNBUFxwEgniG4m9c6ICEkTfq/GKfFiNOA0pBgpbfnmsTsiGGZ+BK+gGwqEc3eIFIzKvChmVBSlbzaspjUVXAS7ggao1PbNL7ef4CwmXGGGpOzZVqq8HAlFMSNl3c0kSRGO0ID0NHIUE+nl02tKeKKdPgwToR9XcOr+nshRLOU4DnRnjNRQztcm5n+1XqbCCy+nPM0U4Xi2KMwYVAmcRAP7VBCs2FgDwoLqv0I8RDoTpQOs6xDs+ZMXwTlrXjbtO6vRuq7SqIFDcAROgQ3OQQvcgjZwAAaP4Bm8gjfjyXgx3o2PWeuSUc0cgD8yPn8AlRabeQ==</latexit>
<latexit sha1_base64="O9GuRVuYx2v+9x/1eESUfYxnZko=">AAAB/XicbVC7TsMwFHXKq5RXADGxWFRITFXCAmwVLIxFIrRSE0WO47RWHTuynYoqqsSvsDAAYuU/2Pgb3DQDtBzpSsfn3Cvfe6KMUaUd59uqrayurW/UNxtb2zu7e/b+wYMSucTEw4IJ2YuQIoxy4mmqGellkqA0YqQbjW5mfndMpKKC3+tJRoIUDThNKEbaSKF95I8Jho/Qx7HQsHzk4Si0m07LKQGXiVuRJqjQCe0vPxY4TwnXmCGl+q6T6aBAUlPMyLTh54pkCI/QgPQN5SglKijK9afw1CgxTIQ0xTUs1d8TBUqVmqSR6UyRHqpFbyb+5/VznVwGBeVZrgnH84+SnEEt4CwLGFNJsGYTQxCW1OwK8RBJhLVJrGFCcBdPXibeeeuq5d45zfZ1lUYdHIMTcAZccAHa4BZ0gAcwKMAzeAVv1pP1Yr1bH/PWmlXNHII/sD5/AIQHlL4=</latexit>
2) take histogram of projected stimuli
3) ML estimate of Poisson rate in each bin is # spikes / # stimuli
projection onto uk
7
<latexit sha1_base64="kzHLhkSGSdIE9++YRvFK95s20wQ=">AAAB/HicbVDLSsNAFJ3UV62v+Ni5GSyCCymJG3VXdOOygrGFJoTJ9KYdOnkwMynGUPwVNy5U3Poh7vwbp20W2nrgwuGce7n3niDlTCrL+jYqS8srq2vV9drG5tb2jrm7dy+TTFBwaMIT0QmIBM5icBRTHDqpABIFHNrB8Hrit0cgJEviO5Wn4EWkH7OQUaK05JsHboHdEVD84LNTnPvMHWPsm3WrYU2BF4ldkjoq0fLNL7eX0CyCWFFOpOzaVqq8ggjFKIdxzc0kpIQOSR+6msYkAukV0+vH+FgrPRwmQles8FT9PVGQSMo8CnRnRNRAznsT8T+vm6nwwitYnGYKYjpbFGYcqwRPosA9JoAqnmtCqGD6VkwHRBCqdGA1HYI9//Iicc4alw371qo3r8o0qugQHaETZKNz1EQ3qIUcRNEjekav6M14Ml6Md+Nj1loxypl99AfG5w8NApPa</latexit>
= stimuli and spike trains such that falls in bin j
<latexit sha1_base64="+nY3WdWTDls0W+HqppxAy1lUp3A=">AAACAHicbVC7TsMwFHXKq5RXgAWJxaJCYqoSFmCrYGEsEqFITRQ5jtNadezIdiqqqCz8CgsDIFY+g42/wU0zQMuRrnR8zr3yvSfKGFXacb6t2tLyyupafb2xsbm1vWPv7t0pkUtMPCyYkPcRUoRRTjxNNSP3mSQojRjpRsOrqd8dEamo4Ld6nJEgRX1OE4qRNlJoH/gjguFDSKGPY6Fh+czDIQztptNySsBF4lakCSp0QvvLjwXOU8I1ZkipnutkOiiQ1BQzMmn4uSIZwkPUJz1DOUqJCoryggk8NkoMEyFNcQ1L9fdEgVKlxmlkOlOkB2rem4r/eb1cJ+dBQXmWa8Lx7KMkZ1ALOI0DxlQSrNnYEIQlNbtCPEASYW1Ca5gQ3PmTF4l32rpouTdOs31ZpVEHh+AInAAXnIE2uAYd4AEMHsEzeAVv1pP1Yr1bH7PWmlXN7IM/sD5/AGsVlcQ=</latexit>
<latexit sha1_base64="1y/OQEwtflwAWPRa9KFhmR3i2GA=">AAAB73icbVA9T8MwFHwpX6V8FRhZLCokpiphAbYKFsYiEVrURpXjOK2p7US2g1RF/RUsDIBY+Tts/BvcNAO0nGTpdHdPfu/ClDNtXPfbqaysrq1vVDdrW9s7u3v1/YN7nWSKUJ8kPFHdEGvKmaS+YYbTbqooFiGnnXB8PfM7T1Rplsg7M0lpIPBQspgRbKz00Oc2GuHB46DecJtuAbRMvJI0oER7UP/qRwnJBJWGcKx1z3NTE+RYGUY4ndb6maYpJmM8pD1LJRZUB3mx8BSdWCVCcaLskwYV6u+JHAutJyK0SYHNSC96M/E/r5eZ+CLImUwzQyWZfxRnHJkEza5HEVOUGD6xBBPF7K6IjLDCxNiOarYEb/HkZeKfNS+b3q3baF2VbVThCI7hFDw4hxbcQBt8ICDgGV7hzVHOi/PufMyjFaecOYQ/cD5/ACdDkCo=</latexit>
