Unraveling the evolutionary history of wild potatoes and...
Transcript of Unraveling the evolutionary history of wild potatoes and...
Unraveling the evolutionary history ofUnraveling the evolutionary history of wild potatoes and tomatoeswild potatoes and tomatoes
FlorFlor RodriguezRodriguez
July 17July 17thth, 2009, 2009
OutlineOutline
Solanum subgenus Potatoe
Wild potato and wild tomato classifications
Previous phylogenetic studies
Materials and methods
Results and discussion
SolanumSolanum subgenus subgenus PotatoePotatoe
Family Family SolanaceaeSolanaceaeGenus Genus SolanumSolanum
Subgenus Subgenus PotatoePotatoeSection Section LycopersiconLycopersiconSection Section EtuberosumEtuberosumSection Section PetotaPetota
Wild potato and tomato distributionsWild potato and tomato distributions
Spooner and Hijmans, 2001 Peralta et al., 2008
potato tomato
Morphological variability in wild potatoMorphological variability in wild potato
Morphological variability in wild tomatoMorphological variability in wild tomato
Hawkes Hawkes phylogenetic/biogeographicphylogenetic/biogeographic hypothesis of hypothesis of genome evolution and migration in genome evolution and migration in SolanumSolanum sect. sect. PetotaPetota
Hawkes, 1990; Spooner and Hawkes, 1990; Spooner and SytsmaSytsma, 1992, 1992
““BB”” genome prototypegenome prototype
““BB”” genome migrationgenome migration
““AA”” genome evolvesgenome evolves““AA””, , ““AAAA””
““AA”” genomegenomemigrationmigration
““AA”” xx ““BB””allopolyploidsallopolyploids
evolveevolve
Previous molecular studiesPrevious molecular studies
HosakaHosaka et alet al., 1984; Spooner & ., 1984; Spooner & SytsmaSytsma, 1992;, 1992;Castillo & Spooner, 1997; Castillo & Spooner, 1997; RodrRodrííguezguez & Spooner, 1997& Spooner, 1997
Clade 2
Clade 3
Clade 4
Clade 1
Outgroup (S. palustre)
all ser. Piuranasome from ser. Megistacroloba & YungasensaDiploids ser. Conicibaccata & TuberosaRemaining SA speciesNA & CA polyploid speciesand S. verrucosum
Most US, Mexican & Central American diploids
S. bulbocastanumS. cardiophyllum subsp. cardiophyllumS. cardiophyllum subsp. ehrenbergii
plastid DNA restriction siteplastid DNA restriction site
Comparison of classifications of wild Comparison of classifications of wild tomatoestomatoes
Series Series LycopersiconLycopersicon
Series Series NeolycopersiconNeolycopersicon
Series Series EriopersiconEriopersicon
Muller,1940Muller,1940
Subgenus Subgenus EulycopersiconEulycopersiconL. L. esculentumesculentumL. L. pimpinellifoliumpimpinellifolium
Subgenus Subgenus EriopersiconEriopersiconL. L. peruvianumperuvianum
L. L. cheesmaniaecheesmaniaeL. L. hirsutumhirsutumL. L. glandulosumglandulosum
Rick, 1979Rick, 1979
EsculentumEsculentum complexcomplex
PeruvianumPeruvianum complexcomplex
L. L. pennelliipennellii
L. L. chmielewskiichmielewskiiL. L. parviflorumparviflorum
L. L. cheesmaniaecheesmaniae
L. L. hirsutumhirsutum
L. L. chilensechilenseL. L. peruvianumperuvianum
L. L. esculentumesculentumL. L. pimpinellifoliumpimpinellifolium
Child, 1990Child, 1990
S. S. pennelliipennellii
S. S. habrochaiteshabrochaitesS. S. chmielewskiichmielewskiiS. S. neorickiineorickii
S. S. lycopersicumlycopersicumS. S. pimpinellifoliumpimpinellifoliumS. S. cheesmaniaecheesmaniae
S. S. chilensechilenseS. S. peruvianumperuvianum
Peralta et al. 2008Peralta et al. 2008
LycopersiconLycopersicon groupgroupS. S. lycopersicumlycopersicumS. S. pimpinellifoliumpimpinellifoliumS. S. cheesmaniaecheesmaniaeS. S. galapagensegalapagense
NeolycopersiconNeolycopersicon groupgroupS. S. pennelliipennellii
EriopersiconEriopersicon groupgroupS. S. habrochaiteshabrochaitesS. S. huaylasensehuaylasenseS. S. corneliomullericorneliomulleriS. S. peruvianumperuvianumS. S. chilensechilense
Arcanum Arcanum groupgroupS. S. arcanumarcanumS. S. chmielewskiichmielewskiiS. S. neorickiineorickii
WaxyWaxyPeralta & Spooner, 2001
S.lycopersicumS.lycopersicumS.pimpinellifoliumS.pimpinellifoliumS.cheesmaniaeS.cheesmaniaeS.peruvianumS.peruvianum NNS.chmielewskiiS.chmielewskiiS.neorickiiS.neorickiiS.peruvianumS.peruvianum SSS.chilenseS.chilenseS.habrochaitesS.habrochaitesS.pennelliiS.pennelliiS.juglandifoliumS.juglandifoliumS.ochranthumS.ochranthumS.lycopersicoidesS.lycopersicoidesS.sitiensS.sitiensS.tuberosumS.tuberosumS.bulbocastanumS.bulbocastanumS.jamesiiS.jamesiiS.etuberosumS.etuberosumS.palustreS.palustreS.muricatumS.muricatum
