Matthew C. Keller Sarah E. Medland Laramie E. Duncan

Transcription

1 Bhv Gn OI / ORIGINL RRCH r n in ily ign Worh h roubl? Coprion o h Bi, Prciion, n ccurcy o Prr i in our in ily Mol Mh C. Kllr rh. Mln Lri. uncn Rciv: 23 Jun 2009 / ccp: 24 Novbr 2009 Ó pringr cinc+buin Mi, LLC 2009 brc h clicl in ign (C) u obrv covrinc ro onozygoic n izygoic in pir o inr h rliv gniu o gnic n nvironnl cu o phnoypic vriion. pi i i u, i i ll knon h h C cn prouc bi i i i ringn upion r no. By oling obrv covrinc o in rliv in iion o in hlv, n in ily ign () ruir l ringn upion, cn i ny or prr o inr, n houl prouc l bi i hn h C. Hovr, r or coplic o u n inrpr, n by ping o i lrg nubr o prr, h prciion o prr i y ur. hi ppr i orl invigion ino ipl uion: I i orhhil o u or copl ol uch in bhviorl gnic? In priculr, copr h bi, prciion, n ccurcy o i ro h C n hr incringly copl. W in h C i by Gi Lubk. M. C. Kllr L.. uncn prn o Pychology n Nurocinc, Univriy o Coloro, Boulr, CO, U M. C. Kllr L.. uncn Iniu or Bhviorl Gnic, Univriy o Coloro, Boulr, CO, U.. Mln Qunln Iniu or Micl Rrch, Bribn, QL, urli M. C. Kllr (&) prn o Pychology n Nurocinc, Munzingr Hll, 345 UCB, Boulr, CO 80309, U -il: h.c.kllr@gil.co URL:.hckllr.co. o cn job o iing bro n hribiliy, bu C i o hr nvironnl c n h rliv ipornc o iiv vru non-iiv gnic vrinc cn b bi, oi illy o. Incringly copl, on h ohr hn, r or ccur n l niiv o upion hn iplr ol. W conclu h rrchr inr in chrcrizing h nvironn or h kup o gnic vriion houl u hn poibl. Kyor Bhvior gnic Mol ipciicion n in ily ign Clicl in ign Prr inrincy Inroucion h obrv covrinc o in, op, n hir ily br r on u o unrn h rliv ipornc o gnic n nvironnl cu o phnoypic vriion. h o coonly u gniclly inoriv ign i h Clicl in ign (C), hich copr h onozygoic (MZ) in covrinc o h izygoic (Z) in covrinc o i h vriion in ri u o uniu nvironnl c (V ) ll ny o o h hr vrinc coponn iiv gnic (V ), oinnc gnic (V ), n coon nvironnl (V C ) h cu ilil iilriy. hr r vrl ppl o h C. or pl, MZ n Z in rv nurl conrol o on nohr, hir i rlivly ipl o collc, n hr nvironnl c r no conoun ih gnic c, hy r in non-in ilil ui (Mrin l. 1997). Nvrhl, i h long bn unroo h h C ur ro vrl iporn liiion (v l.

2 Bhv Gn 1978). or on, ^V, ^V, n ^V C r uully conoun in h C, lloing only o o h hr prr o b i. 1 hi ollo ro h c h i i ipoibl o iulnouly i hr prr ( ^V, ^V, n ^V C ) ro ju o pic o rlvn inorion (h MZ n Z covrinc). o circuvn hi unr-iniicion probl, bhviorl gnici uing h C rouinly u h ihr V = 0 or h V C = 0. Hovr, h r iply upion, unbl uing in lon, born ro h hicl nciy o king h C inii. o h gr h upion r viol, C i o V n o b bi upr n i o V n V C n o b bi onr (Gryon 1989; Hh l. 1985; Kllr n Covnry 2005). con, h C o no ol h c o oriv ing or gn-nvironn covrinc, h prnc o hich ill cr bi in i (.g., ^V ill b oo lo). hir, h C h nohing o y bou h iology o h hr nvironnl c (conribuing o V C ): o h gr r hy p culurlly ro prn o opring n o h gr r hy u o non-prnl cor uch pr inlunc? inlly, h C o no u inorion icinly: or vry in pir rcrui (o n ubjc), only ingl iionl bi o inorion (on covrinc i) i gin uul o oling h cu o ilil iilriy. or h n ohr ron, in h 1970 rrchr bgn ploring n in ily ign (), hich ruir l ringn upion n prouc l bi i hn h C (ulkr 1982). h lrniv ign u on prn o in (v l. 1978; Nl n ulkr 1984) n opring o in (Nnc n Cory 1976) o br rvl gnic noniiviy n h rol o prnl nvironnl c, n u prn o in n pou o in (v 1979) o ol h c o oriv ing. Cloningr l. (1979) ir crib ho o u ll o h rliv yp oghr in ingl ol. hir ol i h orrunnr o h hr crib in hi ppr: h Nuclr in ily ign (N) (Hh l. 1985), h lh ign (ru l. 1994), n h Cc ign (Kllr l. 2009). or or horough hiory o in n ily ign, v (2009). n in ily ign () r h liiion o h C crib bov. Copr o h C, llo or inr grin cripion o h cu o phnoypic vriion, hy prouc l bi prr i, n or inorion (incring iicl por) i gin pr iionl ubjc in (Pohu n Boo 2000). Y, h rucion in bi n or 1 W ollo h convnion h ^V i h i o h populion prr V il inorion oci ih co h co o grly incr copliy. hi copliy i jor probl or inniing h ol ino co. or pl, uch crip rin in M (Nl 1999) cn rch or 50 prin pg or or, king hun rror virul criny rgrl o ho vigiln h rror chcking i. W no, hovr, h n vrion o M, OpnM (hp://opn.pyc.virgini.u/), ill b vilbl pckg or h R iicl lngug in rly 2010, n chng in h OpnM yn houl igniicnly ipliy co. Nvrhl, h copliy o y lo obcur logicl rror h hr o h ign; crin pcion y iply hv bn rong h oling g. urhror, ih ll ol, lo u k upion in orr or hir ol o b inii, n i i poibl h hy y pror b or or hn iplr ol hn h upion r viol. inlly, h copliy o ol n h nubr o prr hy p o i y l o n unccpbl lvl o iprciion in i cu by h high covriion bn h lrg nubr o i prr (ulicolinriy probl). or h ron, o rrchr in bhviorl gnic rin kpicl o h vlu o n vor h u o iplr, i- ol uch h C, hich r y o u n inrpr n ruir l collcion. h gol o hi ppr i o plor h r-o. In priculr, u iulion o gug h bi, prciion, n ccurcy o prr i uing h C n hr in orr o unrn hhr hy ork inn, unr h circunc hir i r bi, i h incr in inorion in co n unccpbl co in prciion, n ho violion o upion c prr i. In iion o iniying h cnrl nncy o h prr i, lo plor hir pr, covriion, n iribuionl hp. uch rul cn hlp rrchr inrpr C n ining ih propr circupcion. In ury, hi ppr i orl invigion ino ipl uion: I i orhhil o u or copl ol uch in bhviorl gnic? Mho Gnrl rgy W k our propri h bi, prciion, ccurcy, n iribuion o prr i riv ro h C, N, lh, n Cc ign. O cour, h prr bi, prciion, ccurcy, n iribuion or givn ign chng pning on h cnrio, o n o ur h propri unr vrl cnrio h