= Poisson firing rate in bin j
Fitting the nonlinearity: derivation
Log-likelihood:<latexit sha1_base64="Wb2tlweIcDBGuHRLC0YxyG2IpmI=">AAACGHicbVBLSwMxGMzWV62vqkcvwSIIYtn1oh6EohcPHiq4ttBdlmyatrF5LElWWEr/hhf/ihcPKl5789+YtnuorQMJw8w3JN/ECaPauO6PU1haXlldK66XNja3tnfKu3uPWqYKEx9LJlUzRpowKohvqGGkmSiCeMxII+7fjP3GM1GaSvFgsoSEHHUF7VCMjJWisnsHr2CgUw5hFlEYMNm1l823UfQET2f4CcRSaBOVK27VnQAuEi8nFZCjHpVHQVvilBNhMENatzw3MeEAKUMxI8NSkGqSINxHXdKyVCBOdDiYbDaER1Zpw45U9ggDJ+psYoC41hmP7SRHpqfnvbH4n9dKTeciHFCRpIYIPH2okzJoJBzXBNtUEWxYZgnCitq/QtxDCmFjyyzZErz5lReJf1a9rHr3bqV2nbdRBAfgEBwDD5yDGrgFdeADDF7AG/gAn86r8+58Od/T0YKTZ/bBHzijX1fqnj8=</latexit>
<latexit sha1_base64="xOihw1X6fiePm2lfAdMk4gwa6P0=">AAACDXicbVC9TsMwGHT4LeUvwMhiUVViqhIWYECqYGFCRSK0UhNFjuO0bu0ksh2kKMoTsPAqLAyAWNnZeBvcNgO0nGTpdPedPn8XpIxKZVnfxtLyyuraem2jvrm1vbNr7u3fyyQTmDg4YYnoBUgSRmPiKKoY6aWCIB4w0g3GVxO/+0CEpEl8p/KUeBwNYhpRjJSWfLPpDpGCLtOJEPkjeAHdSCBcuDLjMPdpWdz4o9I3G1bLmgIuErsiDVCh45tfbpjgjJNYYYak7NtWqrwCCUUxI2XdzSRJER6jAelrGiNOpFdMzylhUyshjBKhX6zgVP2dKBCXMueBnuRIDeW8NxH/8/qZis68gsZppkiMZ4uijEGVwEk3MKSCYMVyTRAWVP8V4iHSdSjdYF2XYM+fvEick9Z5y761Gu3Lqo0aOARH4BjY4BS0wTXoAAdg8AiewSt4M56MF+Pd+JiNLhlV5gD8gfH5AxvMm7o=</latexit>
ML estimate: = # spikes / # stim
• piecewise constant model of nonlinearity f• more histogram bins ⇒ more flexible model
<latexit sha1_base64="sEXpHb6JayD4xH6qcmvpjVZVf2Q=">AAACEXicbVC7TsMwFHXKq5RXgJHFokJqlyphAbYKFsYiEVqpqSLHcVpTO4lsBxGFfgMLv8LCAIiVjY2/wWkzQMuRbB2dc6997/ETRqWyrG+jsrS8srpWXa9tbG5t75i7ezcyTgUmDo5ZLHo+koTRiDiKKkZ6iSCI+4x0/fFF4XfviJA0jq5VlpABR8OIhhQjpSXPbGYehQ/wXt+upBy6HKmR4HknplJOGi7TTwXIu21Cz6xbLWsKuEjsktRBiY5nfrlBjFNOIoUZkrJvW4ka5EgoihmZ1NxUkgThMRqSvqYR4kQO8ulKE3iklQCGsdAnUnCq/u7IEZcy476uLCaW814h/uf1UxWeDnIaJakiEZ59FKYMqhgW+cCACoIVyzRBWFA9K8QjJBBWOsWaDsGeX3mROMets5Z9ZdXb52UaVXAADkED2OAEtMEl6AAHYPAInsEreDOejBfj3fiYlVaMsmcf/IHx+QPFE50Z</latexit>
Model:
or <latexit sha1_base64="rJxGMAFxbCzCJXrF21ep8Vyz2wo=">AAACPnicbVC7TsMwFHXKq5RXgZHFUCGVgSphAQYkBAtjEbRFatrIcRzq4jiR7SAqkz9j4RfYWFkYALEy4rQV4nUkS+eec6+u7/ETRqWy7UerMDE5NT1TnC3NzS8sLpWXV5oyTgUmDRyzWFz4SBJGOWkoqhi5SARBkc9Iy786zv3WNRGSxvxcDRLSidAlpyHFSBnJKzfrVVcPPOpm8Ba6+iZnW/AAuomIA09T6FIOz7x+Bt1QIKydLO9eNyUzSwLk9bu5kEHS1dtfWuaVK3bNHgL+Jc6YVMAYda/84AYxTiPCFWZIyrZjJ6qjkVAUM5KV3FSSBOErdEnahnIUEdnRw/szuGmUAIaxMI8rOFS/T2gUSTmIfNMZIdWTv71c/M9rpyrc62jKk1QRjkeLwpRBFcM8TBhQQbBiA0MQFtT8FeIeMjkpE3nJhOD8PvkvaezU9mvOqV05PBqnUQRrYANUgQN2wSE4AXXQABjcgSfwAl6te+vZerPeR60FazyzCn7A+vgEOIuvWA==</latexit>
