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Optimality model

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178:, a species of mollusk. To break the shell of the mollusk, the crows fly and drop the whelks on rocks. Reto Zach constructed an optimality model to predict the optimal height at which crows drop the whelks. The benefit in this model is the success rate of cracking the whelk's shell, while the primary cost is the energy spent flying. If the crows did not fly high enough, they would have little success in breaking the whelks' shells. However, the crows could waste valuable energy if they climb too high. In his model, Zach predicted the optimal height for crows to drop the whelks. To do this, Zach calculated the total distance each whelk was dropped before it was successfully broken. Whelks dropped from 3 meters and lower actually had traveled high total distances because they had to be dropped numerous times in order to be broken. On the other hand, whelks dropped from 5 meters and 15 meters were dropped approximately the same number of times to initiate a break; however, the crows would obviously have to climb higher to break a whelk at 15 meters than at 5 meters. Zach predicted 5 meters to be the optimal dropping height. The results indicated that the crows do follow this model, as the average dropping height was 5.2 meters. 220:. As starlings gather more leatherjackets, it becomes increasingly difficult and time-consuming to find subsequent leatherjackets with the additional prey in their mouths. Thus at some point, it benefits them to stop expending extra energy to find additional food and return to their nests instead. A graph of this phenomenon, called a loading curve, compares foraging time to the number of prey captured. Alex Kacelnik predicted that the curve would fluctuate depending on the starling's travel time. He predicted that starlings traveling further would spend more time at the foraging site to achieve optimal foraging behavior. It is important that these starlings spend extra time at the foraging ground because it takes a lot of energy to travel back and forth from its nest. On the other hand, he predicted starlings traveling shorter distances to foraging grounds should spend less time foraging, making more frequent trips in order to optimize their behavior. Since these starlings have a shorter distance to travel, they do not need to put as much energy into searching for leatherjackets because it is easier for them to return to the foraging ground. His results were consistent with his predictions. 195:
fertilize 80% of the eggs, while the first male will only fertilize 20%. This creates a dilemma for male dung flies. The longer they remain with a female after copulation, the better they are able to guard her from copulating with other males, hence increasing the likelihood of passing his genes to her offspring. However, the longer the male remains with an individual female, the smaller chance he has of finding other mates. Geoff Parker predicted that an optimality model comparing these two behaviors would be affected by the travel time between two patches. For example, short distances between cowpats should widen the pool of available mates in a specific geographic location. Parker predicted that under this condition, dung flies would be more likely to leave their current mate sooner to find additional mates. But if cowpats are few and far between, it would benefit a male dung fly to spend more time guarding a mate, ensuring his genes are passed on, because he may have difficulty finding an additional mate. The results from Parker's experiment agree with this model.
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their actions. Currency is defined as the variable that is intended to be maximized (ex. food per unit of energy expenditure). It is the driving factor behind an action and usually involves food or other items essential to an organism's survival. Constraints refer to the limitations placed on behavior, such as time and energy used to conduct that behavior, or possibly limitations inherent to their
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To construct an optimality model, the behavior must first be clearly defined. Then, descriptions of how the costs and benefits vary with the way the behavior is performed must be obtained. Examples of benefits and costs include direct fitness measures like offspring produced, change in lifespan, time
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Each time an organism displays a certain behavior, it must weigh the costs and benefits to make a decision. For example, given X amount of time traveling, after catching one bug, would it be better for a bird to continue foraging or to quickly return to its nest to feed chicks? Better understanding
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Optimality models are used to predict optimal behavior (ex. time spent foraging). To make predictions about optimal behavior, cost-benefit graphs are used to visualize the optimality model (see Fig 1). Optimality occurs at the point in which the difference between benefits and costs for obtaining a
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Optimal behavior is defined as an action that maximizes the difference between the costs and benefits of that decision. Three primary variables are used in optimality models of behavior: decisions, currency, and constraints. Decision involves evolutionary considerations of the costs and benefits of
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Dung flies are a polygamous species that mate on cowpats. The copulation behavior of this species can also be modeled using the marginal value theorem. It has been discovered that in cases where two male dung flies copulate with the same female in relatively rapid succession, the second male will
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To determine the optimum time spent on a behavior, one can make a graph showing how benefits and costs change with behavior. Optimality is defined as the point where the difference between benefits and costs for a behavior is maximized, which can be done by graphing the benefits and costs on the
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A currency must also be identified. A test of the predictions generated by the optimality model can be performed to determine which currency the organism maximizes at any given time. For example, when constructing an optimality model for bee foraging time, researchers looked at whether energetic
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Cells exhibit precise behaviors in response to physical cues. This optimality has been modeled by quantifying what information a sensor can learn from its physical surroundings. Over recent decades, experiments observed biophysical optimality in chemosensing, mechanosensing, and light sensing.