3 Bhv Gn igh occur in nur. givn cnrio, or pl, igh iul pcii lvl o iiv gnic, oinn gnic, n coon nvironnl c on o hypohicl ri. h cnrio houl lo viol upion o h our ign o chck hir niivii o upion. o ccoplih h gol, h ir uhor cr progr, Gnvolv, h iul in ily. h ur uppli inpu or vriou prr (.g., h oun o vriion in phnoyp u o vriou yp o gnic n nvironnl c) o iul irn cnrio. W obin iul in ily ro Gnvolv unr vrl irn cnrio h igh occur in rl li n rn M ol ro h our ign bov on hi. W hn copr h i vrinc prr (no by ^V ) riv ro M, o h ru vrinc prr (no by V ) iul uing Gnvolv. W ir hi proc 500 i or ch o 10 irn cnrio. In ol, 20,000 M ol r i (500 irion 9 4 ol pr irion 9 10 cnrio), king ol o *13,000 h o CPU i. cripion o h hr n in ily ign () bl 1 giv h inrprion o h vrinc prr icu in hi ppr ll hich ign cn i hich vrinc prr. or cripion o h C, Ploin l. (2001), n or or il cripion n plnion o h hr, incluing lgbric pcion, Kllr l. (2009). Nuclr in ily ign (N) h N (ig. 1) u on MZ in, Z in, n hir prn. h hr rliv cl provi our pic o inorion ro hich prr r i: h covrinc bn MZ in, C ^VðMZ; MZÞ, h covrinc bn Z in, C ^VðZ; ZÞ, h covrinc bn prn, C ^VðpouÞ, n h covrinc bn prn n chilrn, C ^VðPr; ChilÞ. hi iionl inorion llo h N o i ^V, ^V, n ^V C iulnouly, llo h c o oriv ing on prr i o b ccoun or, n llo piv gn-nvironn covrinc o b irni ro h c o ^V C. Whil hr r ny y h N cn b prriz, ocu hr on prrizion (ig. 1) h ivi ^V C ino h vrinc o c hr bn ibling n in bu no prn ( ^V ) n h vrinc o c h r rni vi vricl rniion ro prn o opring ( ^V ). Bcu only hr pic o, C ^VðMZ; MZÞ, C ^VðZ; ZÞ, n C ^VðPr; ChilÞ, provi inorion on our prr ( ^V, ^V, ^V, n ^V ), on o h prr (ypiclly ^V or ^V ) u b o 0 in ny N ol. Ln vrinc h r no hon in ig. 1, 2, n 3 r ul o 1. lh ign By uing ro MZ n Z in n hir ibling, prn, opring, n pou, 88 -pciic rliv covrinc cn b i. Mny o h 88 rliv cl r inicl cp or -pciic phy. or pl, nph-un covrinc bn on o Z l n hir l Z co-in r irni ro nph-un covrinc h r bn on o Z l n l Z co-in. h lh u h 88 covrinc obrvion o iulnouly i pciic ^V, ^V, ^V, ^V, ^V, n ^V ( bl 1 or hir inrprion) ll iiv gnic vriion uniu o l/l, h c o oriv ing, n covrinc. h lh ol u in hi ppr i iplii by cluing c, rucing h nubr o rliv cl ro 88 o 17. h ph igr or hi lh ol i hon in ig. 2, n i inicl o ig. 1 cp h pou o in n chilrn o in hv bn. o kp h igr unclur, ibling o in r no hon. Cc ign Lik h lh, h Cc u inorion on in n hir ibling, prn, pou, n chilrn o ol ll o h vrinc coponn ol by h lh. Hovr, liiion o h lh i h i ol only on yp o ing (priry phnoypic ing) n only on yp o vricl rniion (ro prnl phnoyp o opring ). h purpo o h Cc i o provi gnrl rork or rling h upion rgring choic n vricl rniion by h lh. hi i on hrough h u o ln phnoyp upon hich pou or upon hich prn inlunc hir chilrn. o kp h nubr o ol coprion ngbl, ocu hr on h ing pc o h Cc rhr hn h vricl rniion pc o i. h only irnc bn ig. 2 (h lh ol) n ig. 3 (h Cc ol) i h iion o h ln phnoyp ( P) upon hich or. pning on h yp o ing or vricl rniion ol bing u, h ph coicin o P r o ihr b ul o h ph coicin o P or o b ul o zro. or pl, o ol ocil hoogy, ll gnic ph coicin o P r o zro ( ¼ 0 n ¼ 0) n ll nvironnl ph coicin o P r conrin o b ul o h vlu o h corrponing ph coicin o P ( ¼, ¼, ¼, n ¼ ). o unrn hhr ocil

