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Whole	
  genome	
  duplica0on	
  and	
  
plant	
  genome	
  diversity	
  
Simon	
  Renny-­‐Byfield	
  
Department	
  of	
  Ecology,	
  Evolu0on	
  and	
  Organismal	
  
Biology	
  
Iowa	
  State	
  University	
  	
  
May	
  12th	
  2014	
  
Outline	
  
•  Brief	
  Introduc0on	
  
•  The	
  role	
  polyploidy	
  in	
  plant	
  evolu0on	
  
•  Repe00ve	
  DNA	
  evolu0on	
  in	
  polyploids	
  
•  Evolu0on	
  of	
  gene	
  duplicates	
  in	
  paleopolyploids	
  
•  Genome	
  diploidisa0on	
  and	
  frac0ona0on	
  in	
  
paleopolyploids	
  
•  CoMon	
  fiber	
  transcriptomics	
  and	
  domes0ca0on	
  
Introduc0on	
  
•  What	
  is	
  polyploidy	
  
(whole	
  genome	
  
duplica0on;	
  WGD)?	
  
•  More	
  than	
  a	
  diploid	
  set	
  
of	
  chromosomes	
  
•  Allo	
  vs	
  auto	
  
•  How	
  to	
  iden0fy	
  
polyploids?	
  
Divergence	
  0me	
  (mya)	
  
Introduc0on	
  
•  Chromosome	
  counts	
  	
  
•  Age	
  es0mates	
  of	
  
duplicated	
  genes	
  
•  Syntenty	
  analysis	
  
Jiao	
  et	
  al.,	
  2011	
  Science	
  
Schnable	
  et	
  al.,	
  2011	
  PNAS	
  
•  The	
  greatest	
  
realiza0on	
  of	
  the	
  
plant	
  genomics	
  
era?	
  
Introduc0on	
  
Stebbins	
  (1950)	
  –	
  35%	
  
Grant	
  (1963,1981)	
  –	
  47%	
  
GoldblaM	
  (1980)	
  –	
  70-­‐80%	
  
Lewis	
  (1980)	
  –	
  70-­‐80%	
  
Current	
  view	
  –	
  	
  100%	
  of	
  
seed	
  plants	
  are	
  polyploid	
  	
  
Three	
  brief	
  stories...	
  
① Diversifica0on	
  of	
  polyploid	
  genomes	
  
② Diversifica0on	
  of	
  duplicated	
  genes	
  following	
  
ancient	
  WGD.	
  
③ How	
  polyploids	
  become	
  more	
  diploid-­‐like	
  
again,	
  and	
  again.	
  
1.	
  Diversifica0on	
  of	
  polyploid	
  genomes	
  
•  Polyploid	
  genomes	
  are	
  highly	
  dynamic	
  
– How	
  do	
  they	
  vary?	
  
– Over	
  what	
  0me	
  scale?	
  
– Do	
  different	
  sub-­‐genomes	
  behave	
  differently?	
  
N.	
  sylvestris	
   x	
   N.	
  tomentosiformis	
  
2n	
  =	
  24	
   2n	
  =	
  24	
  
N.	
  tabacum	
  
Genome	
  doubling	
  
2n	
  =	
  48	
  
2650 MB per 1C 2650 MB per 1C
5200 MB per 1C
1.	
  Diversifica0on	
  of	
  polyploid	
  genomes	
  
•  Es0mate	
  repeat	
  content	
  of	
  
progenitors	
  and	
  allopolyploid	
  
– RepeatExplorer	
  pipeline	
  
– Assess	
  divergence	
  of	
  the	
  
allopolyploid	
  from	
  the	
  diploids	
  
Novak	
  et	
  al.,	
  2010	
  BMC	
  Genomics	
  Renny-­‐Byfield	
  et	
  al.,	
  2011	
  MBE	
  
1.	
  Diversifica0on	
  of	
  polyploid	
  genomes	
  
N.	
  tom	
  
S4	
  synthe0c	
  
tobacco	
  
tobacco	
  
N.	
  tom	
  
Renny-­‐Byfield	
  et	
  al.,	
  2012	
  PLoS	
  One	
  
1.	
  Diversifica0on	
  of	
  polyploid	
  genomes	
  
WGDs	
  and	
  genome	
  diversity	
  
The paternal (N. tomentosiformis) genome
appears to be underrepresented in tobacco
Renny-­‐Byfield	
  et	
  al.,	
  2012	
  MBE	
  
2.	
  Diversifica0on	
  of	
  duplicated	
  genes	
  
following	
  ancient	
  WGD	
  
S.	
  C	
  Harland,	
  1936	
  
2.	
  Diversifica0on	
  of	
  duplicated	
  genes	
  
following	
  ancient	
  WGD	
  
•  Neofunc0onaliza0on	
  (Ohno,	
  1970)	
  