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Some authors have argued that optimality models may be insufficient in explaining an organism's behaviour. The degree of optimization in response to natural selection depends on the rate at which genetic structure changes, the amount of additive variance present at the time of selection,
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are a tool used to evaluate the costs and benefits of different organismal features, traits, and characteristics, including behavior, in the natural world. This evaluation allows researchers to make predictions about an organism's optimal behavior or other aspects of its
33:. It allows for the calculation and visualization of the costs and benefits that influence the outcome of a decision, and contributes to an understanding of adaptations. The approach based on optimality models in biology is sometimes called 215:
model. Researchers compared the amount of time a bird forages to the distance the bird travels to the foraging ground. Birds try to maximize the amount of food they take back to their offspring. Starlings mostly feed their offspring
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Optimal filtering in the salamander retina. F Rieke, WG Owen & W Bialek, in Advances in Neural Information Processing 3, R Lippman, J Moody & D Touretzky, eds, pp 377–383 (Morgan Kaufmann, San Mateo CA,
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Parker, Geoffrey A.; Simmons, Leigh W.; Stockley, Paula; McChristie, Doreen M.; Charnov, Eric L. (April 1999). "Optimal copula duration in yellow dung flies: effects of female size and egg content".
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efficiency (energy gained/energy spent) or net rate of gain ((energy gained − energy spent)/time) was optimized. It was found that the bees maximized energetic efficiency when foraging for nectar.
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and fluctuations in payoff affect optimality. Strict optima may not be reachable due to genetic and environmental changes. Genetic factors limiting the attainment of optimality include
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Frank, S. A. (1998). Foundations of social evolution. Princeton, NJ: Princeton University Press. 4 Prestwich, K. (2007). Notes on optimality theory. Retrieved from
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Davies, N., Krebs, J., & West, S. (2012). Introduction to Behavioural Ecology. (4 ed.). (pp. 432-433). Hoboken, NJ: John Wiley & Sons.
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Skedung, Lisa; Arvidsson, Martin; Chung, Jun Young; Stafford, Christopher M.; Berglund, Birgitta; Rutland, Mark W. (2013-09-12).
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Lucas, Jeffrey R. (August 1983). "The Role of Foraging Time Constraints and Variable Prey Encounter in Optimal Diet Choice".
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Beroz, Farzan; Jawerth, Louise M.; Münster, Stefan; Weitz, David A.; Broedersz, Chase P.; Wingreen, Ned S. (2017-07-18).
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Petkova, Mariela D.; Tkačik, Gašper; Bialek, William; Wieschaus, Eric F.; Gregor, Thomas (February 2019).
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Orzack, S. H., & Sober, E. (2001). Adaptationism and optimality. (1 ed.). Cambridge University Press.
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of the relationships between the values in a model leads to better predictions of organism behavior.
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Schmid-Hempel, Paul (February 1987). "Efficient Nectar-Collecting by Honeybees I. Economic Models".
