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	<title>Physics 380, 2011: Lecture 9 - Revision history</title>
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		<summary type="html">&lt;p&gt;1 revision imported&lt;/p&gt;
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		<author><name>Ilya</name></author>
		
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		<title>nemenman&gt;Ilya: /* Warmup questions */</title>
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		<updated>2011-09-27T13:58:27Z</updated>

		<summary type="html">&lt;p&gt;‎&lt;span dir=&quot;auto&quot;&gt;&lt;span class=&quot;autocomment&quot;&gt;Warmup questions&lt;/span&gt;&lt;/span&gt;&lt;/p&gt;
&lt;p&gt;&lt;b&gt;New page&lt;/b&gt;&lt;/p&gt;&lt;div&gt;{{PHYS380-2011}}&lt;br /&gt;
&lt;br /&gt;
In these lectures, we cover some background on information theory. A good physics style introduction to this problem can be found in the upcoming book by Bialek (Bialek 2010). A very nice, and probably still the best, introduction to information theory as a theory of communication is (Shannow and Weaver, 1949). A standard and very good textbook on information theory is (Cover and Thomas, 2006).&lt;br /&gt;
&lt;br /&gt;
==Warmup questions==&lt;br /&gt;
#Does noise in signal transduction pathways affect information transmission?&lt;br /&gt;
#We would like to characterize how much information is transmitted by a cellular signaling pathway, say the NF-&amp;lt;math&amp;gt;\kappa&amp;lt;/math&amp;gt;B pathway depicted on the right (Cheong et al. 2011) , or in ''E. coli'' transcription (Guet et al., 2002; Ziv et al., 2007), as shown on the left. What characteristics of the system should we measure in order to be able to quantify this? Specifically, do we need:&lt;br /&gt;
#*&amp;lt;math&amp;gt;&amp;lt;r&amp;gt;&amp;lt;/math&amp;gt;, &amp;lt;math&amp;gt;&amp;lt;r|s&amp;gt;&amp;lt;/math&amp;gt; only?&lt;br /&gt;
#*&amp;lt;math&amp;gt;&amp;lt;r&amp;gt;&amp;lt;/math&amp;gt;, &amp;lt;math&amp;gt;&amp;lt;r|s&amp;gt;&amp;lt;/math&amp;gt;, and &amp;lt;math&amp;gt;\sigma^2_r&amp;lt;/math&amp;gt;, &amp;lt;math&amp;gt;\sigma^2_{r|s}&amp;lt;/math&amp;gt; only?&lt;br /&gt;
#*&amp;lt;math&amp;gt;P(r|s)&amp;lt;/math&amp;gt; for all s only?&lt;br /&gt;
#*&amp;lt;math&amp;gt;P(r|s)&amp;lt;/math&amp;gt; for all s and &amp;lt;math&amp;gt;P(s)&amp;lt;/math&amp;gt;, that is, the entire &amp;lt;math&amp;gt;P(r,s)&amp;lt;/math&amp;gt;?&lt;br /&gt;
&lt;br /&gt;
==Main lecture==&lt;br /&gt;
&lt;br /&gt;
*Setting up the problem: How do we measure information transmitted by a biological signaling system?&lt;br /&gt;
*Shannon's axioms and the derivation of entropy: if a variable &amp;lt;math&amp;gt;x&amp;lt;/math&amp;gt; is observed from a distribution &amp;lt;math&amp;gt;P(x)&amp;lt;/math&amp;gt; then the amount of the information we gain from this observation must obey the following properties.&lt;br /&gt;
*#If the cardinality of the distribution grows and the distribution is uniform, then the measure of information grows as well.&lt;br /&gt;
*#The measure of information must be a continuous function of the distribution &amp;lt;math&amp;gt;P(x)&amp;lt;/math&amp;gt;&lt;br /&gt;
*#The measure of information is additive. That is, for a fine graining of &amp;lt;math&amp;gt;x&amp;lt;/math&amp;gt; into &amp;lt;math&amp;gt;\xi&amp;lt;/math&amp;gt;, we should have &amp;lt;math&amp;gt;S[\xi]=S[x]+\sum P(x) S[\xi|x]&amp;lt;/math&amp;gt;.&lt;br /&gt;
Up to a multiplicative constant, the measure of information is then &amp;lt;math&amp;gt;S=-\sum P \log P&amp;lt;/math&amp;gt;, which is also called the Boltzman-Shannon entropy. And we fix the constant by defining the entropy of a uniform binary distribution to be 1. Then &amp;lt;math&amp;gt;S=-\sum P \log_2 P&amp;lt;/math&amp;gt;. The entropy is then measured in ''bits''.&lt;br /&gt;
*Meaning of entropy: Entropy of 1 bit means that we have gained enough information to answer one yes or no (binary) question about the variable &amp;lt;math&amp;gt;x&amp;lt;/math&amp;gt;.&lt;br /&gt;
*Properties of entropy (positive, limited, convex):&lt;br /&gt;
*#&amp;lt;math&amp;gt;0\le S[X]\le \log_2k&amp;lt;/math&amp;gt;, where &amp;lt;math&amp;gt;k&amp;lt;/math&amp;gt; is the cardinality of the distribution. Moreover, the first inequality becomes an equality iff the variable is deterministic (that is, one event has a probability of 1), and the second inequality is an equality iff the distribution is uniform.&lt;br /&gt;