24
70
100
100
100
64
86
97
100
100
ITSITSMarshall & all, 2001
S.pimpinellifoliumS.pimpinellifolium
S.lycopersicumS.lycopersicum
S.lycopersicumS.lycopersicum
S.cheesmaniaeS.cheesmaniae
S.chmielewskiiS.chmielewskii
S.neorickiiS.neorickii
Cestrum, Cestrum, NicotianaNicotiana, Petunia, Petunia
S.habrochaitesS.habrochaites
S.pennelliiS.pennellii
S.lycopersicoidesS.lycopersicoides
S.nigrumS.nigrum
S.peruvianumS.peruvianum NNS.peruvianumS.peruvianum NN
S.peruvianumS.peruvianum SS
S.chilenseS.chilense
54
67
76
85
69
99
99
MorphologyMorphologyPeralta & Spooner, 2005
S.lycopersicumS.lycopersicum
S.pimpinellifoliumS.pimpinellifolium
S.cheesmaniaeS.cheesmaniae
S.galapagenseS.galapagense
S.chmielewskiiS.chmielewskii
S.neorickiiS.neorickii
S.peruvianumS.peruvianum NN
S.peruvianumS.peruvianum SS
S.habrochaitesS.habrochaites
S.chilenseS.chilense
S.pennelliiS.pennellii
S.juglandifoliumS.juglandifolium
S.ochranthumS.ochranthum
S.lycopersicoidesS.lycopersicoides
S.sitiensS.sitiens
87
88
76
72
96
99
99
100
Previous studiesPrevious studies
Molecular phylogeny of potato genomes Molecular phylogeny of potato genomes with emphasis on origins of the with emphasis on origins of the polyploid speciespolyploid species
ObjectivesObjectives
Generate a nuclear molecular data set to reconstruct the phylogeny of wild potatoes
Evaluate previous circumscription of potato genomes
Infer the genomic contributors of potato polyploids
Materials and MethodsMaterials and Methods14 series, 45 ingroups and 3 outgroups
GenBank nitrate reductase tomato sequence (X14060)
Cloning
Exon 1 Exon 4E2 E3i1i1 i2i2 i3i3
1227 bp
NIA3F
NIA3R
74 bp74 bp 847 bp847 bp 436 bp436 bp
NIA 3
Data AnalysisData Analysis
Sequence editing and alignment
Phylogenetic analysis:ParsimonyMaximum LikelihoodBayesian
Hypothesis testing
Summary of sequence variation among Summary of sequence variation among 1) diploids and 1) diploids and 2) diploids and polyploids2) diploids and polyploids
Total characters Variable characters
bp bp
exon 816 95 35 (4%)intron 626 182 81 (13%)DNA 1442 277 116 (8%)gaps 78 78 35 (45%)DNA & gaps 1520 355 151(10%)exon 816 129 64 (8%)intron 629 211 110 (18%)DNA 1445 340 174 (12%)gaps 87 87 42 (48%)DNA & gaps 1532 427 216 (14%)
Diploids and polyploids
Data set Region Number and Percentage of Parsimony Informative characters
Diploids
104 MP treesL = 5112CI = 0.764RI = 0.779
Maximum Parsimony Maximum Parsimony diploidsdiploids
NA & CA diploidsB genome
A genomeRemaining SA diploids
Piurana cladeP, C, D genome
Clade 3
Clade 1+2
Clade 4
ConclusionsConclusions1. Wild potato species are divided into three well-
defined clades
2. The North and Central American B genome species are well distinguish from SA species
3. S. verrucosum clusters with all SA diploid species (A genome)
4. Hawkes’s series PIN and POL are monophyletic
5. S. brevicaule and S. sparsipilum appear to have hybrid origins
6. Species did not group based on Hawkes’s four superseries (flower shape)
NIA data suggests:Monophyly of North and Central American diploid species exclusive of S. verrucosum
Plastid DNA suggested:S. cardiophyllum and S. bulbocastanum cluster with SA diploid species
ries
abz PIU AR 2adr TUB PR 2
chm CON PR 2imt TUB PR 2
sph PIN PS 1blb BUL PS 1
cph PIN PS 1ehr1 PIN PS 1ehr2 PIN PS 1
jam PIN PS 1
pnt PIN PS 1trf PIN PS 1
clr BUL PS 1mrl MOR PS 1les POL PS 1pld POL PS 1ber TUB PR 2tar YNG AS 2ifd1 CUN PR 2blv MEG PR 2mga MEG PR 2chc YNG AS 2gnd TUB PR 2brc1 TUB PR 2ifd2 CUN PR 2spl1 TUB PR 2
buk TUB PR 2
lgl LIG PS 1
mcd TUB PR 2
rap1 MEG PR 2rap2 MEG PR 2vio CON PR 2
brc2 TUB PR 2lph TUB PR 2spl2 TUB PR 2ver TUB AR 2lycetb ETU 1pls ETU 1
92
77
61
92
59
100
100
50
92
98
57
52
56
92
88
6256
100
88
SpeciesSeries
Superseries
EBN
Piurana clade
NA & CA diploids B genome
Remaining SA diploids
NA & CA diploidsB genome
A genomeRemaining SA diploids
Piurana cladeP, C, D genome
Clade 3
Clade 1+2
Clade 4
NA & CA diploidsB genome
A genomeRemaining SA diploids
Piurana cladeP, C, D genome
Clade 3Piurana clade
P, C, D genomePiurana clade
P, C, D genome
Clade 3
Clade 1+2
Clade 4
Maximum Parsimony Maximum Parsimony diploidsdiploids
abz PIU AR 2adr TUB PR 2
chm CON PR 2imt TUB PR 2
sph PIN PS 1blb BUL PS 1
cph PIN PS 1ehr1 PIN PS 1ehr2 PIN PS 1
jam PIN PS 1
pnt PIN PS 1trf PIN PS 1