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http://college.holycross.edu/faculty/kprestwi/behavior/e&be_notes/e&be_07_Optimality.pdf
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http://college.holycross.edu/faculty/kprestwi/behavior/e&be_notes/e&be_07_Optimality.pdf
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Parker, G. A.; Smith, J. Maynard (November 1990). "Optimality theory in evolutionary biology".
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Bautista, Luis M.; Tinbergen, Joost; Wiersma, Popko; Kacelnik, Alex (October 1998).
937: 644: 335: 967: 953:"Optimal Foraging and Beyond: How Starlings Cope with Changes in Food Availability" 917: 890: 828: 820: 779: 771: 730: 722: 681: 671: 627: 622: 600: 571: 563: 522: 514: 473: 465: 408: 373: 323: 300: 288: 113:) plotted versus a measure of the behavior. Optimality occurs where the difference 249:. Complementary strategies to describing and analyzing organism behaviour include 246: 824: 518: 567: 1043: 881:
Zach, Reto (1978). "Selection and Dropping of Whelks By Northwestern Crows".
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Beroz, Farzan; Zhou, Di; Mao, Xiaoming; Lubensky, David K. (2020-10-14).
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Aquino, Gerardo; Wingreen, Ned S.; Endres, Robert G. (March 2016).
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Bialek, W. (1987). "Physical limits to sensation and perception".
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Prestwich, K. (2007). Notes on optimality theory. Retrieved from
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can be predicted using an optimality model, specifically a
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Annual Review of Biophysics and Biophysical Chemistry
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y-axis and a measure of the behavior on the x-axis.
806: 549: 658:Endres, Robert G.; Wingreen, Ned S. (2008-10-14). 131: 105: 85: 353:(4 ed.). Hoboken, NJ: John Wiley & Sons. 49:currency via a particular behavior is maximized. 1041: 809:"Physical limits to sensing material properties" 174:On the Pacific coast of Canada, crows forage on 29:. Optimality modeling is the modeling aspect of 664:Proceedings of the National Academy of Sciences 348: 657: 599:Mora, Thierry; Wingreen, Ned S. (2010-06-18). 858:The Retina: An Approachable Part of the Brain 452:Berg, H. C.; Purcell, E. M. (November 1977). 363: 203:One common use of the optimality model is in 58:spent or gained, or energy spent and gained. 598: 451: 278: 432: 430: 832: 783: 734: 685: 675: 626: 616: 575: 526: 477: 349:Davies, N.; Krebs, J.; West, S. (2012). 274: 272: 270: 207:. For example, the foraging behavior in 185: 165: 68: 855: 427: 1042: 860:. Cambridge: Harvard University Press. 436: 398: 313: 267: 880: 439:Biophysics: Searching for Principles 413:10.1146/annurev.bb.16.060187.002323 351:Introduction to behavioural ecology 161: 152: 13: 14: 1061: 251:phylogenetic comparative methods 1024: 1012: 1003: 994: 944: 901: 874: 864: 849: 800: 751: 702: 651: 592: 52: 628:10.1103/PhysRevLett.104.248101 556:Journal of Statistical Physics 543: 494: 445: 392: 357: 342: 307: 223: 1: 470:10.1016/s0006-3495(77)85544-6 366:The Journal of Animal Ecology 260: 181: 198: 139:for a behavior is maximized. 