4 Bhv Gn bl 1 plnion o vrinc prr in h 10 irn iul cnrio Prr ibl in Inrprion V P, r 2 C, N, lh, Cc Phnoypic vrinc V C*, N**, lh, Cc iiv gnic vrinc; vrinc o rginl or vrg lllic c V C*, N**, lh, Cc oinnc gnic vrinc; vrinc o c ribubl o cobinion o lll h locu V C C*, N V C = V? V ; ypiclly i in C ol n o N ol. In h lh, Cc, n o N ol, i i pli ino V n V V N**, lh, Cc ibling nvironnl vrinc; vrinc in non-gnic c (.g., pr, cohor, chool, prning yl, c.) hr bn ibling n in bu no bn prn n opring V N**, lh, Cc ilil nvironnl vrinc; vrinc in non-gnic c (.g.,, ocil or, ucion) p (vi vricl rniion ) ro prn o opring V N, lh, Cc in nvironnl vrinc; vrinc in non-gnic c (.g., pr, cohor, clroo, in uro c) hr by in bu no ibling V C, N, lh, Cc Uniu or riul nvironnl vrinc; vrinc in non-gnic c (.g., pr, uniu princ, oic uion, urn rror) h r unhr ih ny ohr rliv cl V 9 Non iiv-by-iiv piic gnic vriion; vrinc ribubl o cobinion o lll iiv c irn loci V 9g Non Non-clr iiv gnic-by-g inrcion vrinc; hi cu rucion in iiv gnic covrinc bn rliv uncion o hir g irnc CV(, ) N, lh, Cc Covrinc bn iiv gnic n ilil nvironnl c; ri i vricl rniion (cuing V ) i uncion o h prnl phnoyp bcu,.g., highr vlu on cr highr phnoypic vlu, hich r p o opring vi vricl rniion * h C cn i ny o o V, V,orV C. ** h N cn i ny hr o V, V, V or V hoogy or priry phnoypic ing b i h, h i o h ocil hoogy ol cn b copr o ol o priry phnoypic orn, in hich ¼, ¼, ¼, ¼, ¼, n ¼. iuling in ily Gnvolv (Kllr 2007) i n opn ourc progr rin in h R progring lngug (R vlopn ig. 1 N ph igr P 1 1 /.25 µ P Mo Ph Covrinc Ph i o 1/2 P - Phnoypic vrinc - iiv gnic - oinn gnic - ily nvironn - ibling nvironn - Uniu (non-hr) nvironn - Covrinc bn - - ilil rniion µ - oriv ing coph P 1 P 2

5 P Mo P P 1 P p P 2 P p P Ch P Ch Ph Covrinc Coph Ph i o 1/2 P - Phnoypic vrinc - iiv gnic - oinn gnic - ilil nvironn - ibling nvironn - in nvironn - Uniu (non-hr) nvironn - Covrinc bn - - ilil rniion µ - oriv ing coph µ µ µ 1 /.25 1 / 0 1 ig. 2 lh ph igr P Mo P Mo µ P P P 1 P 1 P p P p P 2 P 2 P p P p P Ch µ µ P Ch Ph Covrinc Coph Ph i o 1/2 P - Phnoypic vrinc (ln) P - Phnoypic vrinc - iiv gnic - oinn gnic - ilil nvironn - ibling nvironn - in nvironn - Uniu (non-hr) nvironn - Covrinc bn - - ilil rniion z - Phy i o z or o 0 µ - oriv ing coph 1 /.25 1 / 0 1 ig. 3 Cc ph igr Bhv Gn

6 Bhv Gn Cor 2009) n vilbl.hckllr. co. Gnvolv ccurly iul gniclly inoriv ll copl ynic in voluionry gnic. Wih coplic cnrio, i i iicul or ipoibl o in pc uilibriu prr vlu nlyiclly (.g., h uilibriu iiv-by-iiv piic gnic vriion in populion ing orivly). oing o hrough iulion, hovr, i righorr. Givn ur inpu, Gnvolv iul h c o lll n nvironn on iniviul ri in populion, n llo hi populion o volv (,, n hv opring, ho,, n hv opring, c. ) or ny gnrion, unil prr rch uilibriu. Currnly, Gnvolv llo ur inpu o 48 irn prr, incluing 21 vrinc n covrinc prr, 3 irn yp o oriv ing, n 3 irn yp o vricl rniion. Gnvolv h n opion o cr in n in rliv phnoyp uring h inl gnrion o h iulion. W u hi opion o ri ou h phnoypic cor o in n hir ibling, pou, prn, n opring o l il (on ro pr ily), hich r hn u inpu ino M ( blo). ch l il conin ol o *15,000 ili (6,500 MZ ili n 8,500 Z ili). lhough hr r ol o 18 ponil rliv yp in ch ily (o in, o prn, our ibling, on pou o in 1, on pou o in 2, our chilrn o in 1, n our chilrn o in 2), ili h n vrg o bou iv non-iing phnoypic cor n ch l il conin ol *70,000 iniviul. h nubr r chon o rlc h pl iz n iingn prn in h cobin urli n Virgini n in b ( Mln n Kllr 2009), hich i h lrg n in ily in inc. Miingn in n in ri hrough iiculi in crinn ll vriion in g o h n nubr o chilrn ihin ili. pl iz o hi gniu r ncry or king in-grin iincion bn prr, pcilly ih rpc o -pciic phy (Hh l. 1985; Mln n Kllr 2009), lhough or o r u or irniing ol h o no ruir irni phy. bl 2 ho ho ch o h n cnrio in in hi projc in. V o.3 or ch cnrio, n ll ohr vrinc prr no hon in bl 2 r o zro. h vrinc coponn inhri by opring V, V, V 9, n V 9g n o incr cro gnrion uncion o oriv ing n/or vricl rniion, n rch uilibriu vlu ihin 5 10 gnrion. W rn ch Gnvolv iulion or 20 gnrion o nur h h prr rch uilibriu. I cn b iicul o pric h uilibriu vlu o h vrinc coponn h bginning o iulion. Our rgy hror o bgin ch Gnvolv iulion uch h ll vrinc coponn u o uniy (V p = 1) h ir gnrion, n o llo h vrinc coponn n V p o incr o hvr hir uilibriu vlu r. h uilibriu vlu or ch vrinc coponn (ro h 20h gnrion) r hon in bl 2; vlu in prnh r h r vlu i irn hn h uilibriu vlu. hu, h uilibriu vrinc coponn i no u o uniy or iv o h ol. W iul hr irn o o oriv ing ( ro 5 8, bl 2). Phnoypic hoogy (lo cll priry phnoypic orn ) occur hn lik ih lik b on h ni phnoyp. or pl, i ll popl choo ohr ll popl bcu hy r ll, hi oul cliy phnoypic hoogy. hi i h o coonly ol yp o oriv ing in h bhviorl gnic n voluionry gnic lirur. bl 2 iul vrinc prr oci ih 10 irn cnrio cnrio V V V V V V 9 V 9g r (pou).m. yp N/ N/ N/ (.15) N/? oriv ing.49 (.40) (.15) Phnoypic hoogy? oriv ing.42 (.40) (.15) ocil hoogy? oriv ing.51 (.40) (.15) Gnic hoogy? oriv ing.35 (.30) (.10) Phnoypic hoogy? 9 pii N/? 9 g In N/ Nubr in prnh r vrinc prr h ir gnrion, hich y chng by h inl (hr 20h) gnrion i vricl rniion or oriv ing occur ( ) V.30 or ll cnrio;.m. oriv ing

