What began as the study of animal social ) Tj 0 -11.25 TD 0.1513 Tc 0.0362 Tw (behavior, has now embraced the study of interactions at all ) Tj T* 0.0936 Tc 1.2189 Tw (biological levels. 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After ) Tj 0 -11.25 TD 0.0774 Tc 2.4539 Tw (group formation, members may reproduce or survive at ) Tj T* 0.1361 Tc 5.355 Tw (different rates leading to within group change. Multilevel selection framework for symbiotic species ) Tj -29.25 -12 TD -0.0127 Tc 0.0127 Tw (groups. BT 54 293.25 TD 0 0 0 rg /F1 9.75 Tf 0.193 Tc 0 Tw (Syn) Tj 15.75 0 TD 0.1205 Tc 0.8587 Tw (ergistic forms of cooperation benefit both the cell and ) Tj -15.75 -11.25 TD 0.1573 Tc 0.1052 Tw (the cell group. All that has changed ) Tj T* 0.2216 Tc 3.3409 Tw (with the study of ev) Tj 95.25 0 TD 0.1237 Tc 2.0888 Tw (olutionary transitions in levels of ) Tj -95.25 -11.25 TD 0.1489 Tc 1.9136 Tw (organization. Tj 78 0 TD /F2 9.75 Tf 0.1884 Tc 0.6241 Tw (Darwinian Dynamics, Evolutionary ) Tj -78 -11.25 TD 0.2322 Tc 3.3303 Tw (Transitions in Fitness and Individuality) Tj 177 0 TD /F1 9.75 Tf 0.0457 Tc 3.5168 Tw (. 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Princeton,) Tj 0 Tc -0.1875 Tw ( ) Tj -177 -11.25 TD 0.0049 Tc -0.1924 Tw (N.J.:Princeton University Pr) Tj 110.25 0 TD 0.537 Tc 0 Tw (ess.) Tj 24.75 0 TD 0 Tc -0.1875 Tw ( ) Tj -24.75 -11.25 TD ( ) Tj 9.75 0 TD 0.1056 Tc 3.6069 Tw (In our view an organism is more than a group of) Tj 0 Tc -0.1875 Tw ( ) Tj -9.75 -11.25 TD 0.15 Tc 0.9958 Tw (cooperating cells, more even then a group of cooperating ) Tj 0 -11.25 TD 0.1669 Tc 1.8956 Tw (cells related through common descent, sa) Tj 176.25 0 TD 0.0249 Tc 0.9126 Tw (y from a single ) Tj -176.25 -11.25 TD 0.115 Tc 1.9475 Tw (cell. The theory of its action was first fully explained by Charles Darwin and is now believed to be the main process that brings about evolution. History. Darwin's grand idea of evolution by natural selection is relatively simple but often misunderstood. The subscript ) Tj T* /F2 9 Tf -0.252 Tc 0 Tw (j) Tj 2.25 0 TD /F1 9 Tf 0.0011 Tc -0.0011 Tw ( refers to the number of non) Tj 100.5 0 TD 0.003 Tc 0 Tw (-) Tj 3 0 TD 0.0083 Tc -0.0083 Tw (mutant cells in the offspring group;) Tj 126.75 0 TD 0 Tc 0 Tw ( ) Tj -232.5 -12 TD /F2 9 Tf -0.252 Tc (j) Tj 2.25 0 TD /F1 9 Tf 0.0174 Tc 2.2326 Tw ( = 0, 1, 2, ...) Tj 54.75 0 TD /F2 9 Tf -0.003 Tc 0 Tw (N) Tj 6 0 TD /F1 9 Tf -0.0395 Tc 0.7895 Tw (, where ) Tj 31.5 0 TD /F2 9 Tf -0.003 Tc 0 Tw (N) Tj 6 0 TD /F1 9 Tf -0.0176 Tc 1.3509 Tw ( is the total number of cells in the ) Tj -100.5 -12 TD 0.0009 Tc 2.9991 Tw (offspring group, termed propagule size ) Tj 3.003 Tc 0 Tw (\() Tj 160.5 0 TD /F2 9 Tf -0.003 Tc (N) Tj 6 0 TD /F1 9 Tf 0.0851 Tc 2.3149 Tw ( = 1 for zygote ) Tj -166.5 -12 TD 0.0411 Tc 1.3089 Tw (reproduction\). When cooperation benefits the cell as well as ) Tj 0 -11.25 TD 0.1447 Tc -0.1655 Tw (the cell group, cooperation is always synergistic, as defined ) Tj T* 0.021 Tc -0.004 Tw (in terms of a payoff matrix. Multi-Level Selection and the Major Transitions in Evolution The body of ideas known loosely as ‘multi-level selection theory’ takes as its A ) Tj 120.75 0 TD /F2 9.75 Tf 0.1768 Tc 0 Tw (self) Tj 13.5 0 TD -0.2467 