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Selection, large deviations and metastability () Dynamics with selection, large deviations and metastability 1 / 35 1. Dynamics with selection () Dynamics with selection, large deviations and metastability 2 / 35 A cell performs complex


  1. Selection, large deviations and metastability () Dynamics with selection, large deviations and metastability 1 / 35

  2. 1. Dynamics with selection () Dynamics with selection, large deviations and metastability 2 / 35

  3. A cell performs complex dynamics: DNA codes for the production of proteins, which themselves modify how the reading is done. A bit like a program and its RAM content. DNA contains about the same amount of information as the TeXShop program for Mac This dynamics admits more than one attractor: same DNA yields liver and eye cells... The dynamical state is inherited. On top of this process, there is the selection associated to the death and reproduction of individual cells () Dynamics with selection, large deviations and metastability 3 / 35

  4. Stern, Dror, Stolovicki, Brenner, and Braun An arbitrary and dramatic rewiring of the genome of a yeast cell: the presence of glucose causes repression of histidine biosynthesis, an essential process Cells are brutally challenged in the presence of glucose, nothing in evolution prepared them for that! () Dynamics with selection, large deviations and metastability 4 / 35

  5. Stern, Dror, Stolovicki, Brenner, and Braun () Dynamics with selection, large deviations and metastability 5 / 35

  6. Stern, Dror, Stolovicki, Brenner, and Braun () Dynamics with selection, large deviations and metastability 6 / 35

  7. the system finds a transcriptional state with many changes two realizations of the experiment yield vastly different solutions the same dynamical system seems to have chosen a different attractor which is then inherited over many generations () Dynamics with selection, large deviations and metastability 7 / 35

  8. If this interpretation is confirmed, we are facing a dynamics in a complex landscape with the added element of selection but note that fitness does not drive the dynamics, it acts on its results the landscape is not the ‘fitness landscape’ () Dynamics with selection, large deviations and metastability 8 / 35

  9. 2. The relation between a) Large Deviations, b) Metastability c) Dynamics with selection and phase transitions () Dynamics with selection, large deviations and metastability 9 / 35

  10. a pendulum immersed in a low-temperature bath () Dynamics with selection, large deviations and metastability 10 / 35

  11. a pendulum immersed in a low-temperature bath θ () Dynamics with selection, large deviations and metastability 11 / 35

  12. Imposing the average angle, the trajectory shares its time between saddles 0 o and 180 o Θ(τ) 90 θ 0 τ phase-separation is a first order transition! () Dynamics with selection, large deviations and metastability 12 / 35

  13. �� t � � 0 θ ( t ′ ) dt ′ − tθ o D [ θ ] P ( trajectory ) δ � � t � 0 θ ( t ′ ) dt ′ D [ θ ] P ( trajectory ) e λ e − λtθ o = dλ � �� � canonical canonical version, with λ conjugated to θ � t � 0 θ ( t ′ ) dt ′ D [ θ ] P ( trajectory ) e λ Z ( λ ) = () Dynamics with selection, large deviations and metastability 13 / 35

  14. • λ is fixed to give the appropriate θ (Laplace transform variable) • a system of walkers with cloning rate λθ ( t ) � d � �� dP T d dt = − dθ + sin( θ ) P − λθ P dθ yields the ‘canonical’ version of the large-deviation function () Dynamics with selection, large deviations and metastability 14 / 35

  15. • the relation is useful for efficient simulations • but also to understand the large deviation function () Dynamics with selection, large deviations and metastability 15 / 35

  16. We wish to simulate an event with an unusually large value of A without having to wait for this to happen spontaneously but without forcing the situation artificially () Dynamics with selection, large deviations and metastability 16 / 35

  17. N independent simulations x x with probability c . A per unit time kill or clone ... continue ... a way to count trajectories weighted with e cA () Dynamics with selection, large deviations and metastability 17 / 35