7 Bhv Gn ocil hoogy rr o iilriy riing ro iilr nvironnl bckgroun. or pl, i popl rry ihin rligion n choic o rligion i no hribl, hn ny iilriy bn pou u o rligion (.g., iilr vi on borion) oul b u o ocil hoogy rhr hn priry phnoypic orn. hir poibiliy, gnic hoogy, occur i choo ch ohr b on h hribl pc o hir phnoyp rhr hn on hir ni phnoyp (ihr 1918; hin n Grgg 1980). lhough ingly ipluibl, hr r o y hi igh occur. h ir i i popl p o conrol or h c o h nvironn hn king choic (.g., H/h i rlly r givn h nvironn hy co ro ). h con i i popl b choic on o hir vribl (.g., ovrll vlu) h i rl o h phnoyp o inr purly gniclly. hi oul b n r or o goo gn hori o hun choic (Millr n o 1998). Conir, or pl, oriv ing or inllignc. I popl choo olly b on vlu (.g., h ir principl coponn o ri uch hlh, hlici, high, cil rcivn, boily rcivn, inllignc, n o orh), n i h inr-rlionhip bn h vlu coponn i gnic in nur, hn iilriy bn pou on inllignc oul b u o gnic hoogy. Our poin i no o rgu h gnic hoogy i or i no likly o o iilriy, bu rhr o no h i i vibl opion h houl b piriclly. O h our in-ily ign icu hr, only h Cc cn ol gnic n ocil hoogy. W lo iul o cnrio h inclu prr h coul no b i in ny ol (ro 9 10, bl 2). h o cnrio llo u o h niiviy o upion or ll ign, incluing h lh n Cc. Mol iing ih M h uhor ro M crip or h C (137 lin o co), h N (189 lin o co), n h Cc ign (2,717 lin o co); h crip or h lh ign (2,780 lin o co) rin by H. M (M l. 2009). h crip r vilbl hp://.hckllr.co/hl/ cc.hl. n vng o h lh crip, no y inni in h Cc crip, i h i i up o i ulivri. h vng o h Cc ign, n i originl purpo, i h iionl libiliy in oling oriv ing n vricl rniion. or ch iul run uing h N, lh, n Cc crip, boh ull n ruc ol r i (no ruc ol r ncry or h C). h ull N ol i ^V, ^V, ^V, ^V, n ihr ^V (i ilil c i in h cnrio) or ^V (i ibling c i). 2 h ull lh n Cc ol i ^V, ^V, ^V, ^V, ^V, n, ^V, (no h C ^Vð; Þ i chniclly non-linr conrin n i no rly i; Kllr l. 2009). h ruc N, lh, n Cc ol i only ho vrinc prr h r ruly non-zro in h givn cnrio. h iing o boh ull n ruc ol on o onr h c o h coon prcic o ropping non-igniicn pricor. or pl, unr h cnrio (op ro, bl 2), ^V, ^V, ^V, ^V, ^V, n ^V r i in h ull lh n Cc ol bu only ^V, ^V, n ^V r i in h ruc ol; ^V, ^V, n ^V r ropp ( ul o 0). Our rgy hror u h no yp-i rror occurr in chooing h ruc ol. Whil no opil, cring progr h h igniicnc o ch i iniviully n ropp non-igniicn on oul hv norou copliy n copuing i ono projc h lry rch boh o h lii. Morovr, i oul hv bn incorrcly rin only *5% o h i (h yp-i rror r), n hror hi rgy inrouc only inor n probbly ngligibl inccurcy ino our ruc ol rul. ining h bi, prciion, n ccurcy o prr i W copr h prr i ro M or ch ign o h ru prr ro Gnvolv or ch iulion run. hi llo u o piriclly rin h bi, prciion, n ccurcy o h prr i, ll hir iribuionl hp n covrinc (Cl n Brgr 1990). h bi o iic i gnrlly in ð ^V V Þ, h pc (i.., n) irnc bn h i prr, ^V, n h ru prr, V. n lrniv i o u h in irnc rhr hn h pc irnc, Mð ^V V Þ, hich i l inlunc by oulir i. W cho hi lr ur o bi bcu vrl oulir ^V in our r probbly ricul u o h uo y h ol r run. lhough icr i ro ol h gv Co R (IIL = 6) in M, hich occur hn conrin cnno b ii n i ypoic o poorly proring iion, inpcion o M oupu l u o conclu h occionlly (*2 8% o h i, pning on h cnrio), M poorly rcr h pc covrinc ri n gv b i vn hn no Co R occurr. uch i r 2 ricly pking, C ^Vð; Þ i nonlinr conrin n i no rly i in. I i rin by, n hlp o rin, i prr by conrining hir inr-rlionhip in y h kp h nir ol inrnlly conin.