•  Subfunc0onaliza0on	
  (Force,	
  Lynch	
  and	
  others)	
  
hMp://www.personal.psu.edu/rua15/Stage3.jpg	
  
2.	
  Diversifica0on	
  of	
  duplicated	
  genes	
  
following	
  ancient	
  WGD	
  
2.	
  Diversifica0on	
  of	
  duplicated	
  genes	
  
following	
  ancient	
  WGD	
  
Dt	
  	
  64	
  	
  	
  37	
  	
  	
  45	
  	
  37	
  	
  65	
  100	
  100	
  	
  0	
  	
  100	
  	
  64	
  	
  	
  51	
  
At	
  	
  36	
  	
  	
  63	
  	
  	
  55	
  	
  	
  63	
  	
  	
  35	
  	
  	
  0	
  	
  	
  	
  	
  0	
  	
  100	
  	
  0	
  	
  	
  	
  36	
  	
  49	
  
Adams	
  et	
  al.,	
  2003	
  
2.	
  Diversifica0on	
  of	
  duplicated	
  genes	
  
following	
  ancient	
  WGD	
  
Renny-­‐Byfield	
  et	
  al.,	
  2014	
  GBE	
  
2.	
  Diversifica0on	
  of	
  duplicated	
  genes	
  
following	
  ancient	
  WGD	
  
2.	
  Diversifica0on	
  of	
  duplicated	
  genes	
  
following	
  ancient	
  WGD	
  
Almost	
  complete	
  divergence	
  in	
  expression	
  aier	
  ca.	
  60	
  my	
  
Renny-­‐Byfield	
  et	
  al.,	
  2014	
  GBE	
  
2.	
  Diversifica0on	
  of	
  duplicated	
  genes	
  
following	
  ancient	
  WGD	
  
Gene	
  (G)	
  effect 	
  Tissue	
  (T)	
  effect	
   	
  G	
  x	
  T	
  interac0on	
  
Renny-­‐Byfield	
  et	
  al.,	
  2014	
  GBE	
  
3.	
  Biased	
  frac0ona0on	
  following	
  WGD	
  
•  What	
  happens	
  to	
  most	
  genes	
  following	
  WGD..	
  
Woodhouse	
  et	
  al.,	
  2010	
  PloS	
  Biology	
  
3.	
  Biased	
  frac0ona0on	
  following	
  WGD	
  
• 	
  CoGe	
  SynMap	
  tool	
  
• 	
  Examine	
  CDS	
  for	
  
colinearity	
  with	
  
reference	
  genome	
  
• 	
  Allows	
  iden0fica0on	
  
of	
  duplicated	
  regions	
  	
  
3.	
  Biased	
  frac0ona0on	
  following	
  WGD	
  
3.	
  Biased	
  frac0ona0on	
  following	
  WGD	
  
• 	
  Ten	
  chromosome	
  level	
  comparisons	
  
• 	
  Significant	
  bias	
  in	
  gene	
  loss	
  in	
  all	
  comparisons	
  
T. cacao
chromosome
G. raimondii
chromosome
(block numbers)
observed predicted !2
p value
2 5 (137,138,139) 929 3641
8 (179,184,185) 642 3641
42.8072 6.1x10-11
6 6 (149,150) 147 2637
9 (190) 580 2637
226.6415 <1x10-15
6 (149,150) 147 2637
10 (33,34,36) 227 2637
15.5573 8x10-5
3.	
  Biased	
  frac0ona0on	
  following	
  WGD	
  
leaf petal seed
0
200
400
600
count(numberofwins)
LF
MF
Over	
  expression	
  of	
  genes	
  on	
  LF	
  chromosomes	
  
leaf petal seed
−2.5
0.0
2.5
5.0
7.5
−5 0 5 −5 0 5 −5 0 5
log(RPKM MF)
log(RPKMLF)
0.02
0.04
0.06
density
3.	
  Biased	
  frac0ona0on	
  following	
  WGD	
  
0
2
4
6
−1000 −500 0 500 1000
distance from transcription start/stop site (bp)
meannumberofmappedreads
Most	
  Frac0onated	
  
Least	
  Frac0onated	
  
24nt	
  siRNAs	
  preferen0ally	
  locate	
  to	
  the	
  MF	
  genome	
  
Current	
  project	
  
• 	
  Two	
  independent	
  domes0ca0on	
  events.	
  