clr BUL PS 1mrl MOR PS 1les POL PS 1pld POL PS 1ber TUB PR 2tar YNG AS 2ifd1 CUN PR 2blv MEG PR 2mga MEG PR 2chc YNG AS 2gnd TUB PR 2brc1 TUB PR 2ifd2 CUN PR 2spl1 TUB PR 2
buk TUB PR 2
lgl LIG PS 1
mcd TUB PR 2
rap1 MEG PR 2rap2 MEG PR 2vio CON PR 2
brc2 TUB PR 2lph TUB PR 2spl2 TUB PR 2ver TUB AR 2lycetb ETU 1pls ETU 1
92
77
61
92
59
100
100
50
92
98
57
52
56
92
88
6256
100
88
SpeciesSeries
Superseries
EBN
Piurana clade
NA & CA diploids B genome
Remaining SA diploids
Maximum Parsimony Maximum Parsimony diploidsdiploids
Monophyly of NA and Central American diploid species
Clade 3Piurana clade
P, C, D genome
Clade 1+2NA & CA diploids
B genome
Clade 4Remaining SA diploidsA genome
ConclusionsConclusions
1. Wild potato species are divided into three clades well defined
2. The North and Central American B genome species are well distinguish from SA species
3. S. verrucosum clusters with all SA diploid species (A genome)
4. Hawkes’s series PIN and POL are monophyletic
5. S. brc and spl appear to have hybrid origins
6. Species did not group based on Hawkes’s four superseries based on flower shape
abz PIU AR 2adr TUB PR 2
chm CON PR 2imt TUB PR 2
sph PIN PS 1blb BUL PS 1
cph PIN PS 1ehr1 PIN PS 1ehr2 PIN PS 1
jam PIN PS 1
pnt PIN PS 1trf PIN PS 1
clr BUL PS 1mrl MOR PS 1les POL PS 1pld POL PS 1ber TUB PR 2tar YNG AS 2ifd1 CUN PR 2blv MEG PR 2mga MEG PR 2chc YNG AS 2gnd TUB PR 2brc1 TUB PR 2ifd2 CUN PR 2spl1 TUB PR 2
buk TUB PR 2
lgl LIG PS 1
mcd TUB PR 2
rap1 MEG PR 2rap2 MEG PR 2vio CON PR 2
brc2 TUB PR 2lph TUB PR 2spl2 TUB PR 2ver TUB AR 2lycetb ETU 1pls ETU 1
92
77
61
92
59
100
100
50
92
98
57
52
56
92
88
6256
100
88
SpeciesSeries
Superseries
EBN
Piurana clade
NA & CA diploids B genome
Remaining SA diploids
Maximum Parsimony Maximum Parsimony diploidsdiploids
Monophyly of ser. TUB?
Monophyly of ser. CON ?
Clade 3Piurana clade
P, C, D genome
Clade 1+2NA & CA diploids
B genome
Clade 4Remaining SA diploidsA genome
Species in clade 3
abz PIU AR 2adr TUB PR 2
chm CON PR 2imt TUB PR 2
sph PIN PS 1blb BUL PS 1
cph PIN PS 1ehr1 PIN PS 1ehr2 PIN PS 1
jam PIN PS 1
pnt PIN PS 1trf PIN PS 1
clr BUL PS 1mrl MOR PS 1les POL PS 1pld POL PS 1ber TUB PR 2tar YNG AS 2ifd1 CUN PR 2blv MEG PR 2mga MEG PR 2chc YNG AS 2gnd TUB PR 2brc1 TUB PR 2ifd2 CUN PR 2spl1 TUB PR 2
buk TUB PR 2
lgl LIG PS 1
mcd TUB PR 2
rap1 MEG PR 2rap2 MEG PR 2vio CON PR 2
brc2 TUB PR 2lph TUB PR 2spl2 TUB PR 2ver TUB AR 2lycetb ETU 1pls ETU 1
92
77
61
92
59
100
100
50
92
98
57
52
56
92
88
6256
100
88
SpeciesSeries
Superseries
EBN
Piurana clade
NA & CA diploids B genome
Remaining SA diploids
Maximum Parsimony Maximum Parsimony diploidsdiploids
Monophyly of series Monophyly of series MEGMEG and and YNG ?YNG ?
Clade 3Piurana clade
P, C, D genome
Clade 1+2NA & CA diploids
B genome
Clade 4Remaining SA diploidsA genome
abz PIU AR 2adr TUB PR 2
chm CON PR 2imt TUB PR 2
sph PIN PS 1blb BUL PS 1
cph PIN PS 1ehr1 PIN PS 1ehr2 PIN PS 1
jam PIN PS 1
pnt PIN PS 1trf PIN PS 1
clr BUL PS 1mrl MOR PS 1les POL PS 1pld POL PS 1ber TUB PR 2tar YNG AS 2ifd1 CUN PR 2blv MEG PR 2mga MEG PR 2chc YNG AS 2gnd TUB PR 2brc1 TUB PR 2ifd2 CUN PR 2spl1 TUB PR 2
buk TUB PR 2
lgl LIG PS 1
mcd TUB PR 2
rap1 MEG PR 2rap2 MEG PR 2vio CON PR 2
brc2 TUB PR 2lph TUB PR 2spl2 TUB PR 2ver TUB AR 2lycetb ETU 1pls ETU 1
92
77
61
92
59
100
100
50
92
98
57
52
56
92
88
6256
100
88
SpeciesSeries
Superseries
EBN
Piurana clade
NA & CA diploids B genome
Remaining SA diploids
Maximum Parsimony Maximum Parsimony diploidsdiploids
ser. PIN is supported as monophyletic with the exclusion of S. bulbocastanum
Clade 3Piurana clade
P, C, D genome
Clade 1+2NA & CA diploids
B genome
Clade 4Remaining SA diploidsA genome
abz PIU AR 2adr TUB PR 2
chm CON PR 2imt TUB PR 2
sph PIN PS 1blb BUL PS 1
cph PIN PS 1ehr1 PIN PS 1ehr2 PIN PS 1
jam PIN PS 1
pnt PIN PS 1trf PIN PS 1
clr BUL PS 1mrl MOR PS 1les POL PS 1pld POL PS 1ber TUB PR 2tar YNG AS 2ifd1 CUN PR 2blv MEG PR 2mga MEG PR 2chc YNG AS 2gnd TUB PR 2brc1 TUB PR 2ifd2 CUN PR 2spl1 TUB PR 2
buk TUB PR 2
lgl LIG PS 1
mcd TUB PR 2
rap1 MEG PR 2rap2 MEG PR 2vio CON PR 2
brc2 TUB PR 2lph TUB PR 2spl2 TUB PR 2ver TUB AR 2lycetb ETU 1pls ETU 1
92
77
61
92
59
100
100
50
92
98
57
52
56
92
88
6256
100
88
SpeciesSeries
Superseries
EBN
Piurana clade
NA & CA diploids B genome
Remaining SA diploids
Maximum Parsimony Maximum Parsimony diploidsdiploids
Monophyly of ser. BUL ?