Tc (-) Tj 3 0 TD 0.2029 Tc 0.3596 Tw (policing modifier) Tj 71.25 0 TD /F1 9.75 Tf 0.2588 Tc 0.3037 Tw ( causes) Tj 30.75 0 TD 0 Tc -0.1875 Tw ( ) Tj -239.25 -11.25 TD 0.1392 Tc 1.8482 Tw (cells in the group to spend time and energy monitoring ) Tj 0 -11.25 TD 0.1892 Tc 3.8019 Tw (other cells and reducing the advantages of defection) Tj 0 Tc 0.5625 Tw ( ) Tj T* 0.2521 Tc 0.3104 Tw (possibly at some cost to the group. BT 54 729 TD 0 0 0 rg /F1 9.75 Tf 0.0493 Tc 0.4061 Tw (uniformly deleterious to selfish\). The neutral theory of molecular evolution contends that at the molecular level, most evolutionary changes and polymorphisms within species are not caused by natural selection, but by random genetic drift. Tj 165.75 0 TD 0 Tc -0.1875 Tw ( ) Tj -165.75 -11.25 TD ( ) Tj 9.75 0 TD 0.084 Tc 0.0098 Tw (The proliferation of cells during development is indicated ) Tj -9.75 -11.25 TD 0.1513 Tc 2.6612 Tw (by the vertical ar) Tj 75.75 0 TD 0.0768 Tc 1.2357 Tw (rows in ) Tj 36 0 TD -0.1179 Tc 0 Tw (Figure) Tj 25.5 0 TD /F1 9 Tf 0 Tc ( ) Tj 4.5 0 TD (1) Tj 4.5 0 TD /F1 9.75 Tf 0.099 Tc 1.5885 Tw (. Evolution of Human Mate Choice David C. Geary, Jacob Vigil, and Jennifer Byrd-Craven University of Missouri – Columbia This article provides a review of evolutionary theory and empirical research on mate choices in non-human species and used as a frame for understanding the how and why of … The question of levels of selection - on which biologists and philosophers have long disagreed - is central to evolutionary theory and to the philosophy of biology. Unlike the classical ) Tj T* 0.0929 Tc 0.6196 Tw (use of modifier models, say in the evolution of dominance ) Tj T* 0.126 Tc 2.2178 Tw (and recombination, the modifiers studied in the present ) Tj T* 0.2309 Tc 1.0816 Tw (paper are not neutr) Tj 80.25 0 TD 0.1631 Tc 0.7208 Tw (al. Michaela Lewis . Tj -204 -11.25 TD 0.099 Tc -0.2865 Tw (Roy. Get this from a library! Tj -80.25 -11.25 TD 0.1368 Tc 3.8423 Tw (The purpose of the modifier model is to study how) Tj 0 Tc -0.1875 Tw ( ) Tj 0 -11.25 TD 0.1206 Tc 2.4419 Tw (evolution may modify these assumptions and parame) Tj 228.75 0 TD 0.0713 Tc -0.2588 Tw (ter ) Tj -228.75 -11.25 TD 0.1327 Tc 3.2155 Tw (values, and further how these modifications affect the) Tj 0 Tc -0.1875 Tw ( ) Tj 0 -11.25 TD 0.082 Tc 0.2555 Tw (transition to a higher level of selection. In addition to studying the focal levels ) Tj -78.75 -11.25 TD 0.0677 Tc 3.776 Tw (of cells and cell groups \(the emerging organism\), we) Tj 0 Tc -0.1875 Tw ( ) Tj 0 -11.25 TD 0.2448 Tc 0 Tw (conside) Tj 31.5 0 TD 0.1504 Tc -0.0046 Tw (r the \223surrounding\224 levels of genes within cells and ) Tj -31.5 -11.25 TD 0.174 Tc 4.2323 Tw (populations of cell groups. Almost everybody feels that they understand DarwinÕs Theory of Evolution by natural selection. Cooperation is now seen as the primary ) Tj T* 0.1102 Tc 0.6666 Tw (creative force behind ever greater levels of co) Tj 188.25 0 TD -0.0682 Tc 0.6307 Tw (mplexity and ) Tj -188.25 -11.25 TD 0.0581 Tc -0.2456 Tw (organization in all of biology.) For example, under ) Tj T* 0.0959 Tc -0.0959 Tw (certain conditions some bacteria associate to form a fruit) Tj 226.5 0 TD 0.0965 Tc 0.466 Tw (ing ) Tj -226.5 -11.25 