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A  @n?" dd@  @@`` ph/t#      "#?B$]&M.# 3-WX )y(B$!]l-7x@6)y(B$'zE<:rlY y(c $ 333@ g4kdkd4Ipppp <4BdBdg}nzn<4!d!dg}nzng4?d?d4Ip@ pp? %O =j)#Why-questions in functional biology$#, $  Arno Wouters  Mayr  \Functional biology How-questions Proximate causes Decoding of genetic program Reductionistic6JJ ]  XEvolutionary biology Why-questions Ultimate causes Origin of genetic program Historical 4DC Y +# My claims  Functional biologists address why-questions in addition to how-questions Functional explanations have a nonreductionist character Functional explanations make sense of organisms without considering their history The source of biology s autonomy is complexity  #Why-questions in functional biology $ Introduction Different issues in functional biology Two types of function function as biological role function as biological advantage Functional explanation ConclusionsVJ=# &33=#  ,Tinbergen s four why s  CImmediate causation Function / survival value Development Evolution D GSix issues with regard to the form and activity of an item or behaviour H Description T!]&M.# 3-WX Zh{gt>Cx_lUƿ]ڥMJK`Ť5!PŌkO3; 1>`o*`b6 >"!V!4[% "&T) bCvSvrN7fX͖"G Uk@"[J36jiBVZfCh^3zl*y\ u2w=6T"vUaZW]M钪1C5fs*ϭUžgk٨qcs]Y~ _ CS 8OR!|/"E&^w`5ʷJ~P63GeItr7K6&оj0Dj'˴V &1YTU\&X}8uLh^EJeҊcl'clB8;)W F3?BFXf䑥ࠦGWz*G3 intlHUU7:piʮQ|6 v:w]47XTFV> 9;ԧ:e|$y f#<)SqZnނxx˲Fqsy"\k|Vf\TݎtL5vˉ2N>6ޅ n2\ߗ*5G5Y>w6Bwq,(z *p wcܯ0V\hP߬3vZtZnZs@eY[Dڋ SQ^/1vc^“ZH'rg7i$5¯сgzpNm7&\7@ 2Sp'붲ul}KC\1H(A !ab3~$??~^y=tS$!,Y֥}999ܷβ,ʜrp._SWuz8]s!9]vd3i;a'f6(X2q_ `8e&֘(uSSߪ>5xʂǓz6Vp_k*\ a\g7[IY"q\ܣW_l0!^VP1./FeWu3o'Gh̻']"(*[f˳*>`Qd6WZ_64QCުe,; ?s/^PAlFzɯa~^A (e'3yn9̆Q}:oPzh.,CXGUݮ|; 6~ Ȱ5pc{UX 'p Y1aCp#7(O]X_@L 4@qy`t{!*";*"te}.Ys>oوK(?u-m⥡^>Ȳ<}h]p]WWWW4A_F.Zُ0һ]zzwW^-yF:EVDEV\-:^>{79iew>6o^Rj Vv$kIvldrN[XߤR Ef?$2eCTb< sF֠R_d'Mfr6E,5(@"2tbQ7SbkATa+1kHp!jC0< \ZD/n=lK"_1s#K劮D-0 L[S-j qQL#Wc#2CL{9wo'1L?O73T? &h?OJQ3"zk&8%3}rɲH})?Qp9ͳw,cӹ4\vЫb(p' | _;T*fokiz>'k T'zE<:rlYo8hx v<VxOHQnm.A nlVb Mnn{iCQDPP<Q7u${7LF,}ćDhj)Se͟xVEZiX9j PiʡhdSL xb+j/8gFh4& sޓ 1qloډn\`~v;U5M.1Bф|V6|j4-钭G6``Cq:ZQk⪅V:K.\}6K W BG*e7Xnz q?쮷3B'add(|g=_ H3C*KT^Iޮen&R64-C М4'ׁݠdP8 JBqqoFNr#nq~78j XUfW\J rMgP >UR UtH-1ݸ,J-KY] pssu pه:BK^$%G8kV۲;|m+ur=:M$ǜ+Ng?ɹjS~pv^(5R#wu90＇1t$seT.@Sʍ$r )g8M1nN8{IDZ4i7&udp#XCzJu8/t2c y+;&S]8~'44od"ϕb^Rm>ӂamk R Biological role (function) Individual level causes and mechanisms Biological value (functional explanation) Development Evolution  Tinbergen versus Wouters  4 Immediate causation Function Development Evolution&5 5  XDescription Biological role Causes and mechanisms Biological value Development Evolution:Y  Y Important distinctions  Pfunction as biological role / function as biological advantage  what is the function? /  why does o have an i? specifying a function / giving a functional explanation6c ;  #Why-questions in functional biology $ Introduction Different issues in functional biology Two types of function function as biological role function as biological advantage Functional explanation ConclusionsJJ=#4=#  Function as biological role   The role of an item / activity is the manner in which that item / activity contributes to the activity of a complex system Nagel (1961), Cummins (1975), Boorse (1976), Craver (2001)   Examples biological role  The lungs have the function of gas exchange Some functions of the circulatory system are to transport oxygen, carbon dioxide, nutrients and heat. The heart propels the blood     Function as biological advantage !  