• 	
  One	
  polyploid	
  and	
  one	
  diploid	
  
•  RNAseq	
  at	
  Four	
  development	
  stages:	
  
– 5	
  ,	
  10,	
  15,	
  and	
  20	
  DPA	
  
•  Wild	
  and	
  domes0cated	
  lines:	
  
– Three	
  in	
  each	
  group	
  
•  Polyploid	
  and	
  diploid	
  groups:	
  
– Wild	
  A1,	
  domes0cated	
  A1	
  
– Wild	
  AD1,	
  domes0cated	
  AD1	
  
Current	
  project	
  
•  Gene	
  expression	
  architecture	
  
–  How	
  do	
  transcrip0onal	
  networks	
  
alter	
  (i.e.	
  similar	
  to	
  Swanson-­‐Wagner	
  et	
  al.,2011)	
  
–  connec0vity,	
  edge	
  weight,	
  
movement	
  of	
  nodes.	
  
–  Superimposi0on	
  of	
  graphs	
  to	
  
compare	
  networks	
  in	
  wild	
  and	
  
domes0cated	
  (Lelandias	
  al.,	
  2006,	
  
Bioinforma0cs)	
  
–  Are	
  there	
  parallel	
  changes	
  in	
  
diploid	
  vs	
  polyploid	
  groups	
  
Current	
  project	
  
hMp://www.georgebassellab.com/wp-­‐content/
uploads/2012/01/seedNet.jpg	
  
Conclusions	
  
•  WGD	
  is	
  ubiquitous	
  in	
  angiosperms	
  
•  Polyploid	
  genomes	
  are	
  highly	
  dynamic	
  
•  Parental	
  sub-­‐genomes	
  can	
  behave	
  differently	
  
•  Gene	
  duplica0on	
  (via	
  WGD)	
  can	
  result	
  in	
  biological	
  
novelty	
  
•  Processes	
  of	
  genome	
  turnover	
  and	
  frac0ona0on	
  
result	
  in	
  diploidiza0on	
  
•  Bias	
  frac0ona0on	
  linked	
  to	
  expression	
  and	
  local	
  TE	
  
coverage	
  

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Whole genome duplication and diversification of plant genomes