TD 0.1324 Tc -0.1699 Tw (body, some amoebae associate to form a slug, some solitary ) Tj 0 -11.25 TD 0.0536 Tc 1.1755 Tw (cells form a colonial group, some normally solitary wasps ) Tj T* 0.1316 Tc -0.0691 Tw (breed cooperatively, some birds associate to form a colony, ) Tj T* 0.0376 Tc 2.4416 Tw (and some mammals form societies. Tj 76.5 0 TD 0 Tc 0 Tw ( ) Tj ET endstream endobj 26 0 obj 42685 endobj 24 0 obj << /Type /Page /Parent 5 0 R /Resources << /Font << /F0 6 0 R /F1 8 0 R /F2 15 0 R /F3 17 0 R /F5 27 0 R >> /ProcSet 2 0 R >> /Contents 25 0 R >> endobj 30 0 obj << /Length 31 0 R >> stream T. Ohta, in Brenner's Encyclopedia of Genetics (Second Edition), 2013. A two ) Tj T* 0.5411 Tc 0 Tw (sp) Tj 9.75 0 TD 0.0823 Tc 3.1588 Tw (ecies group formed by association of members from) Tj 0 Tc -0.9375 Tw ( ) Tj -9.75 -11.25 TD 0.1754 Tc -0.1962 Tw (different species is indicated by the larger touching shapes; ) Tj 0 -11.25 TD 0.1481 Tc 2.7582 Tw (the size of each touching shape indicating the relative) Tj 0 Tc -0.1875 Tw ( ) Tj T* 0.2006 Tc 3.0807 Tw (proportions of each species within the group and the) Tj 0 Tc -0.1875 Tw ( ) Tj T* 0.2049 Tc 0.3576 Tw (shading of each shape indicating th) Tj 146.25 0 TD 0.0902 Tc 0.0973 Tw (e relative proportion of ) Tj -146.25 -11.25 TD 0.1959 Tc -0.1584 Tw (the two phenotypes within each species in the group. Effect of c) Tj 42 0 TD -0.053 Tc 0.053 Tw (ooperation on fitness at cell and organism level. ) It is crucial to distinguish the question concerning the level at which adaptations evolve from the question about the identity of the ultimate beneficiaries of that selection process. Also ignores cases where selection at both levels is in same direction. A two species group is indicated by touching ) Tj -44.25 -12 TD -0.0362 Tc 0.0362 Tw (shapes; the size of each touching shape indicating the relative ) Tj 0 -12 TD 0.0195 Tc -0.0195 Tw (proportions of each species within the group and the shading of ) Tj T* 0.0112 Tc -0.0112 Tw (each shape indicating the relative proportion of the two ) Tj T* 0.1007 Tc -0.1007 Tw (phenotypes within each) Tj 87 0 TD 0.0562 Tc -0.0562 Tw ( species in the group.) If costly, ) Tj T* 0.165 Tc 1.8975 Tw (cooperation is assumed to lower the ce) Tj 168.75 0 TD -0.0019 Tc 1.0644 Tw (ll\222s replication or ) Tj -168.75 -11.25 TD 0.1338 Tc 0.2037 Tw (survival rate. Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? The ) Tj T* 0.1093 Tc 1.096 Tw (evolution of these modifiers are the first ne) Tj 180.75 0 TD 0.1776 Tc 0.3849 Tw (w functions at ) Tj -180.75 -11.25 TD 0.1039 Tc 1.1405 Tw (the organism level. Tj 15.75 0 TD 0 Tc -0.1875 Tw ( ) Tj -139.5 -11.25 TD -0.2813 Tc 0.0938 Tw (3. ) The propagules then ) Tj T* -0.0273 Tc 1.2773 Tw (form offspring groups of the next generation \(dashed arrows\). ) To find out how it works, imagine a population of beetles: Delet) Tj 105 0 TD 0.1017 Tc -0.1392 Tw (erious mutation may occur during ) Tj -105 -11.25 TD 0.1578 Tc -0.0953 Tw (cell division. Selection at the species level is much harder to observe than selection at the cellular, or even the individual, level because species selection takes place over millions of years. 