Topological Pressure of Finite Sequences David Koslicki July 2011 Penn State University Joint work with Dan Thompson Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Symbolic Dynamics Symbolic Dynamics Hadamard: 19th century Symbolic Dynamics Hadamard: 19th century Studying geodesics Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations T :X ØX Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations T :X ØX Smale’s Horseshoe Map Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations X= T :X ØX Smale’s Horseshoe Map Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations X= T :X ØX Smale’s Horseshoe Map Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations X= T :X ØX Smale’s Horseshoe Map Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations X= T :X ØX Smale’s Horseshoe Map Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations X= T T :X ØX Smale’s Horseshoe Map Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations X= A T B C D T :X ØX Smale’s Horseshoe Map Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations . X= A T B C D T :X ØX Smale’s Horseshoe Map Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations . X= A T B C D T :X ØX Smale’s Horseshoe Map . Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations . X= A B C D T :X ØX Smale’s Horseshoe Map . T . Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations . X= A B C D T :X ØX Smale’s Horseshoe Map . T . . Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations . X= A B C D T :X ØX Smale’s Horseshoe Map . T . A B C D . Symbolic Dynamics Hadamard: 19th century Studying geodesics Coding smooth transformations T :X ØX . X= A B C . D T Smale’s Horseshoe Map “CD” is an acceptable word . A B C D . Symbolic Dynamics Symbolic Dynamics = 8a 1 , . . . , a d < Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø Shift Map Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Shift Map Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Shift Map X Õ Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Shift Map X Õ Shift-invariant space Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Shift Map X Õ sH X L X Shift-invariant space Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Shift Map X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Complexity Function Shift Map X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Complexity Function Shift Map w œ X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Complexity Function Shift Map w œ X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X pw HnL # 8u : u n and u appears as a subword of w< Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Complexity Function Shift Map X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X w œ pw HnL # 8u : u n and u appears as a subword of w< Number of n-length subwords of w Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Complexity Function Shift Map X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X w œ pw HnL # 8u : u n and u appears as a subword of w< Number of n-length subwords of w Topological Entropy of a Sequence Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Complexity Function Shift Map X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X w œ pw HnL # 8u : u n and u appears as a subword of w< Number of n-length subwords of w Topological Entropy of a Sequence H top HwL lim nض log pw HnL n Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Complexity Function Shift Map X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X w œ pw HnL # 8u : u n and u appears as a subword of w< Number of n-length subwords of w Topological Entropy of a Sequence H top HwL lim log pw HnL nض Exponential growth rate of the number of n-length subwords as n tends to infinity n Symbolic Dynamics = 8a 1 , . . . , a d < Alphabet One-sided infinite sequences s : Ø sHHai Liœ L Hai+1 Liœ Complexity Function Shift Map X Õ Shift-invariant space H X , sL Symbolic Dynamical System sH X L X w œ pw HnL # 8u : u n and u appears as a subword of w< Number of n-length subwords of w Topological Entropy of a Sequence H top HwL lim log pw HnL nض Exponential growth rate of the number of n-length subwords as n tends to infinity n Traditional Topological Entropy Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… pw HnL n + 1 w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… pw HnL n + 1 Sturmian sequence w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… pw HnL n + 1 H top HwL 0 Sturmian sequence