7: 454:"Physics of chemoreception" 147: 10: 1066: 825:10.1038/s41467-020-18995-4 519:10.1016/j.cell.2019.01.007 568:10.1007/s10955-015-1412-9 960:The American Naturalist 895:10.1163/156853978x00297 856:Dowling, J. E. (1987). 677:10.1073/pnas.0804688105 605:Physical Review Letters 316:The American Naturalist 205:optimal foraging theory 922:10.1006/anbe.1998.1034 237:. Thus, discontinuous 213:marginal value theorem 191: 171: 140: 133: 107: 87: 813:Nature Communications 764:Nature Communications 441:. Princeton U. Press. 255:quantitative genetics 189: 169: 134: 108: 88: 72: 117: 97: 77: 776:10.1038/ncomms16096 670:(41): 15749–15754. 458:Biophysical Journal 437:Bialek, W. (2012). 132:{\displaystyle B-C} 31:optimization theory 715:Scientific Reports 513:(4): 844–855.e15. 192: 172: 141: 129: 103: 83: 727:10.1038/srep02617 106:{\displaystyle C} 86:{\displaystyle B} 35:optimality theory 22:optimality models 1057: 1034: 1028: 1022: 1016: 1010: 1007: 1001: 998: 992: 991: 957: 948: 942: 941: 910:Animal Behaviour 905: 899: 898: 889:(1–2): 134–147. 878: 872: 868: 862: 861: 853: 847: 846: 836: 804: 798: 797: 787: 755: 749: 748: 738: 706: 700: 699: 689: 679: 655: 649: 648: 630: 620: 596: 590: 589: 579: 562:(5): 1353–1364. 547: 541: 540: 530: 498: 492: 491: 481: 449: 443: 442: 434: 425: 424: 396: 390: 389: 361: 355: 354: 346: 340: 339: 311: 305: 304: 293:10.1038/348027a0 276: 162:Crows and whelks 153:Cellular sensing 138: 136: 135: 130: 112: 110: 109: 104: 92: 90: 89: 84: 1065: 1064: 1060: 1059: 1058: 1056: 1055: 1054: 1040: 1039: 1038: 1037: 1029: 1025: 1017: 1013: 1008: 1004: 999: 995: 955: 949: 945: 906: 902: 879: 875: 869: 865: 854: 850: 805: 801: 756: 752: 707: 703: 656: 652: 597: 593: 548: 544: 499: 495: 450: 446: 435: 428: 397: 393: 362: 358: 347: 343: 312: 308: 287:(6296): 27–33. 277: 268: 263: 247:genetic linkage 226: 201: 190:Two dung flies. 184: 164: 155: 150: 118: 115: 114: 98: 95: 94: 78: 75: 74: 55: 12: 11: 5: 1063: 1053: 1052: 1036: 1035: 1023: 1011: 1002: 993: 972:10.1086/286189 966:(4): 543–561. 943: 916:(4): 795–805. 900: 873: 863: 848: 799: 750: 701: 650: 611:(24): 248101. 591: 542: 493: 464:(2): 193–219. 444: 426: 391: 372:(1): 209–218. 356: 341: 328:10.1086/284130 322:(2): 191–209. 306: 265: 264: 262: 259: 225: 222: 218:leatherjackets 200: 197: 183: 180: 163: 160: 154: 151: 149: 146: 128: 125: 122: 102: 82: 54: 51: 9: 6: 4: 3: 2: 1062: 1051: 1048: 1047: 1045: 1033: 1027: 1021: 1015: 1006: 997: 989: 985: 981: 977: 973: 969: 965: 961: 954: 947: 939: 935: 931: 927: 923: 919: 915: 911: 904: 896: 892: 888: 884: 877: 867: 859: 852: 844: 840: 835: 830: 826: 822: 818: 814: 810: 803: 795: 791: 786: 781: 777: 773: 769: 765: 761: 754: 746: 742: 737: 732: 728: 724: 720: 716: 712: 705: 697: 693: 688: 683: 678: 673: 669: 665: 661: 654: 646: 642: 638: 634: 629: 624: 619: 614: 610: 606: 602: 595: 587: 583: 578: 573: 569: 565: 561: 