*#Entropy is a convex function of the distribution&lt;br /&gt;
*#Entropies of independent variables add.&lt;br /&gt;
*#Entropy is an extensive quantity: for a joint distribution &amp;lt;math&amp;gt;P(x_1,x_2,\dots,x_n)&amp;lt;/math&amp;gt;, we can define an entropy ''rate'' &amp;lt;math&amp;gt;S_0=\lim_{n\to\infty} S[X_1,\dots,X_n]/n&amp;lt;/math&amp;gt;.&lt;br /&gt;
*Differential entropy: a continuous variable &amp;lt;math&amp;gt;x&amp;lt;/math&amp;gt; can be discretized with a step &amp;lt;math&amp;gt;\Delta x&amp;lt;/math&amp;gt;, and then the entropy is &amp;lt;math&amp;gt;S[X]=-\sum P(x)\Delta x\log_2 \left(P(x)\Delta x\right)\to \int dx P(x)\log_2P(x)  +\log_21/\Delta x&amp;lt;/math&amp;gt;. This formally diverges at fine discretization: we need infinitely many bits to fully specify a continuous variable. The integral in the above expression is called the ''differential entropy'', and whenever we write &amp;lt;math&amp;gt;S[X]&amp;lt;/math&amp;gt; for continuous variables, we mean the differential entropy.&lt;br /&gt;
*Entropy of a normal distribution with variance &amp;lt;math&amp;gt;\sigma^2&amp;lt;/math&amp;gt; is &amp;lt;math&amp;gt;S=1/2\log_2\sigma^2 + {\rm const}&amp;lt;/math&amp;gt;.&lt;br /&gt;
*Multivariate entropy is defined with summation/integration of log-probability over multiple variables, cf. entropy rate above.&lt;br /&gt;
*Conditional entropy is defined as averaged log-probability of a conditional distribution&lt;br /&gt;
*Mutual information: what if we want to know about a variable &amp;lt;math&amp;gt;x&amp;lt;/math&amp;gt;, but instead are measuring a variable &amp;lt;math&amp;gt;y&amp;lt;/math&amp;gt;. How much are we learning about &amp;lt;math&amp;gt;x&amp;lt;/math&amp;gt; then? This is given by the difference of entropies of &amp;lt;math&amp;gt;x&amp;lt;/math&amp;gt; before and after the measurement: &amp;lt;math&amp;gt;\begin{array}{ll}I[X;Y]&amp;amp;=S[X]-\langle S[X|Y]\rangle_y\\&amp;amp;=S[X]+S[Y]-S[X,Y]\\&amp;amp;=\langle\log_2\frac{P(x,y)}{P(x)P(y)}\end{array}&amp;lt;/math&amp;gt;.&lt;br /&gt;
*Meaning of mutual information: mutual information of 1 bit between two variables means that by querying one of them as much as possible, we can get one bit of information about the other.&lt;br /&gt;
*Properties of mutual information&lt;br /&gt;
*#Limits: &amp;lt;math&amp;gt;0\le I[X;Y]\le \min(S[X],S[X])&amp;lt;/math&amp;gt;. Note that the first inequality becomes an equality iff the two variables are completely statistically independent.&lt;br /&gt;
*#Mutual information is well-defined for continuous variables.&lt;br /&gt;
*#Reparameterization invariance: for any &amp;lt;math&amp;gt;\xi=\xi(x),\, \eta=\eta(y)&amp;lt;/math&amp;gt;, the following is true &amp;lt;math&amp;gt;I[X;Y]=I[\Xi;\Eta]&amp;lt;/math&amp;gt;.&lt;br /&gt;
*#Data processing inequality: For &amp;lt;math&amp;gt;P(x,y,z)=P(x)P(y|x)P(z|y)&amp;lt;/math&amp;gt;, &amp;lt;math&amp;gt;I[X;Z]\le \min (I[X;Y], I[Y;Z])&amp;lt;/math&amp;gt;. That is, information cannot get created in a transformation of a variable, whether deterministic or probabilistic.&lt;br /&gt;
*#Information rate: Information is also an extensive quantity, so that it makes sense to define an information rate &amp;lt;math&amp;gt;I_0=\lim_{n\to\infty}I[X_1,\dots,X_n;Y_1\dots Y_n]/n&amp;lt;/math&amp;gt;.&lt;br /&gt;
*Mutual information of a bivariate normal with a correlation coefficient &amp;lt;math&amp;gt;\rho&amp;lt;/math&amp;gt; is &amp;lt;math&amp;gt;I=1/2 \log_2(1-\rho^2)&amp;lt;/math&amp;gt;.&lt;br /&gt;
*For Gaussian variables &amp;lt;math&amp;gt;y=g(x+\eta)&amp;lt;/math&amp;gt;, where &amp;lt;math&amp;gt;x&amp;lt;/math&amp;gt; is the signal, &amp;lt;math&amp;gt;y&amp;lt;/math&amp;gt; is the response, and &amp;lt;math&amp;gt;\eta&amp;lt;/math&amp;gt; is the noise related to the input, &amp;lt;math&amp;gt;I[X;Y]=\frac{1}{2}\log_2\left(1+\frac{\sigma^2_x}{\sigma^2_\eta}\right)=\frac{1}{2}\log_2(1+SNR)&amp;lt;/math&amp;gt; (see the homework problem).&lt;/div&gt;</summary>
		<author><name>nemenman&gt;Ilya</name></author>
		
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