  18. • A collection of metastable states • each with its own emigration rate • and its cloning/death rates dependent upon the observable We understand the relation between metastability and large deviations () Dynamics with selection, large deviations and metastability 18 / 35

  19. Dynamical phase transitions large deviations of the activity JP Garrahan, RL Jack, V Lecomte, E Pitard, K van Duijvendijk, and Frederic van Wijland () Dynamics with selection, large deviations and metastability 19 / 35

  20. may be obtained with selection proportional to the activity () Dynamics with selection, large deviations and metastability 20 / 35

  21. 3. Large deviations and order () Dynamics with selection, large deviations and metastability 21 / 35

  22. What do extremal trajectories look like? () Dynamics with selection, large deviations and metastability 22 / 35

  23. The baker’s map p p ���� ���� ���� ���� ���� ���� ���� ���� ������ ������ ������ ������ ���� ���� ������ ������ ������ ������ ���� ���� ������ ������ ������ ������ q q ... is as chaotic as you can be. () Dynamics with selection, large deviations and metastability 23 / 35

  24. And yet, orbits minimising a function, e.g. � dt ( q ( t ) − q ∗ ) 2 A ≡ ρ = 3 / 13 ≈ 0 . 231 ρ = 5 / 17 ≈ 0 . 294 q ∗ ∈ [0 . 2360 , 0 . 2362] q ∗ ∈ [0 . 29395 , 0 . 29397] are periodic or quasiperiodic but unstable! Hunt and Ott — Khan-Dang Nguyen Thu Lam, JK , D Levine () Dynamics with selection, large deviations and metastability 24 / 35

  25. In a case like this one: T. Duriez, J..L. Aider, E. Masson, J.E. Wesfreid ; Qualitative investigation of the main flow features over a TGV ; Proceedings of the Euromech Colloquium 509, Vehicle Aerodynamics, Berlin, Allemagne, 2009 , p. 52-57 http://opus.kobv.de/tuberlin/volltexte/2009/2249/ � τ ¯ f τ = 1 0 f ( t ) dt τ () Dynamics with selection, large deviations and metastability 25 / 35

  26. if this metaphor is good, we should see order during exceptional times () Dynamics with selection, large deviations and metastability 26 / 35

  27. 4. Miscellanea advertisement for J. Tailleur’s talk () Dynamics with selection, large deviations and metastability 27 / 35

  28. because planets disturb one another, the dynamics is chaotic () Dynamics with selection, large deviations and metastability 28 / 35

  29. chaotic? 1 Lyapunov time = λ 2.71 ∆ ∆ difference between trajectories multiplies by e = 2 . 71 ... every ∼ 5 M years Laskar λ ≡ 1 / 5 MY rs is called the Lyapunov exponent λ > 0 → chaos () Dynamics with selection, large deviations and metastability 29 / 35

  30. Consider this: If you start a planetary random system in your computer, it often runs into trouble. If you observe a planetary system, many conditions within the observational error imply recent formation or immediate destruction Are there places between large planets where earth-like planets may have relative stability? You need to know rare trajectories () Dynamics with selection, large deviations and metastability 30 / 35

  31. Laskar et al () Dynamics with selection, large deviations and metastability 31 / 35

  32. The problem of transitions peptide helix-coil transition () Dynamics with selection, large deviations and metastability 32 / 35

  33. The problem of transitions may be shown to be a problem of large deviation functions of the (largest) Lyapunov exponent! () Dynamics with selection, large deviations and metastability 33 / 35

  34. Drag, traffic jams, etc: probability larger times (or sizes) average drag � τ f τ = 1 ¯ 0 f ( t ) dt τ () Dynamics with selection, large deviations and metastability 34 / 35

  35. Intermittency in fully developed turbulence: Longitudinal-structure functions p ln | v ( x + R ) − v ( x ) | S p ( R ) = �| v ( x + R ) − v ( x ) | p � = � e � �� � � who is responsible for the large moments? () Dynamics with selection, large deviations and metastability 35 / 35

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