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Multilevel Selection and Population Genetics in Structured Multilevel Selection, Population Genetics and Cooperation in Structured Populations NIMBioS Tutorial: Game Theoretical Modeling of Evolution in Structured Populations Jeremy Van Cleve University of Kentucky 26 April 2016 Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY How do cooperative behaviors evolve? Cooperation occurs when: focal: cost to improve state of its partner partner: beneft from improved state Food improves state ⟶ higher ftness Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY trate into a tightWhen form they known cease as moving,and the a when cells Mexican there ofenvironment hat. is the lacks Then, either slug electrolytes, concen- directional when light it orimmune is system, no before they very are light. shed at moist, thefunctioning rear of simultaneously the slug. asthe liver, slug from kidney, front and toered back innate class picking up of toxins cells and calledand bacteria, ultimately sentinel become cells the stalk. that There is sweep athe through recently discov- constituent cells. Those atthe slug the lacks a front nervous system, direct there are movement differencesmove: among an important advantage to themore quickly social and farther stage. than Though any individual amoebaeffectively could recovering the solitaryrear, stage. and these The cells slug can feed moves sheath on any largely bacteria made they discover, upfrom of it cellulose, it in drops some cells important at ways. the Astoward light and heat it and away from ammonia crawls ( throughwhich a then elongatesAfter slightly a few and hours, beginsa this center to center concentrates into crawl in aconcentrate around a mound, in great process streams of calledhighest dicty cells, aggregation concentration. flowing ( toward As more and more starve, they Figure 3 sexual stage of basal disc and the stalk(b) are Motile formed multicellular of slug formerly moving living towards amoebae light. that (c) have Fruiting died body to consisting form of this a supporting basal structure. disc, (d) a Macrocysts, stalk, the and a sorus, or spores. The Slugs move farther and for a longer time when the This translucent slug looks like a tiny worm, but differs Multicellular stages of D. discoideum. (Courtesy of Owen Gilbert). Dictyostelium discoideum Encyclopedia of Animal Behavior Animal of Encyclopedia Dictyostelium discoideum A Figure 3(a) u Figure 3(b) t h o r ' . (a) Aggregation of formerly independent cells into a multicellular body. Author's personal copy s ). ). p e Dictyostelium, the Social Amoeba 515 r s stalk ( to rise upthe out front of of thein the mass a slug as begin process a to called very form culmination, slender cellulose the but walls cells rigid and that were at price of postage.lected This stock clones center obtainedMany is from studies accessed through can the be stockand performed Cultured center using for previouslyHow the Dictyostelids col- Are Obtained, Collected, fruiting body, then they alsobacteria, perish. or enough other shed cellsduring to their form normal a new, movement. smaller Ifslug. these do Still not encounter otherstoxins were and shed bacteria asOthers from they sacrifice the made themselves rear theirthe as way of through sentinel soil the the cells surface,others slug picking may or rise up up into and sporulate a a millimeter gap or sostructure between above called soil a particles. the fruiting body spores, ( and stalk, at and thethree-quarters top basal or they disk form so comprise hardy of spores. an the At erect cells this point, flow up this stalk, o (2010), vol. 1, pp. 513-519 Thus, some of the cells sacrifice their lives so that the n a Figure 3(c) l c Dictyostelium o p y ). These cells die. The remaining , the Social Amoeba Figure 3(c) ). 515 Cost: dying as part of the stalk Beneft: surviving as part of the spore Figure 3 Multicellular stages of Dictyostelium discoideum. (a) Aggregation of formerly independent cells into a multicellular body. (b) Motile multicellular slug moving towards light. (c) Fruiting body consisting of a basal disc, a stalk, and a sorus, or spores. The basal disc and the stalk are formed of formerly living amoebae that have died to form this supporting structure. (d) Macrocysts, the sexual stage of D. discoideum. (Courtesy of Owen Gilbert). NIVERSITY OF ENTUCKY Multilevel selection, pop gen and games in structured pops U K highest concentration. As more and more starve, they in a process called culmination, the cells that were at concentrate in great streams of dicty cells, flowing toward the front of the slug begin to form cellulose walls and a center in a process called aggregation (Figure 3(a)). to rise