w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… pw HnL n + 1 H top HwL 0 Example 2: Full Shift Sturmian sequence w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… pw HnL n + 1 Sturmian sequence H top HwL 0 Example 2: Full Shift v baaaaabbbaabaababbabbbaababaababbaaaaaaa… w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… pw HnL n + 1 Sturmian sequence H top HwL 0 Example 2: Full Shift v baaaaabbbaabaababbabbbaababaababbaaaaaaa… pw HnL 2n w encodes trajectory Traditional Topological Entropy Example 1: Cutting Sequence of Square Billiard w abababaabababaababababaabababaababababaa… pw HnL n + 1 Sturmian sequence H top HwL 0 Example 2: Full Shift v baaaaabbbaabaababbabbbaababaababbaaaaaaa… pw HnL 2n H top HwL 1 w encodes trajectory Salient Properties: Salient Properties: 1. 0 § H top HwL § 1 Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable Issues with finite sequences: Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable Issues with finite sequences: pw HnL eventually zero Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable Issues with finite sequences: pw HnL eventually zero lim nض log pw HnL n =0 Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable Issues with finite sequences: pw HnL eventually zero lim log pw HnL nض So we need to evaluate at a single n… n =0 Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable Issues with finite sequences: pw HnL eventually zero lim log pw HnL nض So we need to evaluate at a single n… Shape of complexity function: n =0 Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable Issues with finite sequences: pw HnL eventually zero lim log pw HnL nض So we need to evaluate at a single n… Shape of complexity function: n =0 Salient Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable Issues with finite sequences: pw HnL eventually zero lim log pw HnL nض So we need to evaluate at a single n… Shape of complexity function: n =0 Adaptation to Finite Sequences Adaptation to Finite Sequences Shape of complexity function: Adaptation to Finite Sequences Shape of complexity function: For a finite sequence w, of length N there are integers m and M , such that the complexity function pw HnL is strictly increasing in the interval [0,m], non-decreasing in the interval [m,M] and strictly decreasing in the interval [M,N] (in fact pwtHn + 1L - pw HnL - 1 ). Adaptation to Finite Sequences Shape of complexity function: For a finite sequence w, of length N there are integers m and M , such that the complexity function pw HnL is strictly increasing in the interval [0,m], non-decreasing in the interval [m,M] and strictly decreasing in the interval [M,N] (in fact pwtHn + 1L - pw HnL - 1 ). Definition of Topological Entropy for Finite Sequences: Adaptation to Finite Sequences Shape of complexity function: For a finite sequence w, of length N there are integers m and M , such that the complexity function pw HnL is strictly increasing in the interval [0,m], non-decreasing in the interval [m,M] and strictly decreasing in the interval [M,N] (in fact pwtHn + 1L - pw HnL - 1 ). Definition of Topological Entropy for Finite Sequences: For w, a finite sequence over the alphabet (with d ), let n be the unique integer such that d n + n - 1 § w < d n+1 + Hn + 1L - 1 Then we define: Adaptation to Finite Sequences Shape of complexity function: For a finite sequence w, of length N there are integers m and M , such that the complexity function pw HnL is strictly increasing in the interval [0,m], non-decreasing in the interval [m,M] and strictly decreasing in the interval [M,N] (in fact pwtHn + 1L - pw HnL - 1 ). Definition of Topological Entropy for Finite Sequences: For w, a finite sequence over the alphabet (with d ), let n be the unique integer such that d n + n - 1 § w < d n+1 + Hn + 1L - 1 Then we define: H top HwL : logd I pwd1 n n +n -1 HnLM Adaptation to Finite Sequences Shape of complexity function: For a finite sequence w, of length N there are integers m and M , such that the complexity function pw HnL is strictly increasing in the interval [0,m], non-decreasing in the interval [m,M] and strictly decreasing in the interval [M,N] (in fact pwtHn + 1L - pw HnL - 1 ). Definition of Topological Entropy for Finite Sequences: For w, a finite sequence over the alphabet (with d ), let n be the unique integer such that d n + n - 1 § w < d n+1 + Hn + 1L - 1 Then we define: H top HwL : logd I pwd1 n n +n -1 HnLM Adaptation to Finite Sequences Adaptation to Finite Sequences Lemmas: Adaptation