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Manipula ing gene ic a chi ec u e o e eal i ness ela ionships
© The Au ho s, 2014.
Published e sion
Ke ola, Ta mo; Bo a ynski, Zbigniew; Ko iaho, Janne Saka i
Ke ola, T., Bo a ynski, Z., & Ko iaho, J. S. (2014). Manipula ing gene ic a chi ec u e o e eal
i ness ela ionships. P oceedings o Pee age o Science, 2014(Janua y), A icle e1.
h ps://doi.o g/10.14726/p ocpos.2014.e1
2014
Manipula ing gene ic a chi ec u e o e eal i ness
ela ionships
Ta mo Ke olaTa mo Ke ola11 ,, Zbyszek Bo a yńskiZbyszek Bo a yński11,2,2 && Janne S. Ko iahoJanne S. Ko iaho11,3,3
1Depa men o Biological and En i onmen al Science, P.O. Box 35, 40014, Uni e si y o
Jy äskylä, Finland.
[email p o ec ed]
2CIBIO, Cen o de In es igação em Biodi e sidade e Recu sos Gené icos, Uni e sidade do
Po o, Campus Ag á io de Vai ão, 4485-661, Vai ão, Po ugal.
3Na u al His o y Museum, P.O. Box 35, FI-40014 Uni e si y o Jy äskylä, Finland.
Ci a ion: Ke ola T e al. (2014) Manipula ing gene ic a chi ec u e o e eal i ness ela ionships. P ocPoS 1:e1
DOI: 10.14726/p ocpos.2014.e1
Public Pa on #1:
T ai s a e o en claimed ( a he han shown) o be
closely ela ed o i ness. Some imes his leads o
con as ing p edic ions o he di ec ion o he
associa ion be ween he ai alue and i ness, a
p ime example o which is he deba e o e how
(Da winian) i ness links wi h he a e o ene gy
u no e a es (see: Bu on e al. 2011, o
de ini ions). To esol e his deba e Bu on e al.
(2011) sugges ed manipula ion o ai s and
measu ing changes in i ness. In his pe spec i e, we
p opose ha we should do he opposi e – manipula e
gene ic a chi ec u e unde lying i ness and measu e
he ai s esponse.
E olu iona y heo ies equen ly ocus on he ole
o esou ce alloca ion and esou ce ade-o s in
de e mining i ness ai s (Nilsson 2002;Tomkins e
al. 2004;Bo a yński and Ko eja 2010;Ke ola &
Ko iaho 2009). One common empi ical measu e o
esou ce use is es ing me abolic a e, and he e a e
wo a enues o hinking in how es ing me abolic
a e is ela ed o i ness. High es ing me abolic a e
is ei he assumed o allow highe a es o ene gy
u no e when needed, o high es ing me abolic a e
is expec ed o be nega i ely ela ed o i ness as high
me abolic a e pe se can es ic he use o limi ed
ene gy on i ness ai s (Nilsson 2002;Bo a yński
and Ko eja 2010;Ke ola & Ko iaho 2009).
In e es ingly, he e a e only a hand ul o s udies ha
ha e ho oughly add essed he alidi y o hese
hypo heses and mos o he e idence is co ela i e
(Bu on e al. 2011 see also: Whi e & Kea ney 2012).
To o e come he limi a ion o co ela i e s udies
Bu on e al. (2011) igh ly ad oca ed a manipula i e
app oach. Howe e , hey sugges ed using a i icial
selec ion expe imen s o ho monal manipula ions o
modi y es ing me abolic a e and hen measu ing
changes in i ness o de e mine he ou come o hese
manipula ions. Due o di icul ies in measu ing
i ness, labo ious p ocedu es in ol ed in p oduc ion
o selec ion lines and po en ial con ounding ac o s
included in ho monal manipula ions, hese me hods
may be less han op imal. We ully ag ee ha
manipula ions a e called o , bu p opose
manipula ing gene ic a chi ec u e unde lying i ness
and measu ing he ai s alues, a he han
manipula ing es ing me abolic a e and hen
measu ing i ness.
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P oceedings o Pee age o Science | Janua y 2014 | e1
©Au ho , All
Righ s Rese ed.
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One e y po en , bu unde used, i ness manipula ion
is based on di ec ional dominance and e ealing
hidden ha m ul ecessi e alleles by inb eeding. This
me hod elies on a ew well es ablished p inciples
(Falcone & Mackay 1996;Lynch & Walsh 1998).
By de ini ion, i ness is unde di ec ional selec ion,
which is impo an as condi ions o he build-up
o di ec ional dominance would o he wise no exis .