How a trait effects that the fitness of its bearers is higher than the fitness of hypothetical organisms in which this trait is replaced by another one would be  )Examples function as biological advantage * "An advantage of the presence of a circulatory system is that oxygen is transported much faster by means of convection than by means of diffusion alone An advantage of the increased surface area of gills and lungs is that it increases the amount of oxygen intake in a certain amount of time # Advantages are comparative  To find an advantage of trait a relative to b an organisme (with a) is compared to a hypothetical organism in which a is replaced by b The organism with which the real organism is compared need not be real l 2H  $v is an advantage of a relative to b@%  % v is something that is different in organisms with a than in organism in which a is replaced by b that difference is the result of the replacement due to this difference, organisms with a have better life chances than organism in which a is replaced by b2Y1  !Role and value  The advantages of an item / activity having a certain character are often assessed in relation to the biological role of this item The role of an item provides a tacit fitness criterium The value is external to the role    Role vs. Advantage  WRole items / behaviours absolute position in an organization empirical generalisation XT X  -Advantage traits comparative value law-like. % .   Biological role is central  _Position / task of an item Mechanistic explanations Design explanations Selection explanations `   Mechanistic explanation How does a Vertebrate transport oxygen? Functional explanatin (design explanation) Why does a Vertebrate need to transport oxygen actively? Selection explanation How did the circulatory system evolve in the course of Vertebrate history?Why did it so?r)+:Z)+:Z  "#Why-questions in functional biology $ Introduction Different issues in functional biology Two types of function function as biological role function as biological advantage Functional explanation ConclusionsNJ=#I33=    Functional explanation  y Why does this organism have this item / performs this activity? Why does this item / activity has the character it has? z  Structure of design explanton  1) What is the biological role of this item / activity? 2) a) Why does the organism need to perform this role? b) Why is the role performed in the way in which it is performed rather than in some alternative way?  -$.Fick s law of diffusion    Nature of functional explanation !  A functional explanation shows how a trait of an organism functionally depend on other traits of that organism For example: living on land is functionally dependent on having lungs  $#Why-questions in functional biology $ Introduction Different issues in functional biology Two types of function function as biological role function as biological advantage Functional explanation ConclusionsNJ=#I33=   & Tinbergen versus Wouters  4 Immediate causation Function Development Evolution&5 5  XDescription Biological role Causes and mechanisms Biological value Development Evolution:Y  Y (!  *"   Conclusions  Functional biologists address why-questions in addition to how-questions Functional explanations have a nonreductionist character Functional explanations make sense of organisms without considering their history The source of biology s autonomy is complexity  /T#%sx(HHA@J {hhx El2 EGP'd5Ea   ` ̙33` ` ff3333f` 333MMM` f` f` 3>?" dd@(?" dd@  @ ` n?" dd@   @@``PR    @ ` ` p>>  7/(    6 P P V Click to edit Master title style! !   0d  P RClick to edit Master text styles Second level Third level Fourth level Fifth level!     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