8 Bhv Gn ricul in h prn con bcu hy likly coul hv bn vr in o rl li oling con by proviing irn r vlu, ropping prr, or by king ohr ril ur o iprov h i. h prciion o i ur h pr o h i roun hir cnr, n i ypiclly ur by h nr viion or vrinc o h prr i,.g., n 2 iiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiii P 1 ^V i ð ^V Þ. n lrniv hich n 1 i¼1 u or h ron nion bov nly, h ih o onigh oulir h r likly o b ricul i h in bolu viion, or M, hich i ul o M ^V i Mð ^V Þ. h ccurcy o iic cobin inorion on boh bi n prciion o gug ho r y ro h ru vlu n i ypiclly i. hu, n i cn b prci bu nvrhl inccur i i i bi, or cn b unbi bu inccur i i i iprci. ih prciion, ccurcy i on ur uing h vrinc or nr viion, cp h i r jug by ho r y hy r ro h vlu o h ru prr rhr hn h vlu o h n i,.g., iiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiii P 1 n 2 ^V i ðv Þ. In hi iuion, ccurcy 2 = n 1 i¼1 bi 2? prciion 2 uing h ir o ch o h iniion bov. In h prn uy, u h in bolu rror, M ^V i V, o ur ccurcy o o ln h ipc o oulir. Rul Bi, prciion, n ccurcy o prr i h iribuion o our o h prr i or ch o h n cnrio crib in bl 2 r hon in ig. 4, 5, 6, 7, 8, 9, 10, 11, 12, n 13. h igur o no ho ^V, ^V,orC ^Vð; Þ bcu h i n o b o l inr. h igur lo plc C i o ^V C ino h colun rrv or ^V or ^V, hichvr i ppropri givn h cnrio. no bov, no ruc C ol n o b i, n o ruc C i r no hon. Rul or h n cnrio, hich i no viol upion in ny o h our ign, r hon in ig. 4 n 5. hing houl b no. ir, hn upion o h C r no viol (i.., V C in h cnrio n V in h cnrio), i ro h C r unbi n hv cn prciion. con, h ruc ol ro h hr r lo unbi, n hy hv grr prciion hn h C i. Ruc i r or prci bcu hy r b on uch or inorion (covrinc obrvion) hn h C i. hir, h ull ol or h hr ho vrying gr o bi n poorr prciion hn h ohr ol. h bi in h ull ol occur or h unnl ron h bi i in Cholky ol (Cry 2005): vrinc i r orc o b non-ngiv. By chnc, h ull ol pick up ig. 4 cnrio

9 Bhv Gn ig. 5 cnrio ig. 6 cnrio ligh vinc or non-zro vrinc prr h, in ruh, r cully zro (.g., V n V in h cnrio). I h vinc ugg h h i r poiiv, ol i h rly, bu i ngiv, h i hi h zro bounry. hi iblnc pull h ohr i prr (.g., ^V n ^V in h cnrio) in only on ircion, cuing bi. hi ourc o bi, hough inor, coul b rov i h ol llo vrinc i o b ngiv. h lck o prciion in ull, on h ohr hn, cnno b o ily rcii, bu rhr i nurl conunc o ping o i o ny or prr in, pcilly in h lh n Cc ign. igur 6, 7, n 8 ho rul or hr cnrio in hich C upion r viol bcu boh hr nvironnl n non-iiv gnic c inlunc ri iulnouly n, in h inl cnrio, bcu

10 Bhv Gn ig. 7 cnrio ig. 8 & priry phnoypic ing (r =.3) cnrio oriv ing i. Hovr, h cnrio o no viol upion or ny. h C i r highly bi in h pc ircion (Gryon 1989; Kllr n Covnry 2005), ih iiv gnic c bing ovri by bou 50% in h pl n non-iiv gnic c ignor bcu, or ron o iniibiliy, hy coul no b i. hr nvironnl c r unri by h C in h cnrio, bu r ovri in h n & Priry oriv Ming cnrio. hi ovriion i lo pricbl, n occur bcu o h ubnil CV(, ) h i inuc by vricl rniion, hich iic hr nvironn in h C (v l. 1989). pc, h ruc ol o no ho bi hr h ull ol ho ligh bi or h ron icu bov. h lh n Cc i r ui ccur in h cnrio, ypiclly bing ihin.05 poin o h ru prr. N

11 Bhv Gn ig. 9 & priry phnoypic ing (r =.3) cnrio ig. 10 & ocil hoogy (r =.3) cnrio i r l ccur hn boh ^V n ^V r i iulnouly; hi i u o h vry high corrlion bn h o i ( blo). igur 9 ho rul or coplic cnrio in hich V, V, V, V, V, V, n CV(, ) ll conribu o phnoypic vrinc in h con o priry phnoypic oriv ing. Hr, h N upion h ihr V,orV i zro i viol, cuing i h, lhough prci, r ui bi. Bcu ll prr r rin in h ruc ol, h rul or h ull n ruc ol r inicl. ll lh n Cc i r unbi; hovr, ^V ho irly high gr o iprciion u o h corrlion bn ^V n ^V, n bn ^V n ^V ( n cion). igur 10 n 11 ho rul or cnrio inicl o h pic in ig. 8 cp h poul iilriy i u o ocil hoogy (ig. 10) or gnic hoogy (ig. 11). hu, h o cnrio viol upion

12 Bhv Gn ig. 11 & gnic hoogy (r =.3) cnrio ig. 12 & 9 pii (vr =.15) cnrio or vry ign cp or h Cc, n pc, ll ign ohr hn h Cc prouc i h r bi o vrying gr. In priculr, i poul iilriy i u o ocil hoogy rhr hn priry phnoypic orn, h lh ovri V n V n unri V. On h ohr hn, i poul iilriy i u o gnic hoogy rhr hn priry phnoypic orn, h lh ovri V n V unri V. l in h con o h pciic prr vlu iul in h o cnrio, oling oriv ing incorrcly uing h lh i or i ocil hoogy i h ru cu o poul iilriy hn i gnic hoogy i h ru cu o poul iilriy. igur 12 n 13 ho rul or cnrio in hich upion r viol in vry ign. Whn gnic non-iiviy i u o iiv-by-iiv pii rhr hn oinnc (ig. 12), ol n o

