Stochastic Dynamic Programming Illuminates the Link Between Environment, Physiology, and Evolution

I describe how stochastic dynamic programming (SDP), a method for stochastic optimization that evolved from the work of Hamilton and Jacobi on variational problems, allows us to connect the physiological state of organisms, the environment in which they live, and how evolution by natural selection acts on trade-offs that all organisms face. I first derive the two canonical equations of SDP. These are valuable because although they apply to no system in particular, they share commonalities with many systems (as do frictionless springs). After that, I show how we used SDP in insect behavioral ecology. I describe the puzzles that needed to be solved, the SDP equations we used to solve the puzzles, and the experiments that we used to test the predictions of the models. I then briefly describe two other applications of SDP in biology: first, understanding the developmental pathways followed by steelhead trout in California and second skipped spawning by Norwegian cod. In both cases, modeling and empirical work were closely connected. I close with lessons learned and advice for the young mathematical biologists.

[1]  David Rosen,et al.  Foraging and oviposition decisions in the parasitoid Aphytis lingnanensis : distinguishing the influences of egg load and experience , 1991 .

[2]  G. Engelhard,et al.  Scale analysis suggests frequent skipping of the second reproductive season in Atlantic herring , 2005, Biology Letters.

[3]  R. Nash,et al.  Evaluation of the frequency of skipped spawning in Norwegian spring-spawning herring , 2011 .

[4]  M. Mangel,et al.  Steelhead Life History on California's Central Coast: Insights from a State-Dependent Model , 2009 .

[5]  M. Mangel,et al.  Effects of the Emotion System on Adaptive Behavior , 2013, The American Naturalist.

[6]  D. Rosen,et al.  Influence of egg load and host size on host‐feeding behaviour of the parasitoid Aphytis lingnanensis , 1992 .

[7]  Ray Hilborn,et al.  The Ecological Detective , 2013 .

[8]  N. Yaragina Biological parameters of immature, ripening, and non-reproductive, mature northeast Arctic cod in 1984–2006 , 2010 .

[9]  R. Bellman Dynamic programming. , 1957, Science.

[10]  Marc Mangel,et al.  Seasonal dynamic shifts in patch exploitation by parasitic wasps. , 1992 .

[11]  E. Charnov,et al.  Evolution of Host Selection and Clutch Size in Parasitoid Wasps , 1984 .

[12]  R Bellman,et al.  On the Theory of Dynamic Programming. , 1952, Proceedings of the National Academy of Sciences of the United States of America.

[13]  R. Hilborn,et al.  The Ecological Detective: Confronting Models with Data , 1997 .

[14]  Richard Phillips Feynman,et al.  Mainly mechanics, radiation, and heat , 1963 .

[15]  D. E. Stokes Pasteur's Quadrant: Basic Science and Technological Innovation , 1997 .

[16]  Øyvind Fiksen,et al.  State-dependent Energy Allocation in Cod (Gadus Morhua) , 2006 .

[17]  Lawrence M. Dill,et al.  MORTALITY RISK VS. FOOD QUALITY TRADE-OFFS IN A COMMON CURRENCY: ANT PATCH PREFERENCES' , 1990 .

[18]  S. L. Lima,et al.  Behavioral decisions made under the risk of predation: a review and prospectus , 1990 .

[19]  M. Mangel,et al.  State-dependent life history models in a changing (and regulated) environment: steelhead in the California Central Valley , 2009, Evolutionary applications.

[20]  M. Mangel,et al.  Clutch size, offspring performance, and intergenerational fitness , 1994 .

[21]  M. Mangel The Theoretical Biologist's Toolbox: Quantitative Methods for Ecology and Evolutionary Biology , 2006 .

[22]  J. Krebs,et al.  Behavioural Ecology: An Evolutionary Approach , 1978 .

[23]  C. Miaud,et al.  To breed or not to breed: past reproductive status and environmental cues drive current breeding decisions in a long-lived amphibian , 2014, Oecologia.

[24]  S. Karlin,et al.  A second course in stochastic processes , 1981 .

[25]  Arthur J. Krenert A formal approach to stochastic integration and differential equations , 1980 .

[26]  M. Mangel Rate maximizing and state variable theories of diet selection , 1992 .

[27]  U. Dieckmann,et al.  The logic of skipped spawning in fish , 2006 .

[28]  R Bellman,et al.  DYNAMIC PROGRAMMING AND LAGRANGE MULTIPLIERS. , 1956, Proceedings of the National Academy of Sciences of the United States of America.

[29]  M. Mangel A treatment of complex ions in sea water , 1971 .

[30]  Marc Mangel,et al.  Modelling the proximate basis of salmonid life-history variation, with application to Atlantic salmon, Salmo salar L. , 2004, Evolutionary Ecology.

[31]  A. Houston,et al.  The Common Currency for Behavioral Decisions , 1986, The American Naturalist.

[32]  Steven F. Railsback,et al.  Agent-Based and Individual-Based Modeling: A Practical Introduction , 2011 .

[33]  R. Rideout,et al.  Skipped spawning in female iteroparous fishes , 2005 .

[34]  C. Clark,et al.  Dynamic State Variable Models in Ecology , 2000 .

[35]  Ben S Cooper,et al.  Confronting models with data. , 2007, The Journal of hospital infection.

[36]  R. Courant,et al.  Methods of Mathematical Physics , 1962 .

[37]  R. Nash,et al.  Frequent skipped spawning in the world’s largest cod population , 2012, Proceedings of the National Academy of Sciences.

[38]  John M McNamara,et al.  A classification of dynamic optimization problems in fluctuating environments , 2000 .

[39]  Zbigniew Michalewicz,et al.  An Evolutionary Approach , 2004 .

[40]  C. Clark,et al.  Towards a Unifield Foraging Theory , 1986 .

[41]  E. Charnov,et al.  Complementary Approaches to the Understanding of Parasitoid Oviposition Decisions , 1985 .

[42]  D. Ludwig,et al.  Definition and Evaluation of the Fitness of Behavioral and Developmental Programs , 1992 .

[43]  Marc Mangel,et al.  Dynamic models in behavioural and evolutionary ecology , 1988, Nature.

[44]  M. Mangel,et al.  Life expectancy and reproduction , 1993, Nature.

[45]  C. Clark,et al.  Dynamic State Variable Models in Ecology: Methods and Applications , 2001 .

[46]  M. Uschold,et al.  Methods and applications , 1953 .

[47]  M. Mangel Opposition site selection and clutch size in insects , 1987 .

[48]  C. Clark,et al.  Dynamic Modeling in Behavioral Ecology , 2019 .

[49]  M. Mangel,et al.  Contrasts in Habitat Characteristics and Life History Patterns of Oncorhynchus mykiss in California's Central Coast and Central Valley , 2012 .

[50]  Enric Sala,et al.  Natural History: the sense of wonder, creativity and progress in ecology* , 2001 .