Clade 3Piurana clade
P, C, D genome
Clade 1+2NA & CA diploids
B genome
Clade 4Remaining SA diploidsA genome
ConclusionsConclusions
1. Wild potato species are divided into three clades well defined
2. The North and Central American B genome species are well distinguish from SA species
3. S. verrucosum cluster with all SA diploid species (A genome)
4. Hawkes’s series PIN and POL are monophyletic
5. S. brc and spl appear to have hybrid origins
6. Species did not group based on Hawkes’s four superseries based on flower shape
abz PIU AR 2adr TUB PR 2
chm CON PR 2imt TUB PR 2
sph PIN PS 1blb BUL PS 1
cph PIN PS 1ehr1 PIN PS 1ehr2 PIN PS 1
jam PIN PS 1
pnt PIN PS 1trf PIN PS 1
clr BUL PS 1mrl MOR PS 1les POL PS 1pld POL PS 1ber TUB PR 2tar YNG AS 2ifd1 CUN PR 2blv MEG PR 2mga MEG PR 2chc YNG AS 2gnd TUB PR 2brc1 TUB PR 2ifd2 CUN PR 2spl1 TUB PR 2
buk TUB PR 2
lgl LIG PS 1
mcd TUB PR 2
rap1 MEG PR 2rap2 MEG PR 2vio CON PR 2
brc2 TUB PR 2lph TUB PR 2spl2 TUB PR 2ver TUB AR 2lycetb ETU 1pls ETU 1
92
77
61
92
59
100
100
50
92
98
57
52
56
92
88
6256
100
88
SpeciesSeries
Superseries
EBN
Piurana clade
NA & CA diploids B genome
Remaining SA diploids
Maximum Parsimony Maximum Parsimony diploidsdiploids
Monophyly of two distinct alleles from the same species ?
Clade 4a
Clade 4b
Clade 3Piurana clade
P, C, D genome
Clade 1+2NA & CA diploids
B genome
Clade 4Remaining SA diploidsA genome
ConclusionsConclusions1. Wild potato species are divided into three clades
well defined
2. The North and Central American B genome species are well distinguish from SA species
3. S. verrucosum cluster with all SA diploid species (A genome)
4. Hawkes’s series PIN and POL are monophyletic
5. S. brevicaule and S. sparsipilum appear to have hybrid origins
6. Species did not group based on Hawkes’s four superseries based on flower shape
abz PIU AR 2adr TUB PR 2
chm CON PR 2imt TUB PR 2
sph PIN PS 1blb BUL PS 1
cph PIN PS 1ehr1 PIN PS 1ehr2 PIN PS 1
jam PIN PS 1
pnt PIN PS 1trf PIN PS 1
clr BUL PS 1mrl MOR PS 1les POL PS 1pld POL PS 1ber TUB PR 2tar YNG AS 2ifd1 CUN PR 2blv MEG PR 2mga MEG PR 2chc YNG AS 2gnd TUB PR 2brc1 TUB PR 2ifd2 CUN PR 2spl1 TUB PR 2
buk TUB PR 2
lgl LIG PS 1
mcd TUB PR 2
rap1 MEG PR 2rap2 MEG PR 2vio CON PR 2
brc2 TUB PR 2lph TUB PR 2spl2 TUB PR 2ver TUB AR 2lycetb ETU 1pls ETU 1
92
77
61
92
59
100
100
50
92
98
57
52
56
92
88
6256
100
88
SpeciesSeries
Superseries
EBN
Piurana clade
NA & CA diploids B genome
Remaining SA diploids
Maximum Parsimony Maximum Parsimony diploidsdiploids
NA & CA diploidsB genome
Piurana cladeP, C, D genome
A genomeRemaining SA diploids
Clade 3Piurana clade
P, C, D genome
Clade 1+2NA & CAdiploids
B genome
Clade 4Remaining SA diploidsA genome
EBN
Hawkes’s four superseriesbased on flower shape
ConclusionsConclusions
1. Wild potato species are divided into three clades well defined
2. The North and Central American B genome species are well distinguish from SA species (B genome)
3. S. verrucosum cluster with all SA diploid species (A genome)
4. Hawkes’s series PIN and POL are monophyletic
5. S. brc and spl appear to have hybrid origins
6. Species did not group based on Hawkes’s four superseries based on flower shape
BayesianInference tree with the complete database
Diploid speciesPolyploid species ser. LongipedicellataPolyploid species ser. ConnicibaccataPolyploid species Acaulia GroupPolyploid species Iopetalum GroupPolyploid species ser. PiuranaPolyploid species ser. Tuberosa
1.00
0.98
0.98
0.97
1.00
0.78
0.59
0.991.000.740.77
0.78
10.69
0.71
0.95
0.960.98
0.981.00
1.000.85
1.001.00
1
0.830.96
110.94
0.52
1.000.96
1
0.98
0.99
0.96
0.91
0.760.69
0.98
1.000.99
0.97
0.93
0.751.00
0.740.68
1.001.00
1.00
0.79
1.0072
0.68
10.56
1
0.59
0.950.95
0.66
0.96
10.97
0.57
0.57
S. lycopersicumS. acaule 1
S. acaule 2S. demisum 1
S. demisum 2
S. raphanifolium 1S. raphanifolium 2
S. agrimonifolium 1S. colombianum 1
S longiconicum 1S. violaceimarmoratum
S. microdontumS. lignicauleS. berthaultii
S. tarijenseS. infundibuliforme 1
S. sparsipilum 1
S. bolivienseS. megistacrolobum
S. chacoenseS. gandarillasii
S. brevicaule 1S. infundibuliforme 2
S. brachycarpum 1S. verrucosum
S. hjertingii 1S. stoloniferum 1
S. demisum 3S. demisum 4S. fendleri 1S. schenkii 1S. demisum 5
S. demisum 6S. fendleri 2
S. schenkii 2S. oplosence A1S. oplosence A2
S. oplosence A3
S. oplosence A5S. oplosence A6S. oplosence B3S. oplosence B4
S. oplocense B1S. oplosence B2
S. sparsipilum 2
S. brevicaule 2
S. oplosence A4
S. leptophyes
S. agrimonifolium 2S. agrimonifolium 3S. agrimonifolium 4