8
LNP (Linear-Nonlinear-Poisson) cascade model
Characterization Procedure:
1. Fit filter k using maximum likelihood under assumed nonlinearity.
2. Project stimuli onto k, compute spike rate (mean # spikes / stimulus) in each histogram bin.
stimulus filter Poissonspiking
stimulusk f λ(t)
exponentialnonlinearity
projection onto uk
9
• output: Poisson process• problem: assumes spiking depends only on stimulus!
stimulus filter Poissonspiking
stimulusk f λ(t)
conditional intensity(spike rate)
exponentialnonlinearity
LNP (Linear-Nonlinear-Poisson)cascade model
10
conditional intensity(spike rate)
(Truccolo et al 04)
• output: no longer a Poisson process
Poisson GLM with spike-history dependence
post-spike filter
exponentialnonlinearity
probabilisticspiking
stimulus
stimulus filter
+k
h
f
11
filter output
traditional IF
filter output
∞
“hard threshold”
“soft-threshold” IF
spik
e ra
te
• interpretation: “soft-threshold” integrate-and-fire model
Poisson GLM with spike-history dependence
post-spike filter
exponentialnonlinearity
probabilisticspiking
stimulus
stimulus filter
+k
h
f
12
GLM dynamic behaviors
post-spike filter h(t)
stimulus
p(spike)
• irregular spiking
filter outputs(“currents”)
13
GLM dynamic behaviors
post-spike filter h(t)
stimulus
p(spike)
• regular spiking
filter outputs(“currents”)
(Weber & Pillow 2016)
14
GLM dynamic behaviors
post-spike filter h(t)
• bursting
filter outputs(“currents”)
p(spike)
stimulus
15
GLM dynamic behaviors
post-spike filter h(t)
stimulus
filter outputs(“currents”)
p(spike)
• adaptation
16
GLM dynamic behaviors (from Izhikevich)A B C D
E F G H
I J K L
M N O P
tonic spiking phasic spiking tonic bursting phasic bursting
mixed mode type I type II
spike latency resonator integrator rebound spike
rebound burst variabilitybistability I bistability II
50 ms
spike frequencyadaptation
threshold
Figure 6: Suite of dynamical behaviors of Izhikevich and GLM neurons. Each panel,
top to bottom: stimulus (blue), Izhikevich neuron response (black), GLM responses
on five trials (gray), stimulus filter (left, blue), and post-spike filter (right, red). Black
line in each plot indicates a 50 ms scale bar for the stimulus and spike response.