up out of the mass as a very slender but rigid After a few hours, this center concentrates into a mound, stalk (Figure 3(c)). These cells die. The remaining which then elongates slightly and begins to crawl around three-quarters or so of the cells flow up this stalk, toward light and heat and away from ammonia (Figure 3(b)). and at the top they form hardy spores. At this point, This translucent slug looks like a tiny worm, but differs the spores, stalk, and basal disk comprise an erect from it in some important ways. As it crawls through a structure called a fruiting body (Figure 3(c)). sheath largely made up of cellulose, it drops cells at the Thus, some of the cells sacrifice their lives so that the rear, and these cells can feed on any bacteria they discover, others may rise up and sporulate a millimeter or so above effectively recovering the solitary stage. The slug moves the soil surface, or into a gap between soil particles. more quickly and farther than any individual amoeba could Others sacrifice themselves as sentinel cells picking up move: an important advantage to the social stage. Though toxins and bacteria as they made their way through the the slug lacks a nervous system, there are differences among slug. Still others were shed from the rear of the slug the constituent cells. Those at the front direct movement during their normal movement. If these do not encounter and ultimately become the stalk. There is a recently discov- bacteria, or enough other shed cells to form a new, smaller ered class of cells called sentinel cells that sweep through fruiting body, then they also perish. the slug from front to back picking up toxins and bacteria, functioning simultaneously as liver, kidney, and innate immune system, before they are shed at the rear of the slug. How Dictyostelids Are Obtained, Collected, Slugs move farther and for a longer time when the and Cultured environment lacks electrolytes, when it is very moist, and when there is either directional light or no light. Many studies can be performed using previously col- When they cease moving, the cells of the slug concen- lected clones obtained from the stock center for the trate into a tight form known as a Mexican hat. Then, price of postage. This stock center is accessed through Encyclopedia of Animal Behavior (2010), vol. 1, pp. 513-519 gene chromosomes prokaryotes + mitochondria/ eukaryotes chloroplasts clonal sexual reproduction reproduction independent multicellular living cells organisms independent social groups individuals (eusociality) “Major transitions in evolution” or “transitions in individuality” (Maynard Smith and Szathmáry) Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY Cooperation occurs at two “scales” “Within a group” “Between/among groups” Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY Cooperation occurs at two “scales” 1. Plastic behaviors “Within a group” “Between/among groups” 2. Kin & Group selection Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY 1. Plastic behaviors Game theory (Nash equilibrium / ESS) Tit for tat / punishment / reputation / etc Reciprocity or responsiveness 2. Kin & Group selection Multilevel selection combines responsiveness with kin/group processes through measures of population structure such as relatedness Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY Outline The two scales of cooperation Simple model of responsiveness within a population Evolution in structured populations: the Price equation Multilevel model with the Price equation Evolution in structured populations: fxation probability & trait substitution Social games in an island-model using fxation probability Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY Outline The two scales of cooperation Simple model of responsiveness within a population Evolution in structured populations: the Price equation Multilevel model with the Price equation Evolution in structured populations: fxation probability & trait substitution Social games in an island-model using fxation probability Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY Plastic behavior and repeated games Plastic behavior requires repeated interactions (i.e., a repeated game) Cooperate (C) Defect (D) Payoff to Strategies in the repeated game Beneft – Cost – Cost determine how individuals respond Cooperate (C) to the actions of social partners Payoff to Beneft 0 Defect (D) Multilevel selection, pop gen and games in structured pops UNIVERSITY OF KENTUCKY Plastic behavior and repeated games Strategy Description ALLD Always play D “always defect” GRIM Play C but switch to D once opponent defects “grim trigger” TFT Start with C and then repeat the opponent's last move “tit for
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