to Finite Sequences Lemmas: H top HwL : logd I pwd1 n n +n -1 HnLM Adaptation to Finite Sequences Lemmas: H top HwL : A sequence w of length d n + n - 1 over an alphabet with d letters can contain at most d n subwords of length n. logd I pwd1 n n +n -1 HnLM Adaptation to Finite Sequences Lemmas: H top HwL : A sequence w of length d n + n - 1 over an alphabet with d letters can contain at most d n subwords of length n. Conversely, for a sequence w, to contain d n subwords of n length n, it must have length d + n - 1 . logd I pwd1 n n +n -1 HnLM Adaptation to Finite Sequences Lemmas: H top HwL : A sequence w of length d n + n - 1 over an alphabet with d letters can contain at most d n subwords of length n. Conversely, for a sequence w, to contain d n subwords of n length n, it must have length d + n - 1 . Properties: logd I pwd1 n n +n -1 HnLM Adaptation to Finite Sequences H top HwL : Lemmas: A sequence w of length d n + n - 1 over an alphabet with d letters can contain at most d n subwords of length n. Conversely, for a sequence w, to contain d n subwords of n length n, it must have length d + n - 1 . Properties: 1. 0 § H top HwL § 1 logd I pwd1 n n +n -1 HnLM Adaptation to Finite Sequences H top HwL : Lemmas: A sequence w of length d n + n - 1 over an alphabet with d letters can contain at most d n subwords of length n. Conversely, for a sequence w, to contain d n subwords of n length n, it must have length d + n - 1 . Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) logd I pwd1 n n +n -1 HnLM Adaptation to Finite Sequences H top HwL : Lemmas: logd I pwd1 A sequence w of length d n + n - 1 over an alphabet with d letters can contain at most d n subwords of length n. Conversely, for a sequence w, to contain d n subwords of n length n, it must have length d + n - 1 . Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) n n +n -1 HnLM Adaptation to Finite Sequences H top HwL : Lemmas: logd I pwd1 n A sequence w of length d n + n - 1 over an alphabet with d letters can contain at most d n subwords of length n. Conversely, for a sequence w, to contain d n subwords of n length n, it must have length d + n - 1 . Properties: 1. 0 § H top HwL § 1 2. H top HwL º 0 iff w contains few subwords (simple) 3. H top HwL º 1 iff w contains many subwords (complex) 4. For different v and w, H top Hv L and H top HwL are comparable n +n -1 HnLM Examples H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 42 +2-1 w1 H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 42 +2-1 w1 w17 1 H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 42 +2-1 w1 w17 1 CT T CCT C A AGT CT C A AC H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T H top HwL : logd I pwd1 n +n -1 HnLM n w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 42 +2-1 w1 w17 1 CT T CCT C A AGT CT C A AC 8CT , TT , TC , CC , CT , TC , CA , AA , AG , GT , TC , CT , TC , CA , AA , AC < Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 42 +2-1 w1 w17 1 CT T CCT C A AGT CT C A AC 8CT , TT , TC , CC , CA , AA , AG , GT , AC < H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 42 +2-1 w1 w17 1 CT T CCT C A AGT CT C A AC 8CT , TT , TC, CC, CA, AA, AG, GT , AC < = 9 H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 42 +2-1 w1 w17 1 CT T CCT C A AGT CT C A AC 8CT , TT , TC , CC , CA , AA , AG , GT , AC < = 9 pw171 H2L 9 H top HwL : logd I pwd1 n n +n -1 HnLM Examples wCT T CCT C A AGT CT C A ACCGGT T H top HwL : w 22 17 42 + 2 - 1 § w < 43 + 3 - 1 66 n2 42 +2-1 w1 w17 1 CT T CCT C A AGT CT C A AC 8CT , TT , TC, CC, CA, AA, AG, GT , AC < = 9 pw171 H2L 9 H top HwL log4 I pw42 +2-1 H2LM 1 2 = log4 H9L 2 = .792481 logd I pwd1 n n +n -1 HnLM Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Topological Pressure Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: We say that a function y depends on m symbols if: Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: We say that a function y depends on m symbols if: For all v œ m the restriction of y to @vD is a constant function. Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: We say that a function y depends on m symbols if: For all v œ m the restriction of y to @vD is a constant function. There exists w œ Am-1 such that the restriction of y to @wD is not a constant function. Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: We say that a function y depends on m symbols if: For all v œ m the restriction of y to @vD is a constant function. There exists w œ Am-1 such that the restriction of y to @wD is not a constant function. Example: Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: We say that a function y depends on m symbols if: For all v œ m the restriction of y to @vD is a constant function. There exists w œ Am-1 such that the restriction of y to @wD is not a constant function. Example: yHwL w31 a Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: We say that a function y depends on m symbols if: For all v œ m the restriction of y to @vD is a constant function. There exists w œ Am-1 such that the restriction of y to @wD is not a constant function. Example: yHwL w31 a yHwL 1 ‚ wn1 n i ai Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: We say that a function y depends on m symbols if: For all v œ m the restriction of y to @vD is a constant function. There exists w œ Am-1 such that the restriction of y to @wD is not a constant function. Example: Subwords yHwL w31 a yHwL 1 ‚ wn1 n i ai Topological Pressure Intuition: Topological pressure captures the diversity of subwords. Topological pressure weights certain subwords as more/less important. Characterize “importance” through a function called the potential. Function that depends on m symbols: We say that a function y depends on m symbols if: For all v œ m the restriction of y to @vD is a constant function. There exists w œ Am-1 such that the restriction of y to @wD is not a constant function. Example: yHwL Subwords SWn HwL 8u : u n and u Õ w< w31 a yHwL 1 ‚ wn1 n i ai Topological Pressure Topological Pressure Definition of Topological Pressure for finite sequences: Topological Pressure Definition of Topological Pressure for finite sequences: For a word w such that w = n +n - 1 and a function y which depends on m symbols, with n ¥ m then the topological pressure is: Topological Pressure Definition of Topological Pressure for finite sequences: For a word w such that w = n +n - 1 and a function y which depends on m symbols, with n ¥ m then the topological pressure is: 1 PHw, yL log† § n ‚ uœ SWn HwL n-m exp ‚ yIsi uM i=0 Topological Pressure Definition of Topological Pressure for finite sequences: For a word w such that w = n +n - 1 and a function y which depends on m symbols, with n ¥ m then the topological pressure is: 1 PHw, yL log† § n For a w with n +n-1§ w < ‚ n-m uœ SWn HwL n +1 +n exp ‚ yIsi uM let: i=0 Topological Pressure Definition of Topological Pressure for finite sequences: For a word w such that w = n +n - 1 and a function y which depends on m symbols, with n ¥ m then the topological pressure is: 1 PHw, yL log† § n For a w with n +n-1§ w < ‚ n-m uœ SWn HwL n +1 PHw, yL PIw1 +n exp ‚ yIsi uM i=0 let: n +n-1 , yM Topological Pressure Topological Pressure Variational Principle: Topological Pressure Variational Principle: lim max PHw, yL sup :hm + ‡ y dm> nض w: w n m Topological Pressure Variational Principle: lim max PHw, yL sup :hm + ‡ y dm> nض w: w n Where m is a shift-invariant probability measure on m Topological Pressure Variational Principle: lim max PHw, yL sup :hm + ‡ y dm> nض w: w n Where m is a shift-invariant probability measure on and hm is the metric entropy of m m Topological Pressure Variational Principle: lim max PHw, yL sup :hm + ‡ y dm> nض w: w n Where m is a shift-invariant probability measure on and hm is the metric entropy of m Miscellanea: m Topological Pressure Variational Principle: lim max PHw, yL sup :hm + ‡ y dm> nض w: w n m Where m is a shift-invariant probability measure on and hm is the metric entropy of m Miscellanea: For y log j with j > 0 a function that depends on m symbols, we have Topological Pressure Variational Principle: lim max PHw, yL sup :hm + ‡ y dm> nض w: w n m Where m is a shift-invariant probability measure on and hm is the metric entropy of m Miscellanea: For y log j with j > 0 a function that depends on m symbols, we have PHw, yL 1 log† § n ‚ n-m ‰ jIsi uM uœ SWn HwL i=0 Topological Pressure Variational Principle: lim max PHw, yL sup :hm + ‡ y dm> nض w: w n m Where m is a shift-invariant probability measure on and hm is the metric entropy of m Miscellanea: For y log j with j > 0 a function that depends on m symbols, we have PHw, yL 1 log† § n Scaling of a potential: ‚ n-m ‰ jIsi uM uœ SWn HwL i=0 Topological Pressure Variational Principle: lim max PHw, yL sup :hm + ‡ y dm> nض w: w n m Where m is a shift-invariant probability measure on and hm is the metric entropy of m Miscellanea: For y log j with j > 0 a function that depends on m symbols, we have PHw, yL 1 log† § n ‚ n-m ‰ jIsi uM uœ SWn HwL i=0 Scaling of a potential: PHw, logHt jLL n-m log† § Ht L + PHw, log jL n Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Application to DNA Sequences Application to DNA Sequences DNA: Application to DNA Sequences DNA: Application to DNA Sequences DNA: Alphabet: 8 A, C, T , G < Application to DNA Sequences DNA: Alphabet: 8 A, C, T , G < Application to DNA Sequences DNA: Alphabet: 8 A, C, T , G < Human genome: 3,080,000,000 bp Application to DNA Sequences DNA: Alphabet: 8 A, C, T , G < Human genome: 3,080,000,000 bp Only 2% codes for genes (gene length 30002.4 million bp) Application to DNA Sequences Application to DNA Sequences Introns, Exons, and Genes: Application to DNA Sequences Introns, Exons, and Genes: Exons: translated to a protein (gene) Introns: junk? Application to DNA Sequences Introns, Exons, and Genes: Exons: translated to a protein (gene) Introns: junk? Application to DNA Sequences Introns, Exons, and Genes: Exons: translated to a protein (gene) Introns: junk? Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Application to DNA Sequences Application to DNA Sequences Application to Introns and Exons: (D. Koslicki, Bioinformatics, 2011) Application to DNA Sequences Application to Introns and Exons: (D. Koslicki, Bioinformatics, 2011) How well does topological entropy distinguish between introns and exons? Application to DNA Sequences Application to Introns and Exons: (D. Koslicki, Bioinformatics, 2011) How well does topological entropy distinguish between introns and exons? Is the entropy of introns higher or lower than that of exons? Application to DNA Sequences How well does topological entropy distinguish between introns and exons? Application to Introns and Exons: (D. Koslicki, Bioinformatics, 2011) Is the entropy of introns higher or lower than that of exons? LC 1.00 0.99 0.98 Exons Chromosome Chr Y Chr X Chr 22 Chr 21 Chr 19 Chr 18 Chr 17 Chr 16 Chr 15 Chr 14 Chr 13 Chr 12 Chr 11 Chr 10 Chr 9 Chr 8 Chr 7 Chr 6 Chr 5 Chr 4 Chr 3 Chr 2 Chr 1 Chr 20 Introns 0.97 Application to DNA Sequences How well does topological entropy distinguish between introns and exons? Application to Introns and Exons: (D. Koslicki, Bioinformatics, 2011) Is the entropy of introns higher or lower than that of exons? Htop 0.94 LC 0.93 1.00 0.92 0.91 0.99 0.90 0.98 0.89 Exons Introns Exons 0.87 Chr Y Chr X Chr 22 Chr 21 Chr 20 Chr 19 Chr 18 Chr 17 Chr 16 Chr 15 Chr 14 Chr 13 Chr 12 Chr 11 Chr 10 Chr 9 Chr 8 Chr 7 Chr 6 Chr 5 Chr 4 Chr 3 Chr 2 Chr 1 Introns Chromosome 0.88 0.97 Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Gene Distribution Detection Gene Distribution Detection Gene Detection Simplified: Gene Distribution Detection Gene Detection Simplified: 1. Use all 6 open reading frames, look for “stop” and “start” amino acids. 2. Use known genes with pattern matching and neural networks. Gene Distribution Detection Gene Detection Simplified: 1. Use all 6 open reading frames, look for “stop” and “start” amino acids. 2. Use known genes with pattern matching and neural networks. Must have previously known genes Gene Distribution Detection Gene Detection Simplified: 1. Use all 6 open reading frames, look for “stop” and “start” amino acids. 2. Use known genes with pattern matching and neural networks. Must have previously known genes Not very good at finding novel genes Gene Distribution Detection Gene Detection Simplified: 1. Use all 6 open reading frames, look for “stop” and “start” amino acids. 2. Use known genes with pattern matching and neural networks. Must have previously known genes Not very good at finding novel genes Gene Distribution: Gene Distribution Detection Gene Detection Simplified: 1. Use all 6 open reading frames, look for “stop” and “start” amino acids. 2. Use known genes with pattern matching and neural networks. Must have previously known genes Not very good at finding novel genes Gene Distribution: Given a novel genome, estimate the distribution of genes (or coding sequences) Gene Distribution Detection Gene Detection Simplified: 1. Use all 6 open reading frames, look for “stop” and “start” amino acids. 2. Use known genes with pattern matching and neural networks. Must have previously known genes Not very good at finding novel genes Gene Distribution: Given a novel genome, estimate the distribution of genes (or coding sequences) Gene Distribution Detection Gene Detection Simplified: 1. Use all 6 open reading frames, look for “stop” and “start” amino acids. 