13 Bhv Gn ig. 13 & 9 g Inrcion (vr =.15) cnrio ovri V n lighly unri V. Hovr, h ovrll lvl o gnic vriion (V? V? V 9 ) n o b only lighly unri. Morovr, i ^V i conir bro i o non-iiv gnic vrinc rhr hn n i o oinnc vrinc only, i o non-iiv gnic vriion r only lighly unri. Non-clr gn-by-g inrcion (ig. 13) cnb concpuliz irn gn urning on irn g, n oppo o clr gn-by-g inrcion, n o cr gnic covriion bn rliv uncion o h g irnc bn h. Bcu ibling n in n o b clo in g o on nohr, i i nibl h non-clr gn-by-g inrcion l o ovriion o V (no hon) n V n unriion o V in. nohr inring riicion o uch non-clr gn-by-g inrcion i h hy cn l o ngiv vricl rniion phy in (cring poiiv ^V bu cring iilriy bn prn n opring), no uncoon obrvion in piricl ui. In h C, on h ohr hn, non-clr gn-by-g inrcion cu ovriion o V. lhough r r o no ol h hv bn rin o o o, houl b bl o ol non-clr gn-by-g inrcion u o h i vriion in g ihin ili u in. or pl, in M, g irnc bn ch pir o ily br coul b clcul ro iniionl vribl, n h g irnc coul b u o or h pc iiv gnic covrinc bn rliv yp. Rlionhip bn prr i h inorion ruir o i prr i on prilly runn. or pl, boh V n V cu ihin-ily iilriy h rop o uncion o ho in rliv pir i, n o ^V n ^V n o b ngivly rl: on i incr n plin givn prn o obrv covrinc, hr i l inorion l ovr or h ohr i o plin. igur 14 ho h ^V n ^V, n ^V n ^V u prilly runn inorion in h Cc ign n o r highly ngivly rl. ^V n ^V r poiivly rl, bu only in ol h lo i ^V : ^V incr, boh ^V n ^V cr. ^V, on h ohr hn, i nrly inpnn o ^V, ^V n ^V in h Cc. Inorion o i ^V co pririly ro h coprion bn in n ibling covrinc vru prn opring covrinc, n hu o no u inorion h ovrlp ih ny o h ohr i. linr rgrion ol pricing ^V in h Cc ro ^V n ^V unr h cnrio pic in ig. 9 h n r 2 =.969, hich rnl o vrinc 1 inlion cor o 1 r ¼ 32:6. hu, h vrinc o ^V 2 in h Cc ol i 32.6 i highr, n h nr rror o ^V i 5.7 i highr, hn in ol in hich boh ^V n ^V r ropp. iilrly, h nr rror o ^V n ^V r 4.4 n 4.8 i highr, rpcivly, hn hy r in ol in hich hy r i lon. iilr ining occur or h ohr o ol. hi c cn b n in ig. 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, n13, hich ho h h iribuion o prr

14 Bhv Gn ig. 14 Prr corrlion ro Cc ol iing prr unr n & priry phnoypic ing (r =.3) cnrio bl 3 ll ub o h ourc o inorion or iing prr in h Cc n lh ol Prr Incr uncion o ^V C ^VðMZÞ C ^VðZÞ; C ^VðMZ:vunculrÞ C ^VðZ:vunculrÞ; C ^VðMZ:couÞ C ^VðZ:couÞ ^V C ^VðMZÞ C ^VðZÞ; C ^VðZÞ C ^VðPrn; OpringÞ ^V C ^VðZÞ C ^VðMZÞ; C ^VðZÞ C ^VðPrn; OpringÞ; C ^VðZÞ C ^VðMZ:vunculrÞ ^V C ^VðPrn; OpringÞ CVðMZ:vunculrÞ; C ^VðZ:couÞ C ^VðMZ:couÞ ^V C ^VðZÞ C ^VðibÞ i or ull ol r coninly or pr ou hn h iribuion o prr i or ruc ol. Inorion or iing prr in I i uul o hv n o ho obrv covrinc i rnl ino i prr. In h C, i i obviou h h irnc bn C ^VðMZ; MZÞ n C ^VðZ; ZÞ provi ll h inorion n o i ^V n ^V C (in C ol) or ^V n ^V (in ol). Hovr, i bco incringly iicul o icrn ho obrv covrinc i inlunc i prr in incringly copl. or pl, hich covrinc i hlp irni ^V ro ^V in h lh or Cc? Wh inorion llo irniion o ocil hoogy ro priry phnoypic orn in h Cc ol? Unorunly, hr r no ipl nr o h yp o uion in. hug nubr o prilly runn bi o inorion hlp i h unknon prr in, n h c o hi inorion pn on h ol bing i (.g., ho oriv ing i ol) ll on h vlu o h ohr iulnouly i prr (.g., h gr o vricl rniion lr ho obrv covrinc c ^V ). pi h iiculi, bl 3 provi o inigh ino ho obrv covrinc r u o i prr in h Cc n lh ol. h bl i no huiv; or crin prr (pcilly ^V n ^V ), nrly vry covrinc i ply o rol in hir iion. Rhr, bl 3 li o o h o conin ourc o inorion cro ol u in iing h iv vrinc prr h cu ilil rblnc. C ^VðMZ:vunculrÞ rr o h covrinc