557: 553: 546: 538: 534: 529: 524: 520: 516: 512: 508: 504: 497: 489: 485: 480: 475: 471: 467: 463: 459: 455: 448: 440: 433: 431: 422: 418: 414: 410: 406: 402: 395: 387: 383: 379: 375: 371: 367: 360: 352: 345: 337: 333: 329: 325: 321: 317: 310: 302: 298: 294: 290: 286: 282: 275: 273: 271: 266: 258: 256: 252: 248: 244: 240: 236: 235:genetic drift 232: 221: 219: 214: 210: 206: 196: 188: 179: 177: 168: 159: 145: 126: 123: 120: 100: 93:) and costs ( 80: 71: 67: 63: 59: 50: 46: 44: 38: 36: 32: 28: 23: 19: 1026: 1014: 1005: 996: 963: 959: 946: 913: 909: 903: 886: 882: 876: 866: 857: 851: 816: 812: 802: 770:(1): 16096. 767: 763: 753: 718: 714: 704: 667: 663: 653: 608: 604: 594: 559: 555: 545: 510: 506: 496: 461: 457: 447: 438: 404: 400: 394: 378:10.2307/4810 369: 365: 359: 350: 344: 319: 315: 309: 284: 280: 227: 202: 193: 173: 156: 142: 64: 60: 56: 53:Construction 47: 39: 34: 21: 15: 819:(1): 5170. 721:(1): 2617. 407:: 455–478. 224:Limitations 45:abilities. 261:References 239:phenotypes 182:Dung flies 73:Benefits ( 883:Behaviour 618:1002.3907 243:mutations 231:gene flow 209:starlings 199:Starlings 124:− 27:phenotype 1050:Ethology 1044:Category 988:12049476 980:18811363 938:23642915 930:10202088 843:33056989 794:28719577 745:24030568 696:18843108 645:10526866 637:20867338 586:26941467 537:30712870 336:85216873 148:Examples 834:7560877 785:5520107 736:3771396 687:2572938 577:4761375 528:6526179 479:1473391 421:3297091 301:4348920 170:A crow. 43:sensory 18:biology 986:  978:  936:  928:  871:1991). 841:  831:  792:  782:  743:  733:  694:  684:  643:  635:  584:  574:  535:  525:  488:911982 486:  476:  419:  384:  334:  299:  281:Nature 176:whelks 984:S2CID 956:(PDF) 934:S2CID 641:S2CID 613:arXiv 382:JSTOR 332:S2CID 297:S2CID 976:PMID 926:PMID 839:PMID 790:PMID 741:PMID 692:PMID 633:PMID 582:PMID 533:PMID 507:Cell 484:PMID 417:PMID 386:4810 253:and 245:and 968:doi 964:152 918:doi 891:doi 829:PMC 821:doi 780:PMC 772:doi 731:PMC 723:doi 682:PMC 672:doi 668:105 623:doi 609:104 572:PMC 564:doi 560:162 523:PMC 515:doi 511:176 474:PMC 466:doi 409:doi 374:doi 324:doi 320:122 289:doi 285:348 16:In 1046:: 982:. 974:. 962:. 958:. 932:. 924:. 914:57 912:. 887:67 885:. 837:. 827:. 817:11 815:. 811:. 788:. 778:. 766:. 762:. 739:. 729:. 717:. 713:. 690:. 680:. 666:. 662:. 639:. 631:. 621:. 607:. 603:. 580:. 570:. 558:. 554:. 531:. 521:. 509:. 505:. 482:. 472:. 462:20 460:. 456:. 429:^ 415:. 405:16 403:. 380:. 370:56 368:. 330:. 318:. 295:. 283:. 269:^ 257:. 37:. 20:, 990:. 970:: 940:. 920:: 897:. 893:: 845:. 823:: 796:. 774:: 768:8 747:. 725:: 719:3 698:. 674:: 647:. 625:: 615:: 588:. 566:: 539:. 517:: 490:. 468:: 423:. 411:: 388:. 376:: 338:. 326:: 303:. 291:: 127:C 121:B 101:C 81:B

Index

biology
phenotype
optimization theory
sensory


whelks

optimal foraging theory
starlings
marginal value theorem
leatherjackets
gene flow
genetic drift
phenotypes
mutations
genetic linkage
phylogenetic comparative methods
quantitative genetics



doi
10.1038/348027a0
S2CID
4348920
doi
10.1086/284130
S2CID
85216873

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