S. brachycarpum 2S. colombianum 2S. colombianum 3
S longiconicum 2S. tuquerrense 1S. tuquerrense 2
S. andreanumS. paucijugum 1
S. paucijugum 2S. schenkii 5
S. schenkii 6S. albornozii
S. immite
S. paucijugum 4S. tuquerrense 3S. tuquerrense 4
S. paucijugum 3
S. stenophyllidium
S. cardiophyllum subsp.ehrenbergii 2S. cardiophyllum subsp.ehrenbergii 1
S. fendleri 3
S. stoloniferum 3
S. fendleri 4
S. stoloniferum 2S. hjertingii 2
S. stoloniferum 4
S. bulbocastanumS cardiophyllum
S.trifidum
S. jamesiiS. pinnatisectum
S. schenkii 3S. schenkii 4
S. clarumS. morelliformeS. lesteri
S. polyadenium
S. bukasovii
S. chomatophilum
S. palustreS. etuberosum
Clade 1+2Genome B
Clade 4Genome A
Clade 3Piurana CladeGenome C or D or P
BayesianInference tree for the complete database
Diploid speciesPolyploid species ser. LongipedicellataPolyploid species ser. ConnicibaccataPolyploid species Acaulia GroupPolyploid species Iopetalum GroupPolyploid species ser. PiuranaPolyploid species ser. Tuberosa
1.00
0.98
0.98
0.97
1.00
0.78
0.59
0.991.000.740.77
0.78
10.69
0.71
0.95
0.960.98
0.981.00
1.000.85
1.001.00
1
0.830.96
110.94
0.52
1.000.96
1
0.98
0.99
0.96
0.91
0.760.69
0.98
1.000.99
0.97
0.93
0.751.00
0.740.68
1.001.00
1.00
0.79
1.0072
0.68
10.56
1
0.59
0.950.95
0.66
0.96
10.97
0.57
0.57
S. lycopersicumS. acaule 1
S. acaule 2S. demisum 1
S. demisum 2
S. raphanifolium 1S. raphanifolium 2
S. agrimonifolium 1S. colombianum 1
S longiconicum 1S. violaceimarmoratum
S. microdontumS. lignicauleS. berthaultii
S. tarijenseS. infundibuliforme 1
S. sparsipilum 1
S. bolivienseS. megistacrolobum
S. chacoenseS. gandarillasii
S. brevicaule 1S. infundibuliforme 2
S. brachycarpum 1S. verrucosum
S. hjertingii 1S. stoloniferum 1
S. demisum 3S. demisum 4S. fendleri 1S. schenkii 1S. demisum 5
S. demisum 6S. fendleri 2
S. schenkii 2S. oplosence A1S. oplosence A2
S. oplosence A3
S. oplosence A5S. oplosence A6S. oplosence B3S. oplosence B4
S. oplocense B1S. oplosence B2
S. sparsipilum 2
S. brevicaule 2
S. oplosence A4
S. leptophyes
S. agrimonifolium 2S. agrimonifolium 3S. agrimonifolium 4
S. brachycarpum 2S. colombianum 2S. colombianum 3
S longiconicum 2S. tuquerrense 1S. tuquerrense 2
S. andreanumS. paucijugum 1
S. paucijugum 2S. schenkii 5
S. schenkii 6S. albornozii
S. immite
S. paucijugum 4S. tuquerrense 3S. tuquerrense 4
S. paucijugum 3
S. stenophyllidium
S. cardiophyllum subsp.ehrenbergii 2S. cardiophyllum subsp.ehrenbergii 1
S. fendleri 3
S. stoloniferum 3
S. fendleri 4
S. stoloniferum 2S. hjertingii 2
S. stoloniferum 4
S. bulbocastanumS cardiophyllum
S.trifidum
S. jamesiiS. pinnatisectum
S. schenkii 3S. schenkii 4
S. clarumS. morelliformeS. lesteri
S. polyadenium
S. bukasovii
S. chomatophilum
S. palustreS. etuberosum
Clade 1+2Genome B
Clade 4Genome A
Clade 3Piurana CladeGenome C or D or P
BayesianInference tree for the complete database
Diploid speciesPolyploid species ser. LongipedicellataPolyploid species ser. ConnicibaccataPolyploid species Acaulia GroupPolyploid species Iopetalum GroupPolyploid species ser. PiuranaPolyploid species ser. Tuberosa
1.00
0.98
0.98
0.97
1.00
0.78
0.59
0.991.000.740.77
0.78
10.69
0.71
0.95
0.960.98
0.981.00
1.000.85
1.001.00
1
0.830.96
110.94
0.52
1.000.96
1
0.98
0.99
0.96
0.91
0.760.69
0.98
1.000.99
0.97
0.93
0.751.00
0.740.68
1.001.00
1.00
0.79
1.0072
0.68
10.56
1
0.59
0.950.95
0.66
0.96
10.97
0.57
0.57
S. lycopersicumS. acaule 1
S. acaule 2S. demisum 1
S. demisum 2
S. raphanifolium 1S. raphanifolium 2
S. agrimonifolium 1S. colombianum 1
S longiconicum 1S. violaceimarmoratum
S. microdontumS. lignicauleS. berthaultii
S. tarijenseS. infundibuliforme 1
S. sparsipilum 1
S. bolivienseS. megistacrolobum
S. chacoenseS. gandarillasii
S. brevicaule 1S. infundibuliforme 2
S. brachycarpum 1S. verrucosum
S. hjertingii 1S. stoloniferum 1
S. demisum 3S. demisum 4S. fendleri 1S. schenkii 1S. demisum 5
S. demisum 6S. fendleri 2
S. schenkii 2S. oplosence A1S. oplosence A2
S. oplosence A3
S. oplosence A5S. oplosence A6S. oplosence B3S. oplosence B4
S. oplocense B1S. oplosence B2
S. sparsipilum 2
S. brevicaule 2
S. oplosence A4
S. leptophyes
S. agrimonifolium 2S. agrimonifolium 3S. agrimonifolium 4
S. brachycarpum 2S. colombianum 2S. colombianum 3
S longiconicum 2S. tuquerrense 1S. tuquerrense 2
S. andreanumS. paucijugum 1
S. paucijugum 2S. schenkii 5
S. schenkii 6S. albornozii
S. immite
S. paucijugum 4S. tuquerrense 3S. tuquerrense 4
S. paucijugum 3
S. stenophyllidium
S. cardiophyllum subsp.ehrenbergii 2S. cardiophyllum subsp.ehrenbergii 1
S. fendleri 3
S. stoloniferum 3