15 Bhv Gn bn h chilrn o on MZ in n h ohr (vunculr) MZ co-in, hr C ^VðMZ:couÞ rr o h covrinc bn couin ho prn r MZ co-in. Wih rpc o oriv ing in h Cc ol, in-l r priculrly hlpul or irniing ocil ro phnoypic hoogy. or pl, unr ocil hoogy, hr i no pc irnc bn MZ in-l corrlion n Z in-l corrlion, hr unr phnoypic hoogy, in-l rlionhip houl ir by zygoiy u. icuion Our rul ho h ork ign. hy r gnrlly unbi hn upion r, n unlik h C, hy r no ovrly niiv o violion o upion o long ^V i inrpr broly, n i o gnic non-iiviy in gnrl (incluing gn-by-g inrcion c) rhr hn oinnc in priculr. Our rul lo highligh h h ky ro in uing i on o copliy vru ccurcy. By ping o i lrg nubr o prr, ny o hich u ovrlpping inorion, h prciion o i ur ( h ull ol i in ig. 4, 5, 6, 7, 8, 9, 10, 11, 12, n 13 n prr covrinc in ig. 14). h i in ig. 8, or pl, r uch l prci hn ho ro h C. Nvrhl, i n o b unbi unr uch ir rng o cnrio hn C i, n bcu o hi, r lo univrlly or ccur hn r C i. hi iprov ccurcy cn b unii by piricl rrchr uing by copring goon o i in o n only iing prr (.g., ^V, ^V, n ^V ) vru n iing ll prr. h irnc bn h o i inic provi n i o ho iporn uing n i ovr iplr ol (.g., h C) givn h phnoyp in uion. h rn o incring ccurcy ih incring copliy rp il ihin h ol: lh i r ccur cro ir rng o cnrio hn r N i (ig. 6), n Cc i r ccur cro ir rng o cnrio hn r lh i (ig. 10, 11). or pl, h n ccurcy vlu (lor bing or ccur) cro h n cnrio or ^V r.140 or h C,.069 or h N,.049 or h lh, n.045 or h Cc. pc, h Cc n lh rul r virully inicl cp in c hr upion rgring ing in h lh r viol. Nvrhl, h uion rin: givn h incr iiculy in iing h ol n collcing h ruii, i i orh i o u? Our rul cnno provi n nr o hi uion, bu hy o provi guinc. or ll h probl oci ih h C, h cobin C prr o ^V þ ^V o provi cn i o bro n hribiliy. I rrchr gol i pririly o unrn bro n hribiliy, or o unrn bro n gnic covrinc in ulivri ing, h C i u, n uing i probbly no orh h hl unl n ily lry i. o h gr h ny gnic non-iiviy or poul iilriy i, hovr, C ol cn illy unr- or ovr-i hr nvironnl c ( ig. 7, 8, 9, 10, n 11). hu, i on inr i in chrcrizing h c o h nvironn in ny y incluing rguing h hr nvironnl c r ll h C i ingulrly b ho. iilrly, i on inr i in unrning h rliv ipornc o iiv vru non-iiv gnic vriion, h C provi lil hlp. In h lr iuion, rrchr houl riouly conir h u o. h concluion r no rly b on h iulion rul o hi ppr. Covnry n Kllr (2005) copr h prr i o vry vilbl lh ol run up o h i o h i h oul hv bn obin uing h C on h n phnoyp. Conin ih pricion, hy oun h C rul gv pricbly ior picur o h kup o gnic vriion n h kup n ipornc o h coon nvironn. or rrchr ho lry hv h n o i h lh or Cc ol, our rul ugg h Cc ol houl b u ovr iing ol. Hovr, n rgun coul b ro our rul h h N rprn goo coproi bn h ccurcy o h Cc n h ipliciy o h C. N i n o b l prci n lighly or bi hn Cc i, bu h irnc r inor copr o h irnc bn h i group n h C i. O cour, h jor liiion o h N i h h ourc o hr nvironnl c (u o ibling c or vricl rniion ro prn) cnno b icrn, n hn boh hr nvironnl ourc r prn, i prr ill b bi. In pr pic (Mln n Kllr 2009), icu hich rliv yp provi h o por or cing irn prr in h Cc, hich houl b o rvic o invigor inr in collcing n or ny ( lo Hh l. 1985). Hill l. (2008) rcnly rgu h o gnic vrinc in o ri i iiv in nur. I V 0 or o ri, hn C i o V n V C houl b ccur in h bnc o oriv ing n vricl rniion, n hu oul on b ovrkill.

16 Bhv Gn Whil gr ih Hill l. (2008) concluion h gnic vriion i likly o b oly iiv in nur or o ri, igr ih ponil concluion rn ro hi ppr (.g., Whlbrg 2009) h noniiv gnic vrinc i ypiclly ll n inigniicn. -nlyi o rul ro h lh ign (Covnry n Kllr 2005) oun h ypiclly ^V [[ 0 n, on vrg cro 38 phnoyp, ^V nrly lrg ^V, bing ull 75% o ^V. h lh rul hoing vinc or ubnil non-iiv gnic vrinc r uch or convincing hn Hill l. (2008) in-only nlyi, in hich corrlion o onozygoic n izygoic in r copr cro 86 phnoyp: hv hon (ig. 4, 5, 6, 7, 8, 9, 10, 11, 12, n 13), h rliv gniu o V vru V cnno b ccurly crin uing in lon. Morovr, bcu nurl lcion ro iiv gnic vriion uch r hn non-iiv gnic vrinc, hory ugg h ri rl o rinin in houl hv rlivly high gr o non-iiv gnic vriion (Hln 1932; Wrigh 1929), n in piricl rvi ho h non-iiv gnic vrinc in nonhun nil i iilr in gniu o iiv gnic vrinc ong in-rl ri (Crnokrk n Ro 1995). hu, ihou piricl invigion, hink i oul b prur o k olc in h hop h noniiv gnic vrinc i lo nough or o ri or C i o b gnrlly unbi. hr r vrl liiion ih h currn pproch o unrning h bi, prciion, n ccurcy o prr i ro in-ily ign. ir, nion bov, our procur or uoing ol iing n h h rul ro ruc ol r opiiic. Hovr, rgu in h Mho cion, hi probbly prouc ngligibl gr o bi in our rul. or iporn ourc o bi in our rul, hich ork in h oppoi ircion, i h hun coul no gui ch iing proc inrcivly u o h uo y ol r i. non-ngligibl nubr (2% o 8% pning on h cnrio) o ol run prouc oulir i, poorly rprouc h obrv covrinc ric, n probbly il o in h ru iu liklihoo i. n princ olr coul hv c h iuion n kn ril ur, uch chnging r vlu, o iprov h i o h ol. hi ugg h h rul prn in hi ppr ppr l prci hn hy ill b hn i inrcivly on rl. nohr liiion o h currn pproch h invig only vry ll porion o h pc o poibl prr h igh i in h rl orl. or pl, i no invig lrniv o o vricl rniion or poul iilriy u o convrgnc, boh o hich cn b ol in h Cc. W lo i no invig ny nubr o lrniv cnrio h igh occur n cu bi in ll h ol invig hr, uch i ol o oriv ing (Rynol l. 2000), iionl yp o gn nvironn inrcion n corrlion, highr-orr pii, in uro c, n pcil MZ-in nvironn. hi lr iu i priculrly iporn. h hr o ll in ol, incluing, i h coprion bn MZ n Z in. I o non-gnic cor uch in uro c incr MZ in rblnc, ll ol crib in hi ppr ill ovri ^V n pcilly ^V. urhror, or ipliciy, i no invig pciic i in hi ppr, hich oul hv h iilr bi bu lor prciion hn ho prn hr. Givn hi, non bu h lrg -pciic c r likly o b cbl ih vn h lrg vilbl n in ily. inl liiion o our uy i h only univri ol r invig. lhough univri prr i r inring in hir on righ, ol bco or inring in ulivri con. or pl, prnl rh y b ngivly oci ih olcn prion in chilrn (Oprrio l. 2006), bu h ron or hi ociion r unclr. ol cn icrn hhr hi ociion i u o h gn cing boh rh n prion rik or o prnl rh bing culurlly rni o opring in h or o lor prion rik. Our ppr i no h prr chrcriic in uch ulivri ol, lhough hr i no ron o bliv h h uliy o ulivri prr i oul b ubnilly irn hn univri on. pi h liiion, h currn ppr rprn h ull plorion o o ho irn rl orl cnrio c i ro in-ily ign. W hv rgu h h o coonly u ign in bhviorl gnic, h C, i inu or unrning h rliv gniu o hr nvironnl c or h rio o iiv o non-iiv gnic vriion. Our rul onr h, irrpciv o por or pl iz, i o h o unii ro C cnno b inrpr ih ny gr o coninc unl rong upion no oriv ing, no gn-nvironn covrinc, n h ihr noniiv gnic vrinc or hr nvironnl vrinc i zro hv bn vrii., on h ohr hn, provi unbi n irly ccur i o hi inorion. Mor copl, uch h Cc, r unbi unr n vn ir rng o cnrio n provi iionl il on h kup o hr nvironnl c h y il b o inr. h principl ron hy r rrly u in bhviorl