  • 1. Whole  genome  duplica0on  and   plant  genome  diversity   Simon  Renny-­‐Byfield   Department  of  Ecology,  Evolu0on  and  Organismal   Biology   Iowa  State  University     May  12th  2014  
  • 2. Outline   •  Brief  Introduc0on   •  The  role  polyploidy  in  plant  evolu0on   •  Repe00ve  DNA  evolu0on  in  polyploids   •  Evolu0on  of  gene  duplicates  in  paleopolyploids   •  Genome  diploidisa0on  and  frac0ona0on  in   paleopolyploids   •  CoMon  fiber  transcriptomics  and  domes0ca0on  
  • 3. Introduc0on   •  What  is  polyploidy   (whole  genome   duplica0on;  WGD)?   •  More  than  a  diploid  set   of  chromosomes   •  Allo  vs  auto   •  How  to  iden0fy   polyploids?  
  • 4. Divergence  0me  (mya)   Introduc0on   •  Chromosome  counts     •  Age  es0mates  of   duplicated  genes   •  Syntenty  analysis   Jiao  et  al.,  2011  Science   Schnable  et  al.,  2011  PNAS  
  • 5. •  The  greatest   realiza0on  of  the   plant  genomics   era?   Introduc0on   Stebbins  (1950)  –  35%   Grant  (1963,1981)  –  47%   GoldblaM  (1980)  –  70-­‐80%   Lewis  (1980)  –  70-­‐80%   Current  view  –    100%  of   seed  plants  are  polyploid    
  • 6. Three  brief  stories...   ① Diversifica0on  of  polyploid  genomes   ② Diversifica0on  of  duplicated  genes  following   ancient  WGD.   ③ How  polyploids  become  more  diploid-­‐like   again,  and  again.  
  • 7. 1.  Diversifica0on  of  polyploid  genomes   •  Polyploid  genomes  are  highly  dynamic   – How  do  they  vary?   – Over  what  0me  scale?   – Do  different  sub-­‐genomes  behave  differently?  
  • 8. N.  sylvestris   x   N.  tomentosiformis   2n  =  24   2n  =  24   N.  tabacum   Genome  doubling   2n  =  48   2650 MB per 1C 2650 MB per 1C 5200 MB per 1C 1.  Diversifica0on  of  polyploid  genomes  
  • 9. •  Es0mate  repeat  content  of   progenitors  and  allopolyploid   – RepeatExplorer  pipeline   – Assess  divergence  of  the   allopolyploid  from  the  diploids   Novak  et  al.,  2010  BMC  Genomics  Renny-­‐Byfield  et  al.,  2011  MBE   1.  Diversifica0on  of  polyploid  genomes  
  • 10. N.  tom   S4  synthe0c   tobacco   tobacco   N.  tom   Renny-­‐Byfield  et  al.,  2012  PLoS  One   1.  Diversifica0on  of  polyploid  genomes  
  • 11. WGDs  and  genome  diversity   The paternal (N. tomentosiformis) genome appears to be underrepresented in tobacco Renny-­‐Byfield  et  al.,  2012  MBE  
  • 12. 2.  Diversifica0on  of  duplicated  genes   following  ancient  WGD   S.  C  Harland,  1936  
  • 13. 2.  Diversifica0on  of  duplicated  genes   following  ancient  WGD   •  Neofunc0onaliza0on  (Ohno,  1970)   •  Subfunc0onaliza0on  (Force,  Lynch  and  others)   hMp://www.personal.psu.edu/rua15/Stage3.jpg  
  • 14. 2.  Diversifica0on  of  duplicated  genes   following  ancient  WGD  
  • 15. 2.  Diversifica0on  of  duplicated  genes   following  ancient  WGD   Dt    64      37      45    37    65  100  100    0    100    64      51   At    36      63      55      63      35      0          0    100    0        36    49   Adams  et  al.,  2003  
  • 16. 2.  Diversifica0on  of  duplicated  genes   following  ancient  WGD  
  • 17. Renny-­‐Byfield  et  al.,  2014  GBE   2.  Diversifica0on  of  duplicated  genes   following  ancient  WGD  
  • 18. 2.  Diversifica0on  of  duplicated  genes   following  ancient  WGD   Almost  complete  divergence  in  expression  aier  ca.  60  my   Renny-­‐Byfield  et  al.,  2014  GBE  
  • 19. 2.  Diversifica0on  of  duplicated  genes   following  ancient  WGD   Gene  (G)  effect  Tissue  (T)  effect    G  x  T  interac0on   Renny-­‐Byfield  et  al.,  2014  GBE  
  • 20. 3.  Biased  frac0ona0on  following  WGD   •  What  happens  to  most  genes  following  WGD..   Woodhouse  et  al.,  2010  PloS  Biology  
  • 21. 3.  Biased  frac0ona0on  following  WGD   •   CoGe  SynMap  tool   •   Examine  CDS  for   colinearity  with   reference  genome   •   Allows  iden0fica0on   of  duplicated  regions    
  • 22. 3.  Biased  frac0ona0on  following  WGD  
  • 23. 3.  Biased  frac0ona0on  following  WGD   •   Ten  chromosome  level  comparisons   •   Significant  bias  in  gene  loss  in  all  comparisons   T. cacao chromosome G. raimondii chromosome (block numbers) observed predicted !2 p value 2 5 (137,138,139) 929 3641 8 (179,184,185) 642 3641 42.8072 6.1x10-11 6 6 (149,150) 147 2637 9 (190) 580 2637 226.6415 <1x10-15 6 (149,150) 147 2637 10 (33,34,36) 227 2637 15.5573 8x10-5
  • 24. 3.  Biased  frac0ona0on  following  WGD   leaf petal seed 0 200 400 600 count(numberofwins) LF MF Over  expression  of  genes  on  LF  chromosomes   leaf petal seed −2.5 0.0 2.5 5.0 7.5 −5 0 5 −5 0 5 −5 0 5 log(RPKM MF) log(RPKMLF) 0.02 0.04 0.06 density
  • 25. 3.  Biased  frac0ona0on  following  WGD   0 2 4 6 −1000 −500 0 500 1000 distance from transcription start/stop site (bp) meannumberofmappedreads Most  Frac0onated   Least  Frac0onated   24nt  siRNAs  preferen0ally  locate  to  the  MF  genome  
  • 26.
  • 27. Current  project   •   Two  independent  domes0ca0on  events.   •   One  polyploid  and  one  diploid  
  • 28. •  RNAseq  at  Four  development  stages:   – 5  ,  10,  15,  and  20  DPA   •  Wild  and  domes0cated  lines:   – Three  in  each  group   •  Polyploid  and  diploid  groups:   – Wild  A1,  domes0cated  A1   – Wild  AD1,  domes0cated  AD1   Current  project  
  • 29. •  Gene  expression  architecture   –  How  do  transcrip0onal  networks   alter  (i.e.  similar  to  Swanson-­‐Wagner  et  al.,2011)   –  connec0vity,  edge  weight,   movement  of  nodes.   –  Superimposi0on  of  graphs  to   compare  networks  in  wild  and   domes0cated  (Lelandias  al.,  2006,   Bioinforma0cs)   –  Are  there  parallel  changes  in   diploid  vs  polyploid  groups   Current  project   hMp://www.georgebassellab.com/wp-­‐content/ uploads/2012/01/seedNet.jpg  
  • 30. Conclusions   •  WGD  is  ubiquitous  in  angiosperms   •  Polyploid  genomes  are  highly  dynamic   •  Parental  sub-­‐genomes  can  behave  differently   •  Gene  duplica0on  (via  WGD)  can  result  in  biological   novelty   •  Processes  of  genome  turnover  and  frac0ona0on   result  in  diploidiza0on   •  Bias  frac0ona0on  linked  to  expression  and  local  TE   coverage