S. fendleri 4
S. stoloniferum 2S. hjertingii 2
S. stoloniferum 4
S. bulbocastanumS cardiophyllum
S.trifidum
S. jamesiiS. pinnatisectum
S. schenkii 3S. schenkii 4
S. clarumS. morelliformeS. lesteri
S. polyadenium
S. bukasovii
S. chomatophilum
S. palustreS. etuberosum
Clade 1+2Genome B
Clade 4Genome A
Clade 3Piurana CladeGenome C or D or P
BayesianInference tree for the complete database
Diploid speciesPolyploid species ser. LongipedicellataPolyploid species ser. ConnicibaccataPolyploid species Acaulia GroupPolyploid species Iopetalum GroupPolyploid species ser. PiuranaPolyploid species ser. Tuberosa
1.00
0.98
0.98
0.97
1.00
0.78
0.59
0.991.000.740.77
0.78
10.69
0.71
0.95
0.960.98
0.981.00
1.000.85
1.001.00
1
0.830.96
110.94
0.52
1.000.96
1
0.98
0.99
0.96
0.91
0.760.69
0.98
1.000.99
0.97
0.93
0.751.00
0.740.68
1.001.00
1.00
0.79
1.0072
0.68
10.56
1
0.59
0.950.95
0.66
0.96
10.97
0.57
0.57
S. lycopersicumS. acaule 1
S. acaule 2S. demisum 1
S. demisum 2
S. raphanifolium 1S. raphanifolium 2
S. agrimonifolium 1S. colombianum 1
S longiconicum 1S. violaceimarmoratum
S. microdontumS. lignicauleS. berthaultii
S. tarijenseS. infundibuliforme 1
S. sparsipilum 1
S. bolivienseS. megistacrolobum
S. chacoenseS. gandarillasii
S. brevicaule 1S. infundibuliforme 2
S. brachycarpum 1S. verrucosum
S. hjertingii 1S. stoloniferum 1
S. demisum 3S. demisum 4S. fendleri 1S. schenkii 1S. demisum 5
S. demisum 6S. fendleri 2
S. schenkii 2S. oplosence A1S. oplosence A2
S. oplosence A3
S. oplosence A5S. oplosence A6S. oplosence B3S. oplosence B4
S. oplocense B1S. oplosence B2
S. sparsipilum 2
S. brevicaule 2
S. oplosence A4
S. leptophyes
S. agrimonifolium 2S. agrimonifolium 3S. agrimonifolium 4
S. brachycarpum 2S. colombianum 2S. colombianum 3
S longiconicum 2S. tuquerrense 1S. tuquerrense 2
S. andreanumS. paucijugum 1
S. paucijugum 2S. schenkii 5
S. schenkii 6S. albornozii
S. immite
S. paucijugum 4S. tuquerrense 3S. tuquerrense 4
S. paucijugum 3
S. stenophyllidium
S. cardiophyllum subsp.ehrenbergii 2S. cardiophyllum subsp.ehrenbergii 1
S. fendleri 3
S. stoloniferum 3
S. fendleri 4
S. stoloniferum 2S. hjertingii 2
S. stoloniferum 4
S. bulbocastanumS cardiophyllum
S.trifidum
S. jamesiiS. pinnatisectum
S. schenkii 3S. schenkii 4
S. clarumS. morelliformeS. lesteri
S. polyadenium
S. bukasovii
S. chomatophilum
S. palustreS. etuberosum
Clade 1+2Genome B
Clade 4Genome A
Clade 3Piurana CladeGenome C or D or P
BayesianInference tree for the complete database
Diploid speciesPolyploid species ser. LongipedicellataPolyploid species ser. ConnicibaccataPolyploid species Acaulia GroupPolyploid species Iopetalum GroupPolyploid species ser. PiuranaPolyploid species ser. Tuberosa
1.00
0.98
0.98
0.97
1.00
0.78
0.59
0.991.000.740.77
0.78
10.69
0.71
0.95
0.960.98
0.981.00
1.000.85
1.001.00
1
0.830.96
110.94
0.52
1.000.96
1
0.98
0.99
0.96
0.91
0.760.69
0.98
1.000.99
0.97
0.93
0.751.00
0.740.68
1.001.00
1.00
0.79
1.0072
0.68
10.56
1
0.59
0.950.95
0.66
0.96
10.97
0.57
0.57
S. lycopersicumS. acaule 1
S. acaule 2S. demisum 1
S. demisum 2
S. raphanifolium 1S. raphanifolium 2
S. agrimonifolium 1S. colombianum 1
S longiconicum 1S. violaceimarmoratum
S. microdontumS. lignicauleS. berthaultii
S. tarijenseS. infundibuliforme 1
S. sparsipilum 1
S. bolivienseS. megistacrolobum
S. chacoenseS. gandarillasii
S. brevicaule 1S. infundibuliforme 2
S. brachycarpum 1S. verrucosum
S. hjertingii 1S. stoloniferum 1
S. demisum 3S. demisum 4S. fendleri 1S. schenkii 1S. demisum 5
S. demisum 6S. fendleri 2
S. schenkii 2S. oplosence A1S. oplosence A2
S. oplosence A3
S. oplosence A5S. oplosence A6S. oplosence B3S. oplosence B4
S. oplocense B1S. oplosence B2
S. sparsipilum 2
S. brevicaule 2
S. oplosence A4
S. leptophyes
S. agrimonifolium 2S. agrimonifolium 3S. agrimonifolium 4
S. brachycarpum 2S. colombianum 2S. colombianum 3
S longiconicum 2S. tuquerrense 1S. tuquerrense 2
S. andreanumS. paucijugum 1
S. paucijugum 2S. schenkii 5
S. schenkii 6S. albornozii
S. immite
S. paucijugum 4S. tuquerrense 3S. tuquerrense 4
S. paucijugum 3
S. stenophyllidium
S. cardiophyllum subsp.ehrenbergii 2S. cardiophyllum subsp.ehrenbergii 1
S. fendleri 3
S. stoloniferum 3
S. fendleri 4
S. stoloniferum 2S. hjertingii 2
S. stoloniferum 4
S. bulbocastanumS cardiophyllum
S.trifidum
S. jamesiiS. pinnatisectum
S. schenkii 3S. schenkii 4
S. clarumS. morelliformeS. lesteri
S. polyadenium
S. bukasovii
S. chomatophilum
S. palustreS. etuberosum
Clade 1+2Genome B
Clade 4Genome A
Clade 3Piurana CladeGenome C or D or P
BayesianInference tree for the complete database