(Differing timescales reflect timescales used for each behavior in original Izhikevich
paper (Izhikevich, 2004)). Stimulus filter and post-spike filter plots all have 100 ms
duration.
19
(Weber & Pillow 2017)
Izhikevich neuron
GLM spikes
stimulus
GLM parameters
17
multi-neuron GLM
exponentialnonlinearity
probabilisticspiking
stimulus
neuron 1
neuron 2
post-spike filter
stimulus filter
+
+
18
multi-neuron GLM
exponentialnonlinearity
probabilisticspiking
coupling filters
stimulus
neuron 1
neuron 2
post-spike filter
stimulus filter
+
+
19
...
time t
GLM equivalent diagram:
spike rate
20
Uzzell et al (J Neurophys 04)
• stimulus = binary flicker• parasol retinal ganglion cell spike responses
Example dataset
21
Uzzell et al (J Neurophys 04)
• stimulus = binary flicker• parasol retinal ganglion cell spike responses
Example dataset
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YX~k
time
time lag
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model
Stimulus-only GLM
spike responsedesign matrix
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time <latexit sha1_base64="T7110r2UjToEat5gBhsCYBUeHmw=">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</latexit>
model model
stimulus portion
spike-historyportion
Stimulus + SpikeHistory GLM
YX~k
spike responsedesign matrix
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time <latexit sha1_base64="T7110r2UjToEat5gBhsCYBUeHmw=">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</latexit>
model model
stimulus portion
neuron 1spike-hist
neuron 2spike-hist
neuron 3spike-hist
neuron 4spike-hist
YX~k
spike responsedesign matrix
Stimulus + History + 3 Neuron Coupling GLM
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Fitting: Maximum Likelihood
• find filters that maximize the log-conditional probability of the observed data
GLM parametersData
• log-likelihood is concave • no local maxima [Paninski 04]
logP (Y |X) =X
t
yt log �t � �t
• smooth basis for coupling filters
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• Concavity: having everywhere downward curvature
• Convexity: having everywhere upward curvature
• “f = concave” iff “-f = convex”
• Maximizing concave function = minimizing a convex function
• both preclude (non-global) local optima
convexity and concavity
concave convex
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• A function f is convex if, for any x1, x2, a in [0,1]:
• for continuous scalar functions:
• for vector functions: all eigenvalues of Hessian are ≥0, or
convexity: formal definition
properties of convex functions:• affine maps:
• sums:
• linear functions: concave and convex
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log-concavity of GLM likelihood
Theorem (Paninski 2004)
The GLM log-likelihood is concave in the parameters {k,h}, for any stimulus X and any spike data Y, if the nonlinearity f satisfies:
• f is convex• f is log-concave (i.e., log f(x) is a concave function)
Proof
weighted sum of concave funcs: concave
sum of convex funcs: convexconcave function
negative of convex func: concave
sum of two concave functions is concave ⇒ log-likelihood is concave!
log-likelihood
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log-concavity of GLM likelihood
Theorem (Paninski 2004)
The GLM log-likelihood is concave in the parameters {k,h}, for any stimulus X and any spike data Y, if the nonlinearity f satisfies:
• f is convex• f is log-concave (i.e., log f(x) is a concave function)
Examples of acceptable nonlinearities:
• • • condition: f must grow at least linearly and at most exponentially
Why this matters: no restriction on choice of stimuli! Can easily find ML fits to GLM parameters for any stimuli + spikes
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