2. Use known genes with pattern matching and neural networks. Must have previously known genes Not very good at finding novel genes Gene Distribution: Given a novel genome, estimate the distribution of genes (or coding sequences) Ensemble genome browser Gene Distribution Detection Gene Distribution Detection Potentials: Gene Distribution Detection Potentials: Restrict attention to y = log j which depend on 3 symbols. Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? When is a sequence “important”? Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? When is a sequence “important”? Codons, Proteins… Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? When is a sequence “important”? Procedure: Codons, Proteins… Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? Codons, Proteins… When is a sequence “important”? Procedure: Obtain a potential via maximizing the correlation of PHw, log jL with a given set of biological data. Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? Codons, Proteins… When is a sequence “important”? Procedure: Obtain a potential via maximizing the correlation of PHw, log jL with a given set of biological data. Analyze biological relevance Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? Codons, Proteins… When is a sequence “important”? Procedure: Obtain a potential via maximizing the correlation of PHw, log jL with a given set of biological data. Analyze biological relevance Compare to other possible potentials Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? Codons, Proteins… When is a sequence “important”? Procedure: Obtain a potential via maximizing the correlation of PHw, log jL with a given set of biological data. Analyze biological relevance Compare to other possible potentials Intuition: Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? Codons, Proteins… When is a sequence “important”? Procedure: Obtain a potential via maximizing the correlation of PHw, log jL with a given set of biological data. Analyze biological relevance Compare to other possible potentials Intuition: View a DNA sequence as concatenations of symbolic dynamical systems, and then use pressure to differentiate between the systems. Gene Distribution Detection Potentials: Problem: Restrict attention to y = log j which depend on 3 symbols. Which potential to use? Codons, Proteins… When is a sequence “important”? Procedure: Obtain a potential via maximizing the correlation of PHw, log jL with a given set of biological data. Analyze biological relevance Compare to other possible potentials Intuition: View a DNA sequence as concatenations of symbolic dynamical systems, and then use pressure to differentiate between the systems. Gene Distribution Detection Gene Distribution Detection Sliding Window Output: Gene Distribution Detection Sliding Window Output: Chromosome 5 (180,915,260 bp) Window size 65543 Gene Distribution Detection Sliding Window Output: Chromosome 5 (180,915,260 bp) Window size 65543 TopologicalPressure 90 85 80 75 70 65 60 500 1000 1500 2000 2500 Window Location Gene Distribution Detection Gene Distribution Detection Smoothing: Gene Distribution Detection Smoothing: Convolution with a Gaussian kernel Moving Average Gaussian Filter Etc. all give similar results Gene Distribution Detection Smoothing: Convolution with a Gaussian kernel Moving Average Gaussian Filter Etc. all give similar results Radius selection based on Gaussian kernel density estimation with bandwidth selected according to Silverman’s rule Gene Distribution Detection Smoothing: Convolution with a Gaussian kernel Moving Average Gaussian Filter Etc. all give similar results Radius selection based on Gaussian kernel density estimation with bandwidth selected according to Silverman’s rule TopologicalPressure 77.5 77.0 76.5 76.0 500 1000 1500 2000 2500 Window Location Gene Distribution Detection Smoothing: Convolution with a Gaussian kernel Moving Average Gaussian Filter Etc. all give similar results TopologicalPressure Radius selection based on Gaussian kernel density estimation with bandwidth selected according to Silverman’s rule Window size of 65,543 on Chr5 77.5 77.0 76.5 76.0 500 1000 1500 2000 2500 Window Location Gene Distribution Detection Gene Distribution Detection Correlation between pressure and CDS distribution: Gene Distribution Detection Correlation between pressure and CDS distribution: Hg18, March 2006 assembly. Window size: 65543 Gene Distribution Detection Correlation between pressure and CDS