Diploid speciesPolyploid species ser. LongipedicellataPolyploid species ser. ConnicibaccataPolyploid species Acaulia GroupPolyploid species Iopetalum GroupPolyploid species ser. PiuranaPolyploid species ser. Tuberosa
1.00
0.98
0.98
0.97
1.00
0.78
0.59
0.991.000.740.77
0.78
10.69
0.71
0.95
0.960.98
0.981.00
1.000.85
1.001.00
1
0.830.96
110.94
0.52
1.000.96
1
0.98
0.99
0.96
0.91
0.760.69
0.98
1.000.99
0.97
0.93
0.751.00
0.740.68
1.001.00
1.00
0.79
1.0072
0.68
10.56
1
0.59
0.950.95
0.66
0.96
10.97
0.57
0.57
S. lycopersicumS. acaule 1
S. acaule 2S. demisum 1
S. demisum 2
S. raphanifolium 1S. raphanifolium 2
S. agrimonifolium 1S. colombianum 1
S longiconicum 1S. violaceimarmoratum
S. microdontumS. lignicauleS. berthaultii
S. tarijenseS. infundibuliforme 1
S. sparsipilum 1
S. bolivienseS. megistacrolobum
S. chacoenseS. gandarillasii
S. brevicaule 1S. infundibuliforme 2
S. brachycarpum 1S. verrucosum
S. hjertingii 1S. stoloniferum 1
S. demisum 3S. demisum 4S. fendleri 1S. schenkii 1S. demisum 5
S. demisum 6S. fendleri 2
S. schenkii 2S. oplosence A1S. oplosence A2
S. oplosence A3
S. oplosence A5S. oplosence A6S. oplosence B3S. oplosence B4
S. oplocense B1S. oplosence B2
S. sparsipilum 2
S. brevicaule 2
S. oplosence A4
S. leptophyes
S. agrimonifolium 2S. agrimonifolium 3S. agrimonifolium 4
S. brachycarpum 2S. colombianum 2S. colombianum 3
S longiconicum 2S. tuquerrense 1S. tuquerrense 2
S. andreanumS. paucijugum 1
S. paucijugum 2S. schenkii 5
S. schenkii 6S. albornozii
S. immite
S. paucijugum 4S. tuquerrense 3S. tuquerrense 4
S. paucijugum 3
S. stenophyllidium
S. cardiophyllum subsp.ehrenbergii 2S. cardiophyllum subsp.ehrenbergii 1
S. fendleri 3
S. stoloniferum 3
S. fendleri 4
S. stoloniferum 2S. hjertingii 2
S. stoloniferum 4
S. bulbocastanumS cardiophyllum
S.trifidum
S. jamesiiS. pinnatisectum
S. schenkii 3S. schenkii 4
S. clarumS. morelliformeS. lesteri
S. polyadenium
S. bukasovii
S. chomatophilum
S. palustreS. etuberosum
Clade 1+2Genome B
Clade 4Genome A
Clade 3Piurana CladeGenome C or D or P
ConclusionsConclusions
Species in the polyploid series Conicibaccata, Longipedicellata, and the Iopetalum group are derived from allopolyploidization
S. verrucosum is the maternal genome donor for the polyploid species in ser. Longipedicellata, in the Iopetalum group and of S. demissum
The North and Central American B genome species are well distinguish from species of South America
NIA is useful in wild potato phylogenetic analysis
Do potatoes and tomatoes have a Do potatoes and tomatoes have a single evolutionary history? What single evolutionary history? What proportion of the genome supports proportion of the genome supports this history?this history?
Identify COS appropriate for wild potato and wild tomato phylogenetic studies
Generate a molecular data set to reconstruct the phylogeny of diploid wild tomato and wild potato
Reconstruct the evolutionary history of representatives of 14 wild potato and all 13 wild tomato species
ObjectivesObjectives
Materials and methodsMaterials and methods
IngroupIngroup: 13 species of wild tomatoes (section Lycopersicon)14 species of wild potatoes (section Petota)
OutgroupOutgroup:2 species of Solanum section Juglandifolium2 species of Solanum section Lycopersicoides2 species Solanum section Etuberosum1 species Solanum section Dulcamara1 species of Datura
COS selection criteriaCOS selection criteria
Chromosome coverage
Single-copy at least in potato and tomato
Sequence amplification length
Intron and exon content
MethodsMethods
COS fragments were amplified using PCR and direct cycle sequencing and cloning when necessary
COS sequences were edited using STADEN software, and were aligned with CLUSTALX and further manually aligned in MacClade
The sequence analyses were performed using Maximum Parsimony, Maximum Likelihood, Bayesian, and Concordance Analyses
Analyzing data from different sourcesAnalyzing data from different sources
There are 3 different traditional approaches:
1. The total evidence approach
2. The consensus approach
3. The prior agreement approach
Potato Potato datasetdataset
Individual COS
At1g50020 1,100 82.8 IAt1g30580 1000 92.3 IIAt3g03100 850 82.0 IIIAt2g24270 800 86.0 VIIAt1g14000 750 80.7 VAt1g77470 700 83.1 VI
ChromosomeIntron%COS marker Total length
bp
COS selection criteriaCOS selection criteria
Summary of potato phylogenetic Summary of potato phylogenetic analyses by MPanalyses by MP
Locus Exon bp
Exon PI
Intron bp
Intron PI
DNA bp
DNA PI Gap Gap
PITotal
CharactersTotal
PINo.