47) Tj 37.5 0 TD -0.2467 Tc 0 Tw (-) Tj 3 0 TD -0.0292 Tc 4.3417 Tw (92. Scholars generally ) Tj -24.75 -11.25 TD 0.0871 Tc 2.3921 Tw (viewed cooperation to be of limited interest, of special ) Tj 0 -11.25 TD 0.188 Tc -0.2391 Tw (relevance to certain groups of organisms to be sure, but not ) Tj T* 0.1249 Tc -0.1078 Tw (of general significance to life on earth. We are interested in) Tj 0 Tc 0.5625 Tw ( ) Tj T* 0.1598 Tc 2.6527 Tw (understanding these first true emergent level functions ) Tj T* 0.032 Tc 5.7805 Tw (during the origin of the multicellular organism. When humans manage this process, its called artificial selection. Tj 184.5 0 TD 0 Tc -0.1875 Tw ( ) Tj -119.25 -32.25 TD /F0 12 Tf -0.2052 Tc 0.2052 Tw (Heritability of F) Tj 79.5 0 TD 0.097 Tc 0 Tw (itness) Tj 29.25 0 TD 0 Tc ( ) Tj -174 -18 TD /F1 9.75 Tf 0.1612 Tc 2.3179 Tw (We wish to understand how heritability of fitness, the ) Tj 0 -11.25 TD 0.0956 Tc -0.1331 Tw (defining characteristic of a unit of selection, may increase at ) Tj T* 0.1712 Tc 4.308 Tw (the group level, so that the group may become an) Tj 0 Tc 0.5625 Tw ( ) Tj T* 0.1135 Tc 3.3419 Tw (evolutionary individual or unit of selection. Almost everybody feels that they understand DarwinÕs Theory of Evolution by natural selection. Tj 102 0 TD 0 Tc -0.1875 Tw ( ) Tj -102 -11.25 TD ( ) Tj 9.75 0 TD 0.1261 Tc 1.3739 Tw (To be able to adapt to their environment, evolutionary) Tj 229.5 0 TD 0 Tc -0.1875 Tw ( ) Tj -239.25 -11.25 TD 0.1297 Tc 0.0995 Tw (individuals must be units of selection, and satisfy Darwin\222s ) Tj 0 -11.25 TD 0.1128 Tc 1.033 Tw (conditions of heritability and variation in fitness. The propagules produced ) Tj T* 0.1558 Tc 1.1567 Tw (by the adult form offspring groups of th) Tj 169.5 0 TD 0.0985 Tc 0.464 Tw (e next generation ) Tj -169.5 -11.25 TD 0.1332 Tc 3.9293 Tw (\(dashed arrows\). H�b```f``�``e``/a`@ �����H#S�I,�,z,�~�,��4�N�L�,�(�4Q�����풸�Lq Tj 231 0 TD 0 Tc -0.1875 Tw ( ) Tj -231 -11.25 TD ( ) Tj 9.75 0 TD 0.0642 Tc 0.6483 Tw (The modifier allele can affect virtually any aspect of the ) Tj -9.75 -11.25 TD 0.0725 Tc 0.34 Tw (model, in particular we have studied modifiers that create a ) Tj 0 -11.25 TD 0.3498 Tc 0 Tw (ger) Tj 12.75 0 TD 0.0935 Tc 0.219 Tw (m line, that police other cells, or that cause cell death of ) Tj -12.75 -11.25 TD 0.1498 Tc 0.996 Tw (renegade dividing cells. A theory of evolutionary ) Tj 0 -11.25 TD 0.1612 Tc -0.3487 Tw (transitions. ) In the case of aggregation, the ) Tj T* -0.0311 Tc 0.4811 Tw (offspring group is formed by aggregation of cells. Using a ) Tj T* -0.0684 Tc 0 Tw (multi) Tj 19.5 0 TD -0.2467 Tc (-) Tj 3 0 TD -0.0169 Tc 7.3294 Tw (level selection framework) Tj 115.5 4.5 TD /F1 6 Tf 0 Tc 0 Tw (1) Tj 3 0 TD 0.252 Tc (-) Tj 2.25 0 TD 0 Tc (5) Tj 3 -4.5 TD /F1 9.75 Tf 0.1561 Tc 6.5939 Tw (, it is possible to) Tj 0 Tc 0.5625 Tw ( ) Tj -146.25 -11.25 TD 0.1072 Tc 0.3719 Tw (understand how Darwin\222s properties emerge at a new level ) Tj 0 -11.25 TD -0.1385 Tc 0.951 Tw (\(for example, the mult) Tj 87.75 0 TD 0.0091 Tc 0.5534 Tw (icellular organism\) from the evolution ) Tj -87.75 -11.25 TD 0.0771 Tc 2.2354 Tw (of interactions in groups of lower level individuals \(for ) Tj 0 -11.25 TD 0.0411 Tc -0.1604 Tw (example, cells\). 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