distribution: Hg18, March 2006 assembly. Window size: 65543 ndardized Value 2 1 500 1000 1500 2000 2500 Window Location -1 Topological Pressure Known Coding Sequence Density Gene Distribution Detection Correlation between pressure and CDS distribution: ndardized Value Hg18, March 2006 assembly. Window size: 65543 Correlation: 0.979481 2 1 500 1000 1500 2000 2500 Window Location -1 Topological Pressure Known Coding Sequence Density Gene Distribution Detection Correlation between pressure and CDS distribution: ndardized Value Hg18, March 2006 assembly. Window size: 65543 Correlation: 0.979481 2 1 500 1000 1500 2000 2500 Window Location -1 Topological Pressure Known Coding Sequence Density Gene Distribution Detection Gene Distribution Detection Comparative Analysis: Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Similar parameter values Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Similar parameter values Euclidean Distance Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Similar parameter values Euclidean Distance Pairwise Euclidean Distance 1 5 10 15 20 24 1 1 5 5 10 10 15 15 20 20 24 24 1 5 10 15 20 24 Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Similar parameter values Euclidean Distance Pairwise Euclidean Distance 1 5 10 15 20 24 1 1 5 5 10 10 15 15 20 20 24 24 1 5 10 15 Max: 0.252 (chrY, chr7) 20 24 Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Similar parameter values Euclidean Distance Pairwise Euclidean Distance 1 5 10 15 20 24 1 1 5 5 10 10 15 15 20 20 24 24 1 5 10 15 Max: 0.252 (chrY, chr7) Mean: 0.1815 20 24 Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Similar parameter values Euclidean Distance Pairwise Euclidean Distance 1 5 10 15 20 24 1 1 5 5 10 10 15 15 20 20 24 24 1 5 10 15 Max: 0.252 (chrY, chr7) Mean: 0.1815 Expected: 1.41421 20 24 Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Similar parameter values Euclidean Distance Pairwise Euclidean Distance 1 5 10 15 20 24 1 1 5 5 10 10 15 15 20 20 24 24 1 5 10 15 Max: 0.252 (chrY, chr7) Mean: 0.1815 Expected: 1.41421 20 24 Median Parameter Values: Gene Distribution Detection Comparative Analysis: 24 potentials with 64 variables each Similar parameter values Euclidean Distance Pairwise Euclidean Distance 1 5 10 15 20 24 1 1 5 5 10 10 15 15 20 Median Parameter Values: 2nd Base U UUU HPheL U UUC HPheL C UCU HSerL UCC HSerL A UAU HTyrL UAC HTyrL G UGU HCysL UGC HCysL UUA HLeuL UCA HSerL UAA Stop UGA Stop UUG HLeuL UCG HSerL UAG Stop UGG HTrpL CUU HLeuL CCU HProL CAU HHisL CGU HArgL C CUC HLeuL CCC HProL CAC HHisL CGC HArgL CUA HLeuL CCA HProL CAA HGlnL CGA HArgL CUG HLeuL CCG HProL CAG HGlnL CGG HArgL 1st AUU HIleL baseA AUC HIleL ACU HThrL ACC HThrL AAU HAsnL AAC HAsnL AGU HSerL AGC HSerL AUA HIleL AUG HMetL GUU HValL G GUC HValL ACA HThrL ACG HThrL GCU HAlaL GCC HAlaL AAA HLysL AAG HLysL GAU HAspL GAC HAspL AGA HArgL AGG HArgL GGU HGlyL GGC HGlyL GUA HValL GCA HAlaL GAA HGluL GGA HGlyL GUG HValL GCG HAlaL GAG HGluL GGG HGlyL 20 24 24 1 5 10 15 Max: 0.252 (chrY, chr7) Mean: 0.1815 Expected: 1.41421 20 24 Unit Square Gene Distribution Detection Gene Distribution Detection Compared to Mouse Genome: Gene Distribution Detection Compared to Mouse Genome: Use the median potential j obtained before Gene Distribution Detection Compared to Mouse Genome: Use the median potential j obtained before Compare associated sliding window PHw, log jL to the mouse genome (chromosome 1) Gene Distribution Detection Compared to Mouse Genome: Standardized Value Use the median potential j obtained before Compare associated sliding window PHw, log jL to the mouse genome (chromosome 1) Topological Pressure Known mm Coding Sequence Density 2 1 500 -1 -2 1000 1500 2000 2500 Window Location Gene Distribution Detection Compared to Mouse Genome: Standardized Value Use the median potential j obtained before Compare associated sliding window PHw, log jL to the mouse genome (chromosome 1) Topological Pressure Known mm Coding Sequence Density 2 1 500 -1 -2 1000 1500 2000 2500 Window Location Correlation: 0.827 Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Topological Entropy of Finite Sequences Topological Entropy Topological Pressure Application to DNA Sequences Intron and Exons Gene Distribution Detection Equilibrium Measures Measure of Coding Potential Measure of Coding Potential Measure from parameter values: Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 Conceptually: PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Conceptually: m favors sequences similarly to log j while still respecting entropy constraints. Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Conceptually: m favors sequences similarly to log j while still respecting entropy constraints. Rigorously: Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Conceptually: m favors sequences similarly to log j while still respecting entropy constraints. Rigorously: Gibbs measure (in the sense of Bowen) Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Conceptually: m favors sequences similarly to log j while still respecting entropy constraints. Rigorously: Gibbs measure (in the sense of Bowen) $ M s.t. M -1 mH@wDL § §M n-3 i exp 9-nPH , log jL + ⁄i=0 log jHs wL= Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Conceptually: m favors sequences similarly to log j while still respecting entropy constraints. Rigorously: Gibbs measure (in the sense of Bowen) $ M s.t. M So, -1 mH@wDL § §M n-3 i exp 9-nPH , log jL + ⁄i=0 log jHs wL= Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Conceptually: m favors sequences similarly to log j while still respecting entropy constraints. Rigorously: Gibbs measure (in the sense of Bowen) $ M s.t. M -1 mH@wDL § §M n-3 i exp 9-nPH , log jL + ⁄i=0 log jHs wL= So, n-3 mH@wDL º ‰jIsi wM i=0 Measure of Coding Potential Measure from parameter values: Potential that depends on the first m letters gives ris to a Markov measure m of memory m-1. m is the equilibrium measure for log j max n w: w 4 +n-1 PHw, log jL sup :hm + ‡ log j dm> = hm + ‡ log j dm m Conceptually: m favors sequences similarly to log j while still respecting entropy constraints. Rigorously: Gibbs measure (in the sense of Bowen) $ M s.t. M -1 mH@wDL § §M n-3 i exp 9-nPH , log jL + ⁄i=0 log jHs wL= So, n-3 mH@wDL º ‰jIsi wM i=0 Utilizing The Measure m Utilizing The Measure m Single sequence measure of coding potential: Utilizing The Measure m Single sequence measure of coding potential: Given a single (short) sequence, measure the likelihood that said sequence is (part of) a coding sequence. Utilizing The Measure m Single sequence measure of coding potential: Given a single (short) sequence, measure the likelihood that said sequence is (part of) a coding sequence. Comparative measures have enjoyed success (RNAcode, Washietl et al. 2011) Utilizing The Measure m Single sequence measure of coding potential: Given a single (short) sequence, measure the likelihood that said sequence is (part of) a coding sequence. Comparative measures have enjoyed success (RNAcode, Washietl et al. 2011) Lin, Jungreis, Kellis (Bioinformatics, 2011): “Measures of coding potential based on primary sequence composition are still lacking “ Utilizing The Measure m Single sequence measure of coding potential: Given a single (short) sequence, measure the likelihood that said sequence is (part of) a coding sequence. Comparative measures have enjoyed success (RNAcode, Washietl et al. 2011) Lin, Jungreis, Kellis (Bioinformatics, 2011): “Measures of coding potential based on primary sequence composition are still lacking “ Current Work: Utilizing The Measure m Single sequence measure of coding potential: Given a single (short) sequence, measure the likelihood that said sequence is (part of) a coding sequence. Comparative measures have enjoyed success (RNAcode, Washietl et al. 2011) Lin, Jungreis, Kellis (Bioinformatics, 2011): “Measures of coding potential based on primary sequence composition are still lacking “ Current Work: Evaluating usefulness of the measure m in determining coding potential of short sequences Summary Summary Topological Pressure: Summary Topological Pressure: Leads to new biological insight Summary Topological Pressure: Leads to new biological insight Relevant thermodynamics properties hold Summary Topological Pressure: Leads to new biological insight Relevant thermodynamics properties hold Symbolic dynamics fruitful when applied to biology Summary Topological Pressure: Leads to new biological insight Relevant thermodynamics properties hold Symbolic dynamics fruitful when applied to biology Accurately approximates coding sequence density distribution Summary Topological Pressure: Leads to new biological insight Relevant thermodynamics properties hold Symbolic dynamics fruitful when applied to biology Accurately approximates coding sequence density distribution Clear and comprehensible application of thermodynamic concepts Thank you
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