trees
COS 003 246 1 207 12 453 13 1 0 454 13 2
COS 005 146 4 850 47 996 51 30 13 1026 64 3
COS 008 200 3 102 11 302 14 13 6 315 20 2
COS 009 53 1 1781 44 1834 45 23 9 1857 54 5
COS 07B 16 0 806 40 822 40 9 7 831 47 4
COS 09B 24 0 1157 54 1181 54 14 5 1195 59 5
COS 10B 84 1 576 33 660 34 8 4 668 38 18
COS 011 250 8 402 40 652 48 13 7 665 55 2
COS 01C 171 5 522 40 693 45 12 5 705 50 1
COS 03C 79 2 706 18 785 20 8 1 793 21 1
COS 05C 46 0 1224 52 1270 52 21 9 1291 61 3
COS 02C 217 5 864 28 1081 33 5 4 1086 37 2
Number of total characters and number of Parsimony Informative characters
Summary of potato phylogenetic Summary of potato phylogenetic analyses by MPanalyses by MP
Locus Potato Mexican Clade1+2 blb-bst Piurana
Clade 3Clade
4aClade
4b Clade 4Clade3
with Clade4
Clade2 with
Clade4
Clade3 with
Clade2COS 003 61 56 87 100 no no 68 65 no no
COS 005 99 100 no <51 yes 100 no unres. no no
COS 008 95 66 no 55 yes 89 100 59 no no
COS 009 100 unres. 96 93 yes 98 95 81 no no
COS 07B 100 100 54 100 no no 100 no 90 no
COS 09B 100 60 88 100 yes 64 95 no no 60
COS 10B 100 unres. unres. unres. yes 97 100 no no 100
COS 011 100 100 76 62 unres. unres. 95 100 no no
COS 01C 100 98 76 100 yes 99 60 no <51 no
COS 03C 100 55 53 93 no no 93 no 69 no
COS 05C 100 no no no no no 95 no no no
COS 02C 100 unres. <51 no no <51 no no no no
Bootstrap clade values (%)
Clade 1&2
blb
bst
trf
pld
abz
adr
rap
brc
ver
Clade 3
Clade 4
Potato Potato datasetdataset
Combined COS
Results of ILD test in potatoesResults of ILD test in potatoes
At α= 0.0008 (0.05/66)
COS 3 5 8 9 7bR 09b 10b 11 1c 3c 5c 2c3 _ 0.22 0.12 1 0.48 0.68 0.11 1 0.54 0.72 0.047 0.005
5 0.27 _ 0.08 0.06 0.0001 0.06 0.0016 0.27 0.11 0.14 0.0001 0.09
8 0.17 0.06 _ 0.03 0.07 0.08 0.04 0.15 0.07 0.10 0.0046 0.0037
9 1.00 0.04 0.02 _ 0.0002 0.70 0.15 1.00 0.0065 0.15 0.02 0.03
7bR 0.39 0.0001 0.05 0.0003 _ 0.56 0.01 0.07 0.02 1.00 0.0001 0.0001
09b 0.66 0.04 0.10 0.70 0.48 _ 1.00 0.07 1.00 0.82 0.02 0.0061
10b 0.11 0.0017 0.10 0.12 0.0022 1.00 _ 0.0006 0.26 0.28 0.0009 0.0014
11 1.00 0.21 0.15 1.00 0.03 0.04 0.0006 _ 0.0064 0.63 0.0002 0.03
1c 0.60 0.14 0.14 0.02 0.02 1.00 0.36 0.0063 _ 0.48 0.0001 0.0002
3c 0.73 0.18 0.09 0.19 1.00 0.85 0.42 0.75 0.41 _ 0.19 0.02
5c 0.0421 0.0001 0.0043 0.0074 0.0004 0.0102 0.0018 0.0001 0.0001 0.1663 _ 0.0037
2c 0.0032 0.13 0.0047 0.0224 0.0001 0.0062 0.0014 0.0057 0.0001 0.0149 0.0028 _
Phylogenetic trees based on a combined Phylogenetic trees based on a combined analysisanalysis
12 COS 6 COS 3 COS
2 COS1 COS
Concordance analysesConcordance analyses
Tomato datasetTomato dataset
Combined COS
Phylogenetic Phylogenetic tree tree based on a combined analysisbased on a combined analysis
Concordance analysesConcordance analyses
ConclusionsConclusions
Intron contents more than 60% are the best to investigate relationships among closely related species
The total evidence approach produce a completely resolved potato phylogeny
In potato, three COS are enough to obtain a resolved phylogeny
The “prior agreement” approach and “concordance analysis” identified two histories in potato, one of them is the same as that obtained by “total evidence”
ConclusionsConclusions
COS data support the segregation of S. peruvianuminto at least three species
The total evidence approach completely resolved 19 of the 21 nodes in tomato
COS are useful for potato and tomato phylogenetic studies
Concordance analyses is useful to summarize discordance among genes
AcknowledgementsAcknowledgements
Dr. David Spooner
People in Spooner’s lab
Dr. David Baum and Cecile Ane
Alberto Salas
University of Wisconsin Plant Breeding and Plant Genetics Program
US Potato and Tomato genebanks