Wi h a neu al ai , he expec a ion is ha oughly
equal numbe o alleles has posi i e and nega i e
e ec s (Lynch & Walsh 1998, p.257, 270). When
a ai wi h such gene ic backg ound is subjec ed o
di ec ional selec ion o inc eased ai alue,
ecessi e alleles inc easing he ai alue (due o
di ec ion o selec ion: bene icial alleles) will be ixed
apidly. Recessi e alleles ha dec ease ai alue
(ha m ul ecessi es) will be much ha de o emo e
because hey a e e ealed o selec ion only in a small
subse o o sp ing in he popula ion. The e o e, he
likelihood o ha m ul ecessi e alleles emains high
in ai s unde di ec ional selec ion (Falcone &
Mackay 1996;Lynch & Walsh 1998). When he e
is di ec ional selec ion, di ec ional dominance
de elops, and, when e ealed, ecessi e alleles will
always change ai alues in he di ec ion opposing
he long e m pas selec ion, i.e. away om i ness
(Falcone & Mackay 1996;Lynch & Walsh 1998).
The e o e, expe imen ally manipula ed inb eeding
can be used o in e whe he he ocal ai is ela ed
o i ness and i smalle o la ge ai alues ha e
inc eased he i ness.
Al hough, we ad oca e he use o inb eeding me hod
o e ealing i ness associa ions, one needs o be
cau ious when applying i . Fo example he ecessi e
alleles in e y impo an i ness ai s may ha e been
pu ged by s ong di ec ional selec ion, which can
subs an ially lowe inb eeding dep ession and i is
clea ha he me hod should no be applied on
al eady inb ed popula ions, whe e inb eeding
manipula ion does no cause inc eased inb eeding
dep ession. Mo eo e , while inb eeding can be
use ul in e ealing he cou se o pas di ec ional
selec ion, i is also clea ha i ness may be a esul
o op imiza ion o ai exp ession in se e al ai s
(Falcone & Mackay 1996). Explo ing inb eeding on
ai o ai in e ac ions can be used o de e mine
whe he inb eeding is associa ed wi h concu en
changes in o he ai s, as was o example done in
Malle & Chippindale (2011) and in Ke ola &
Ko iaho (2012), es ing also exis ence o non-linea
selec ion behind he i ness. In he simples case,
calcula ing an in e ac ion be ween wo (mean
s anda dized) ai s p o ides a new index ha can
be es ed o a linea associa ion wi h i ness. Fo
example, i selec ion has been agains high me abolic
a e in la ge indi iduals, and agains low me abolic
a es in small indi iduals he magni ude o his index
is inc eased when indi iduals a e inb ed. Thus, he
exis ence o non-linea selec ion is no p e en ing he
use o his me hod. On he con a y, i p o ides ex a
in o ma ion on how ai s in e ac i ely a ec i ness,
ex ending applicabili y o he me hod o mul iple
ai s ha wo k in conce o be e i ness (Ke ola &
Ko iaho 2012).
The inb eeding me hod has been used ew imes
be o e in he con ex o iden i ying sexually selec ed
ai s (see: Co on e al. 2004), in compa ing he
s eng h o selec ion on male and emale ai s
(Malle & Chippindale 2011), and in de e mining
whe he high main enance me abolism inc eases o
dec eases i ness (Ke ola & Ko iaho 2009, 2012;
Bo a yński e al. 2012). Cu en , albei limi ed,
e idence sugges s ha inb eeding inc eases he
main enance me abolism and g ea ly ele a es
exp ession o main enance genes (Ke ola & Ko iaho
2009, 2012;Bo a yński e al. 2012;Ma ila e al.
2012;K is ensen e al. 2005). Thus, i seems ha high
le els o main enance me abolism may be connec ed
o dec eased i ness.
Al hough i ness is he mos impo an concep in
e olu iona y biology, i is a bugge o a concep no
easily amenable o empi ical measu emen . To
o e come he p oblem o measu emen , he
inb eeding me hod desc ibed abo e is use ul and will
p o ide impo an insigh s in o he na u e o he
i ness ela ed ai s. Despi e he a e usage o
inb eeding o esol ing he ai ’s ela ionship wi h
i ness, he e is a la ge body o li e a u e sugges ing
ha li e-his o y ai s ( ha a e closely ela ed o
i ness) a e indeed a ec ed mo e by inb eeding han
o example mo phological ai s ( e iewed in:
DeRose & Ro 1999) showing ha inb eeding
manipula ion is a po en , bu clea ly unde used, ool
in e olu iona y biology.
ACKNOWLEDGEMENTSACKNOWLEDGEMENTS
We hank Academy o Finland o unding and R.
Hegna o e ising he language.
P oceedings o Pee age o Science | Janua y 2014 | e1
©Au ho , All
Righ s Rese ed.
Pu chase a Public Pa on license: h ps://www.pee ageo science.o g/
p oceedings/ge license.php?ms=14 2 o 3
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