Maxwell's Demon: Implications for Evolution and Biogenesis Avshalom C. Elitzur Iyar, The Israeli Institute for Advanced Research Copyleft 2010 The Relevance of Thermodynamics to Life Sciences 1. Thermodynamics is a discipline that studies energy, entropy, and information Brillouin’s Information: Information=(Initial Uncertainty)–(Final Uncertainty) For several equally possible states, P0 I 0 With information reducing the possible states to P1: P0 I K ln K (ln P0 ln P1 ) P1 Ideally, for P1=1: I K ln P0 Shannon’s Information: Uncertainty = Entropy Boltzmann’s Entropy For all states being equiprobable: S k ln W Otherwise: w S k p1 ln p1 j i 27 Information of one English letter: i k p1 ln p1 j 1 For a string of G letters: jm I G i Gk p1 ln p1 j i The Relevance of Thermodynamics to Life Sciences 1. Thermodynamics is a discipline that studies energy, entropy, and information 2. Its jurisdiction is ubiquitous, regardless of the system’s chemical composition or type of energy Whence the entropy difference between animate and inanimate systems ? The Common Textbook Answer: “Living organisms are open systems” ? Open Systems: Rocks Chairs Blackboards Trash cans (!) etc. The Thesis: Adaptation = Information Maxwell’s Demon Attempts at Exorcizing 1. 2. 3. 4. Kelvin: The devil is alive Von Smoluchowski: It’s intelligent Szilard, Brillouin: It uses information Bennett & Landauer: It erases information Information and Energy Information Costs Energy ergo Information can Save Energy With information, you can do work with less energy, applied at the right time and/or place “Less energy, at the right time/place” “Less energy, at the right time/place”: Comparison between two methods of kill Considerable mechanical energy: Crushing the entire prey’s body Minute chemical energy: Neurotoxin (cobrotoxin) molecules reach the synapses with enormous precision The Demon Vs. the Living Organism: The Analogy 1) Life increases energy’s efficiency, up the thermodynamic scale 2) It does that with the aid of information Ek Et Ec Ee Et Ec + Ee Ec'> Ec Ec + Ee Ek The Demon Vs. the Living Organism: The Disanalogy 1) The real environment is never completely disordered but complex 2) The organism does not create order but complexity Ordered, Random, Complex Measures of Orderliness 1. Divergence from equiprobability (Gatlin) (Are there any digits in the sequence that are more common?) 2. Divergence from independence (Gatlin) (Is there any dependence between the digits?) 3. Redundancy (Chaitin) (Can the sequence be compressed into any shorter algorithm?) a. 3333333333333333333333333333333333333333333333333333333333 333333333333333333333333333333333333333333 1860271194945955774038867706591873856869843786230090655440 136901425331081581505348840600451256617983 0123456789012345678901234567890123456789012345678901234567 890123456789012345678901234567890123456789 6180339887498948482045868343656381177203091798057628621354 486227052604628189024497072072041893911374 b. c. d. 5 1 2 Sequence d is Sequence d is highly informative complex Bennett’s Measure of Complexity Given the shortest algorithm, how much computation is required to produce the sequence from it? And conversely: How much computation is required to encode a sequence into its shortest algorithm? complexity High order Low order The Ski-Lift Pathway: Thermodynamically Unique, Biologically Ubiquitous Goren Gordon & Avshalom C. Elitzur High Order Requires Energy Spontaneous Low Order How do you get to some desired state? High Order Requires Energy Spontaneous Initial State Low Order Desired State Step 1: Use Ski-Lift, get to the top How do you get to some desired state? High Order Requires Energy Spontaneous Initial State Low Order Desired State Step 1: Use Ski-Lift, get to the top How do you get to some desired state? High Order Requires Energy Spontaneous Step 1: Use Ski-Lift, get to the top Step 2: Ski down Initial State Low Order Desired State The Ski-Lift Conjecture (Gordon & Elitzur, 2009): Life approaches complexity “from above,” i.e., from the highorder state, and not “from below,” from the low-order state. Though the former route seems to require more energy, the latter requires immeasurable information, hence unrealistic energy. Dynamical evolution of complex states How to reach a complex state? 