17 Bhv Gn gnic i h hy r or iicul o u n h lil n in ily i uibl or hir u. W hop h h currn ppr clrii h rionl or uing n ncourg rrchr o collc n in ily hn circunc rrn hir u. Rrnc Cry G (2005) Cholky probl. Bhv Gn 35: Cl G, Brgr RL (1990) iicl inrnc. Worh, Blon Cloningr CR, Ric J, Rich (1979) Mulicoril inhrinc ih culurl rniion n oriv ing II: gnrl ol o cobin polygnic n culurl inhrinc. J Hu Gn 31: Covnry WL, Kllr MC (2005) iing h n o prr bi in h clicl in ign: coprion o prr i ro n in-ily n clicl in ign. in R Hu Gn 8: Crnokrk P, Ro (1995) oinnc vriion: ociion ih lcion n in. Hriy 75: v LJ (1979) h u o in in h nlyi o oriv ing. Hriy 43: v LJ (2009) Puing h hun bck in gnic: oling h n kinhip o in. in R Hu Gn 12:1 7 v LJ, L K, Young P, Mrin NG (1978) Mol-iing pproch o h nlyi o hun bhvior. Hriy 41: v LJ, ynck HJ, Mrin JM () (1989) Gn, culur, n pronliy: n piricl pproch. cic Pr, Lonong ihr R (1918) h corrlion bn rliv on h uppoiion o Mnlin inhrinc. rn Roy oc inb 52: ulkr W (1982) nion o h clicl in ho. In Bonné- ir B, Cohn, Goon RM () Hun gnic, pr : h unoling gno (Progr in clinicl n biologicl rrch 103). ln R Li, N York, pp Gryon (1989) in rr oghr: iniizing hr nvironnl c. Bhv Gn 19: Hln JB (1932) h cu o voluion. Princon Univriy Pr, Princon, N.J. Hh C, Knlr K, v LJ, Mrkll (1985) h roluion o culurl n biologicl inhrinc: inorivn o irn rlionhip. Bhv Gn 15: Hill WG, Gor M, Vichr PM (2008) n hory poin o inly iiv gnic vrinc or copl ri. PLo Gn 4:1 10 Kllr, M. C. (2007). Pvolv: iulor o gniclly inoriv ipln in R. nnul ing o h bhvior gnic ociion, r Kllr MC, Covnry WL (2005) Quniying n ring prr inrincy in h clicl in ign. in R Hu Gn 8: Kllr MC, Mln, uncn L, Hi PK, Nl MC, M HMM l (2009) Moling n in ily I: cripion o h cc ol. in R Hu Gn 12:8 18 M HMM, Nl MC, Mln, Kllr MC, Mrin NG, Hh C l (2009) libl M pciicion o vriou n in kinhip ign. in R Hu Gn 12:26 34 Mrin NG, Boo I, Mchin G (1997) in-prong ch on copl ri. N Gn 17: Mln, Kllr MC (2009) Moling n in ily II: por oci ih irn ily rucur. in R Hu Gn 12:19 25 Millr G, o PM (1998) M choic urn cogniiv. rn Cogn ci 2: Nnc W, Cory L (1976) Gnic ol or h nlyi o ro h ili o inicl in. Gnic 83: Nl MC (1999) MX: iicl olling, 5h n. prn o Pychiry, Richon, V Nl MC, ulkr W (1984) bivri ph nlyi o r on in n hir prn. c Gnic Mic Gllol (Ro) 33: Oprrio, chnn J, lor, Brig M (2006) Bri rpor: ociion o prnl rh, pr uppor, n gnr ih olcn oionl ir. J olc 29(2): Ploin R, ri JC, McClrn G, McGuin P (2001) Bhviorl gnic, 4h n. Worh Publihr, N York Pohu, Boo I (2000) no on h iicl por in n in ign. Bhv Gn 30: R Cor vlopn (2009) R: lngug n nvironn or iicl copuing. R ounion or iicl Copuing, Vinn, uri Rynol C, Bkr L, Prn NL (2000) Mulivri ol o i orn: phnoypic orn n ocil hoogy or ucion n lui biliy. Bhv Gn 30(6): hin, Grgg B (1980) Hun oriv ing n gnic uilibriu: n voluionry prpciv. hol ociobiol 1: ru KR, v LJ, Wlr, Hh C, Hi JK, Myr JM l (1994) ol y or nlyi o ily rblnc in n kinhip o in. Bhv Gn 24:35 49 Whlbrg P (2009) Chickn gnoic-linkg n QL pping. igil coprhniv uri o Uppl. irion ro h culy o Micin Wrigh (1929) ihr hory o oinnc. N 63: