ID C5-Sy23-i031-R Addressing,clinalvariaon...

1
Significant differen.a.on was found between popula.ons from the three loca.ons, even a;er 6 genera.ons in a common environment (pairwise F st , P<0.05). In contrast, no differences were observed between popula.ons of the same origin across genera.ons (pairwise F st ,, P >0.05). Nevertheless, the J and U chromosomes significantly changed the frequency of their inversions between genera.ons 2 and 6 in Montpellier popula.ons (MANOVA tests, P<0.05). In par.cular, J 1 , J ST and U 1+2 showed consistent changes in two out of three replicates. Also, the O 3+4 arrangement presented a consistent increase in two of the Adraga replicates (see arrows in figure above; Post hoc DunneQ test, P <0.05). Addressing clinal varia<on: the rela<ve role of evolu<onary history and adapta<on to a novel environment Simões, P. , Fragata, I., Lima, M., LopesCunha, M., Kellen, B., Bárbaro A. M., Santos, J. and Matos, M. Faculdade de Ciências da Universidade de Lisboa, Centro de Biologia Ambiental, Departamento de Biologia Animal, Lisboa, Portugal Final remarks Acknowledgments This study was partially financed by “Fundação para a Ciência e Tecnologia” (FCT) project nº PTDC/BIA-BEC/098213/2008. I.F. has a PhD grant (SFRH/BD/60734/2009) and J.S. had a PhD grant (SFRH/BD/28498/2006) from FCT. P.S. has a Post-Doc grant, M.C. has a BI grant and B. K. and M. L. had BTI grants from FCT. References Balanyà, J., J. M. Oller, R. B. Huey, G. W. Gilchrist & L. Serra (2006) Global genetic change tracks global climate warming in Drosophila subobscura. Science 313: 5794. La.tudinal clines are wellknown phenomena, although it remains unclear how and why they are maintained. Chromosomal inversions (arrangements) in Drosophila are one of the most emblema.c cases of such clinal varia.on and strong evidence supports an adap.ve cause for the evolu.on of these polymorphisms. Local adapta.on to heterogeneous environments has been advanced as a possible explana.on. Furthermore, recent work shows that these arrangements are responding to global warming at a worldwide scale (Balanyà et al. 2006). Nevertheless, the underlying adap.ve processes are s.ll largely unknown. The study of popula.ons ini.ally differen.ated throughout a cline, when adap.ng to a novel common environment may enlighten this issue. Will they converge? Here we present the first snapshot of a longer term study addressing temporal paQerns of varia.on in inversion frequencies of Drosophila subobscura popula.ons during laboratory adapta.on. Inversions in time and space A Principal Component Analysis was also applied to the frequencies of all inversions a;er transforma.on (Balanyà et al., 2006). The first axis clearly separates popula.ons according to their loca.on of origin, with Adraga popula.ons more differen.ated from the others. The second axis is associated with the number of genera.ons in the laboratory, with Adraga again sugges.ng a higher temporal varia.on. Exploring Inversions Drosophila subobscura individuals were collected in Adraga (Portugal), Montpellier (France) and Groningen (Netherlands) along a la.tudinal cline. Popula.ons were scored for inversion polymorphism at genera.ons 2 and 6 a;er laboratory introduc.on. Popula.ons were maintained at 18 o C with controlled densi.es, and were threefold replicated at genera.on 4. To score the inversion polymorphisms, 60 or more males of each popula.on were crossed with females of the cherry curled strain that is homokaryo.pic for the ‘standard’ arrangement in all chromosomes except the O which is O 3+4. When analyzing the weight of different arrangements in the popula.on differen.a.on we observe that the standard inversions are clearly associated with the northern popula.ons of Groningen, while warmer adapted arrangements such as A 2 , J 1 and O 3+4+7 help to explain the higher differen.a.on of the southern popula.ons of Adraga from the other 2 founda.ons. U 1+2 and rare arrangements of the O chromosome appear more closely related with the Montpellier founda.on. ID C5-Sy23-i031-R [email protected] WILL THIS LEAD TO CONVERGENT EVOLUTION? ONLY MORE GENERATIONS WILL TELL... There are clear ini.al differences in inversion polymorphism which have a nonlinear associa.on with la.tude. Is this due to historical effects of geographic barriers? Modest temporal changes occur in inversion frequencies a;er 6 genera.ons of laboratory evolu.on. Contras.ng paQerns between founda.ons changes in the O chromosome for Adraga and J and U for Montpellier Principal Component Analysis Ad1_G6 Ad2_G6 Ad3_G6 Mo1_G6 Mo2_G6 Mo3_G6 Gro1_G6 Gro2_G6 Gro3_G6 Ad_G2 Mo_G2 Gro_G2 -10 -8 -6 -4 -2 0 2 4 6 8 Factor 1: 47.90% -6 -5 -4 -3 -2 -1 0 1 2 3 4 Factor 2: 15.40% Ad1_G6 Ad2_G6 Ad3_G6 Mo1_G6 Mo2_G6 Mo3_G6 Gro1_G6 Gro2_G6 Gro3_G6 Ad_G2 Mo_G2 Gro_G2 Projection of the arrangements on the factor-plane ( 1 x 2) A1 A2 Ast E1+2 E1+2+9 E1+2+9+12 E1+2+9+3 E8 Est J1 Jst O3+4 O3+4+1 O3+4+12 O3+4+13+12 O3+4+2 O3+4+22 O3+4+23+2 O3+4+23+22 O3+4+6 O3+4+7 O3+4+8 O6 O7 Ost U1 U1+2 U1+2+8 Ust -1,0 -0,5 0,0 0,5 1,0 Factor 1 : 47.90% -1,0 -0,5 0,0 0,5 1,0 Factor 2 : 15.40% A1 A2 Ast E1+2 E1+2+9 E1+2+9+12 E1+2+9+3 E8 Est J1 Jst O3+4 O3+4+1 O3+4+12 O3+4+13+12 O3+4+2 O3+4+22 O3+4+23+2 O3+4+23+22 O3+4+6 O3+4+7 O3+4+8 O6 O7 Ost U1 U1+2 U1+2+8 Ust

Transcript of ID C5-Sy23-i031-R Addressing,clinalvariaon...

Page 1: ID C5-Sy23-i031-R Addressing,clinalvariaon ...mmmatos/Eseb2011_inv.pdfSignificantdifferen.aon)was)found)between)populaons)from)the)three)locaons,)even)aer)6)generaons)in)acommon)environment(pairwise)

Significant   differen.a.on   was   found   between   popula.ons   from   the   three   loca.ons,   even   a;er   6   genera.ons   in   a   common   environment   (pairwise   Fst,   P<0.05).   In   contrast,   no  

differences  were   observed   between   popula.ons   of   the   same   origin   across   genera.ons   (pairwise   Fst,,  P   >0.05).   Nevertheless,   the   J   and  U   chromosomes   significantly   changed   the  

frequency  of  their  inversions  between  genera.ons  2  and  6  in  Montpellier  popula.ons  (MANOVA  tests,  P<0.05).  In  par.cular,  J1,  JST  and  U1+2  showed  consistent  changes  in  two  out  of  

three  replicates.  Also,  the  O3+4  arrangement  presented  a  consistent  increase  in  two  of  the  Adraga  replicates  (see    arrows  in  figure  above;  Post  hoc  DunneQ  test,  P  <0.05).  

 Addressing  clinal  varia<on:  the  rela<ve  role  of  evolu<onary  

history  and  adapta<on  to  a  novel  environment        

Simões,  P.,  Fragata,  I.,  Lima,  M.,  Lopes-­‐Cunha,  M.,  Kellen,  B.,  Bárbaro  A.  M.,  Santos,  J.  and  Matos,  M.    

Faculdade  de  Ciências  da  Universidade  de  Lisboa,  Centro  de  Biologia  Ambiental,  Departamento  de  Biologia  Animal,  Lisboa,  Portugal    

Final remarks

Acknowledgments This study was partially financed by “Fundação para a Ciência e Tecnologia” (FCT) project nº PTDC/BIA-BEC/098213/2008. I.F. has a PhD grant (SFRH/BD/60734/2009) and J.S. had a PhD grant (SFRH/BD/28498/2006) from FCT. P.S. has a Post-Doc grant, M.C. has a BI grant and B. K. and M. L. had BTI grants from FCT. References Balanyà, J., J. M. Oller, R. B. Huey, G. W. Gilchrist & L. Serra (2006) Global genetic change tracks global climate warming in Drosophila subobscura. Science 313: 5794.

La.tudinal  clines  are  well-­‐known  phenomena,  although  it  remains  unclear  how  and  why  they  are  maintained.  Chromosomal  inversions  (arrangements)   in  

Drosophila   are   one   of   the  most   emblema.c   cases   of   such   clinal   varia.on   and   strong   evidence   supports   an   adap.ve   cause   for   the   evolu.on   of   these  

polymorphisms.  Local  adapta.on  to  heterogeneous  environments  has  been  advanced  as  a  possible  explana.on.  Furthermore,  recent  work  shows  that  these  

arrangements  are  responding  to  global  warming  at  a  worldwide  scale  (Balanyà  et  al.  2006).  Nevertheless,  the  underlying  adap.ve  processes  are  s.ll  largely  

unknown.  The  study  of  popula.ons  ini.ally  differen.ated  throughout  a  cline,  when  adap.ng  to  a  novel  common  environment  may  enlighten  this  issue.  Will  

they  converge?  Here  we  present  the  first  snapshot  of  a  longer  term  study  addressing  temporal  paQerns  of  varia.on  in  inversion  frequencies  of  Drosophila  

subobscura  popula.ons  during  laboratory  adapta.on.  

Inversions in time and space

A   Principal   Component   Analysis   was   also   applied   to   the  

frequencies  of  all  inversions  a;er  transforma.on  (Balanyà  

et   al.,   2006).   The   first   axis   clearly   separates   popula.ons  

according   to   their   loca.on   of   origin,   with   Adraga  popula.ons   more   differen.ated   from   the   others.   The  

second  axis  is  associated  with  the  number  of  genera.ons  

in  the  laboratory,  with  Adraga  again  sugges.ng  a  higher  temporal  varia.on.  

Exploring Inversions

Drosophila   subobscura   individuals   were   collected   in   Adraga   (Portugal),  Montpellier   (France)   and   Groningen   (Netherlands)   along   a   la.tudinal   cline.  Popula.ons   were   scored   for   inversion   polymorphism   at   genera.ons   2   and   6  a;er   laboratory   introduc.on.   Popula.ons   were   maintained   at   18oC   with  controlled  densi.es,  and  were  three-­‐fold  replicated  at  genera.on  4.  

 

To   score   the   inversion   polymorphisms,   60   or  more  males   of   each   popula.on  were  crossed  with  females  of  the  cherry  curled  strain  that  is  homokaryo.pic  for  the  ‘standard’  arrangement  in  all  chromosomes  except  the  O  which  is  O3+4.    

When  analyzing   the  weight  of  different  arrangements   in   the  popula.on  

differen.a.on   we   observe   that   the   standard   inversions   are   clearly  

associated  with  the  northern  popula.ons  of  Groningen,  while  warmer-­‐

adapted   arrangements   such   as  A2,   J1   and  O3+4+7   help   to   explain   the  

higher  differen.a.on  of   the   southern  popula.ons  of  Adraga   from   the  

other  2  founda.ons.  U1+2  and  rare  arrangements  of  the  O  chromosome  

appear  more  closely  related  with  the  Montpellier  founda.on.  

ID C5-Sy23-i031-R [email protected]

WILL  THIS  LEAD  TO  CONVERGENT  EVOLUTION?    ONLY  MORE  GENERATIONS  WILL    TELL...  

•   There  are  clear  ini.al  differences  in  inversion  polymorphism  which  have  a  non-­‐linear  associa.on  with  la.tude.    

è Is  this  due  to  historical  effects  of  geographic  barriers?  

•   Modest  temporal  changes  occur  in  inversion  frequencies  a;er  6  genera.ons  of  laboratory  evolu.on.  è Contras.ng  paQerns  between  founda.ons  -­‐  changes  in  the  O  chromosome  for  Adraga  and  J  and  U  for    Montpellier    

Principal Component Analysis

Ad1_G6

Ad2_G6

Ad3_G6

Mo1_G6 Mo2_G6

Mo3_G6

Gro1_G6

Gro2_G6

Gro3_G6

Ad_G2

Mo_G2

Gro_G2

-10 -8 -6 -4 -2 0 2 4 6 8

Factor 1: 47.90%

-6

-5

-4

-3

-2

-1

0

1

2

3

4

Fact

or 2

: 15.

40%

Ad1_G6

Ad2_G6

Ad3_G6

Mo1_G6 Mo2_G6

Mo3_G6

Gro1_G6

Gro2_G6

Gro3_G6

Ad_G2

Mo_G2

Gro_G2

Projection of the arrangements on the factor-plane ( 1 x 2)

A1

A2 Ast

E1+2 E1+2+9

E1+2+9+12

E1+2+9+3

E8

Est

J1

Jst

O3+4

O3+4+1

O3+4+12

O3+4+13+12

O3+4+2

O3+4+22

O3+4+23+2 O3+4+23+22

O3+4+6

O3+4+7

O3+4+8 O6

O7

Ost

U1

U1+2

U1+2+8

Ust

-1,0 -0,5 0,0 0,5 1,0

Factor 1 : 47.90%

-1,0

-0,5

0,0

0,5

1,0

Fact

or 2

: 15

.40%

A1

A2 Ast

E1+2 E1+2+9

E1+2+9+12

E1+2+9+3

E8

Est

J1

Jst

O3+4

O3+4+1

O3+4+12

O3+4+13+12

O3+4+2

O3+4+22

O3+4+23+2 O3+4+23+22

O3+4+6

O3+4+7

O3+4+8 O6

O7

Ost

U1

U1+2

U1+2+8

Ust