1. Probabilistic 2. Deterministic Entropy 1. Direct path Ski-lift 2. Ski-lift theorem Initial state Direct path Final state Direct Path Perform a transformation on the initial state to arrive at the final state Ti!f (???) Initial state unknown For each transformation only one initial state transforms to final state Hilbert Space Final state Initial state Direct Path: Probabilistic Perform a transformation on the initial state to arrive at the final state Ti!f (???) Initial state unknown For each transformation only one initial state transforms to final state Hilbert Space Perform transformation once Final state Energy cost: E= Probability of success: P=1/Ni=e-S(i)¿ 1 Initial state Direct Path: Deterministic Perform a transformation on the initial state to arrive at the final state Ti!f (???) Initial state unknown For each transformation only one initial state transforms to final state Repeat transformation until final state is reached Probability of success: P=1 Average energy cost: E= eS(i)À 1 Hilbert Space Final state Initial state Direct Path: Information Perform a transformation on the initial state to arrive at the final state Ti!f If one has information about initial state Ii=S(i) And information about final state (environment) If=S(f) Hilbert Space Then can perform the right transformation once Probability of success: P=1 Energy cost: E= Information required: I=S(i)+S(f) Final state Initial state Ski-lift Path Two stages path: Stage 1: Increase order S-i! order Ends with a specific, known state Probability of success: P1=1 Energy cost: E1=S(i) Hilbert Space Final state Initial state Ski-lift Path Two stages path: Stage 1: Increase order S-i! order Ends with a specific, known state Probability of success: P1=1 Energy cost: E1=S(i) Stage 2: Controlled transformation Torder!f Ends with the specific, final state Probability of success: P2=1 Energy cost: E2= Hilbert Space Final state Initial state Ski-lift Path: Information Requires information on final state (environment), in order to apply the right transformation on ordered-state Probability of success: P=1 Energy cost: Eski-lift=S(i)+ Information required: I=S(f) Hilbert Space Final state Initial state Comparison between paths Direct Path 1. Probabilistic 1. Low probability 2. Low energy 2. Deterministic: 1. High probability 2. High energy 3. Information: • • • • • Ski-lift Deterministic Controlled Reproducible Costs low energy Requires only environmental information 1. Requires much information 2. Low energy Ski-lift uses ordered-state and environmental information to obtain controllability and reproducibility How does Complexity Emerge? And How is it Maintained? Order Information/Complexity Disorder Bennett’s Measure of Complexity Given the shortest algorithm, how much computation is required to produce the sequence from it? And conversely: How much computation is required to encode a sequence into its shortest algorithm? complexity High order Low order Biological examples • • • • Cell formation Apoptosis Embryonic development Ecological development The Morphotropic State as the Cellular Progenitor of Complexity Minsky A, Shimoni E, Frenkiel-Krispin D. (2002) “Stress, Nat. Rev. Mol. Cell Biol. Jan;3(1):50-60. order and survival.” Maintaining the complexity of civilization necessitates huge reservoirs of order Schrödinger’s “What is life?” revisited Hilbert Space Requires energy High order Redundancy High complexity (specific environment) Requires information High entropy High information BIBLIOGRAPHY 1. Leff, H. S., & Rex, A. F. (2003) Maxwell’s Demon 2: Entropy, Classical and Quantum Information, Computing. Bristol: Institute of Physics Publishing. 2. Dill, K.A. , & Bromberg, S. (2003) Molecular Driving Forces: Statistical Thermodynamics in Chemistry and Biology. New York: Garland Science. 3. Di Cera, E., Ed. (2000) Thermodynamics in Biology” Oxford: Oxford University Press. 4. Gordon, G., & Elitzur, A. C. (2008) The Ski-Lift Pathway: Thermodynamically unique, biologically ubiquitous. http://www.a-c-elitzur.co.il/site/siteArticle.asp?ar=214
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