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Gene discovery using massively parallel pyrosequencing to develop ESTs forthe flesh fly Sarcophaga crassipalpis
BMC Genomics 2009, 10:234 doi:10.1186/1471-2164-10-234
Daniel A Hahn ([email protected])Gregory J Ragland ([email protected])
D. DeWayne Shoemaker ([email protected])David L Denlinger ([email protected])
ISSN 1471-2164
Article type Research article
Submission date 13 February 2009
Acceptance date 19 May 2009
Publication date 19 May 2009
Article URL http://www.biomedcentral.com/1471-2164/10/234
Like all articles in BMC journals, this peer-reviewed article was published immediately uponacceptance. It can be downloaded, printed and distributed freely for any purposes (see copyright
notice below).
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© 2009 Hahn et al. , licensee BioMed Central Ltd.This is an open access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
" /"3"/"
Gene discovery using massively parallel
pyrosequencing to develop ESTs for the flesh fly
Sarcophaga crassipalpis
"
Fcpkgn"C0"Jcjp"3¸."Itgiqt{"L0"Tcincpf3."F0"FgYc{pg"Ujqgocmgt4."cpf"Fcxkf"N0"
Fgpnkpigt5"
"
30"Fgrctvogpv"qh"Gpvqoqnqi{"cpf"Pgocvqnqi{."Vjg"Wpkxgtukv{"qh"Hnqtkfc."R0Q0"Dqz"
332842."Ickpguxknng."Hn."54833/2842."WUC0""
"
40"WUFC"CTU."Egpvgt"hqt"Ogfkecn."Citkewnvwtcn."cpf"Xgvgtkpct{"Gpvqoqnqi{."3822"
UY"45tf"Ftkxg."Ickpguxknng."HN"5482:."WUC0"
"
50"Fgrctvogpv"qh"Gpvqoqnqi{."Qjkq"Uvcvg"Wpkxgtukv{."622"Ctqpqhh"Ncdqtcvqt{."53:"
Yguv"34vj"Cxgpwg."Eqnwodwu."QJ"65432."WUC0"
"
¸Eqttgurqpfkpi"cwvjqt"
"
Gockn"cfftguugu<"
FCJ<"fcjcjpBwhn0gfw"
ILT<"itcincpfBwhn0gfw"
FFU<"Fgyc{pg0UjqgocmgtBctu0wufc0iqx""
FNF<"fgpnkpigt03Bquw0gfw
" /"4"/"
Cduvtcev""
Background
Hnguj"hnkgu"kp"vjg"igpwu"Sarcophaga ctg"korqtvcpv"oqfgnu"hqt"kpxguvkicvkpi"
gpfqetkpqnqi{."fkcrcwug."eqnf"jctfkpguu."tgrtqfwevkqp."cpf"koowpkv{0"Fgurkvg"vjg"
rtqokpgpeg"qh"Sarcophaga"hnguj"hnkgu"cu"oqfgnu"hqt"kpugev"rj{ukqnqi{"cpf"
dkqejgokuvt{."cpf"kp"hqtgpuke"uvwfkgu."nkvvng"igpqoke"qt"vtcpuetkrvqoke"fcvc"ctg"
cxckncdng"hqt"ogodgtu"qh"vjku"igpwu0"Yg"wugf"ocuukxgn{"rctcnngn"r{tqugswgpekpi"qp"
vjg"Tqejg"676/HNZ"rncvhqto"vq"rtqfweg"c"uwduvcpvkcn"GUV"fcvcugv"hqt"vjg"hnguj"hn{"
Sarcophaga crassipalpis0"Vq"oczkok¦g"ugswgpeg"fkxgtukv{."yg"rqqngf"TPC"gzvtcevgf"
htqo"yjqng"dqfkgu"qh"cnn"nkhg"uvcigu"cpf"pqtocnk¦gf"vjg"eFPC"rqqn"chvgt"tgxgtug"
vtcpuetkrvkqp0"
Results
Yg"qdvckpgf"429.332"GUVu"ykvj"cp"cxgtcig"tgcf"ngpivj"qh"463"dr0"Vjgug"tgcfu"
cuugodngf"kpvq"42.;;7"eqpvkiu"cpf"53.278"ukpingvqpu0"Wukpi"DNCUV"ugctejgu"qh"vjg"
PT"cpf"PV"fcvcdcugu"yg"ygtg"cdng"vq"kfgpvkh{"33.979"wpkswg"igpg"gngogpvu"
*G>202223+"tgrtgugpvkpi"crrtqzkocvgn{";.222"kpfgrgpfgpv"vtcpuetkrvu0""Eqorctkuqp"qh"
vjg"fkuvtkdwvkqp"qh"S. crassipalpis wpkigpgu"coqpi"IQ"Dkqnqikecn"Rtqeguu"hwpevkqpcn"
itqwru"ykvj"vjcv"qh"vjg"Drosophila melanogaster"vtcpuetkrvqog"uwiiguvu"vjcv"qwt"
GUVu"ctg"dtqcfn{"tgrtgugpvcvkxg"qh"vjg"hnguj"hn{"vtcpuetkrvqog0"Kpugtvkqp"cpf"fgngvkqp"
gttqtu"kp"676"ugswgpekpi"rtgugpv"c"ugtkqwu"jwtfng"vq"eqorctcvkxg"vtcpuetkrvqog"
cpcn{uku0"Ckfgf"d{"c"pgy"crrtqcej"vq"eqttgevkpi"hqt"vjgug"gttqtu."yg"rgthqtogf"c"
eqorctcvkxg"cpcn{uku"qh"igpgvke"fkxgtigpeg"cetquu"IQ"ecvgiqtkgu"coqpi"S.
crassipalpis."D. melanogaster."cpf"Anopheles gambiae0"Vjg"tguwnvu"uwiiguv"vjcv"pqp/
u{pqp{oqwu"uwduvkvwvkqpu"qeewt"cv"ukoknct"tcvgu"cetquu"ecvgiqtkgu."cnvjqwij"igpgu"
" /"5"/"
tgncvgf"vq"tgurqpug"vq"uvkownk"oc{"gxqnxg"unkijvn{"hcuvgt0"Kp"cffkvkqp."yg"kfgpvkhkgf"
qxgt"722"rqvgpvkcn"oketqucvgnnkvg"nqek"cpf"oqtg"vjcp"34.222"UPRu"coqpi"qwt"GUVu0""
Conclusions
Qwt"fcvc"rtqxkfgu"vjg"hktuv"nctig/uecng"GUV/rtqlgev"hqt"hnguj"hnkgu."c"owej/pggfgf"
tguqwteg"hqt"gzrnqtkpi"vjku"oqfgn"urgekgu0"Kp"cffkvkqp."yg"kfgpvkhkgf"c"nctig"pwodgt"qh"
rqvgpvkcn"oketqucvgnnkvg"cpf"UPR"octmgtu"vjcv"eqwnf"dg"wugf"kp"rqrwncvkqp"cpf"
u{uvgocvke"uvwfkgu"qh"S. crassipalpis"cpf"qvjgt"hnguj"hnkgu0"
Background Hnguj"hnkgu"kp"vjg"igpwu"Sarcophaga"*Fkrvgtc<"Ucteqrjcikfcg+"jcxg"nqpi"dggp"
korqtvcpv"oqfgnu"kp"kpugev"rj{ukqnqi{"cpf"dkqejgokuvt{."rctvkewnctn{"ykvj"tgurgev"vq"
vjg"uvwf{"qh"gpfqetkpqnqi{"]3."4_."fkcrcwug"]5."6_."eqnf"jctfkpguu"]7."8_."tgrtqfwevkqp"
]9.:_."cpf"koowpkv{"];_0"Nctxcg"qh"Sarcophaga"hnkgu"v{rkecnn{"hggf"qp"ecttkqp"qt"hgegu."
rnc{kpi"cp"korqtvcpv"geqnqikecn"tqng"kp"fgeqorqukvkqp"cpf"hgcvwtg"rtqokpgpvn{"kp"
jwocp"hqtgpuke"uvwfkgu"]32."33_0"Sarcophaga urgekgu"ecp"ecwug"o{kcuku"kp"jwocpu"cpf"
nkxguvqem"cpf"vjgtghqtg"ctg"qh"korqtvcpeg"vq"ogfkecn"cpf"xgvgtkpct{"gpvqoqnqi{"]34_0"
Kp"cffkvkqp."Sarcophaga"hnkgu"ctg"pcvwtcn"jquvu"qh"vjg"rctcukvke"ycur"Nasonia"
vitripennis"cpf"ctg"qhvgp"wugf"kp"ncdqtcvqt{"uvwfkgu"cpf"hqt"ycur"ewnvwtg0"Nasonia"
vitripennis"rctcukvk¦gu"rwrcg"qh"ocp{"urgekgu"qh"jkijgt"hnkgu."kpenwfkpi"jqwug"hnkgu"cpf"
qvjgt"hknvj"hnkgu."ocmkpi"vjgo"geqpqokecnn{"korqtvcpv"hqt"dkqnqikecn"eqpvtqn"]35_0"
Hwtvjgtoqtg."N. vitripennis"ku"c"rqrwnct"oqfgn"hqt"wpfgtuvcpfkpi"jquv/rctcukvg"
tgncvkqpujkru."dgjcxkqtcn"geqnqi{."fgxgnqrogpv."cpf"gxqnwvkqp."vjwu"qhhgtkpi"cp"
korgvwu"hqt"ugswgpekpi"vjg"igpqog"qh"vjku"ycur"cpf"vyq"eqpigpgtu"N. giraulti cpf"N.
longicornis"]36."37."38_0"
"
" /"6"/"
Fgurkvg"kvu"rtqokpgpeg"cu"c"oqfgn"kp"ugxgtcn"uwd/fkuekrnkpgu."vjgtg"ku"ewttgpvn{"pq"
igpqog"rtqlgev"hqt"cp{"Sarcophaga"urgekgu."cpf"igpqoke"tguqwtegu"ctg"nkokvgf0"
Ugxgtcn"ncdqtcvqtkgu"jcxg"rgthqtogf"nqy/vjtqwijrwv"GUV"rtqlgevu"uwej"cu"fkhhgtgpvkcn"
fkurnc{"]39_"cpf"uwdvtcevkxg"j{dtkfk¦cvkqp"]3:_."dwv"vjg"ockpuvc{"qh"hwpevkqpcn"
igpqoke"tgugctej"qp"urgekgu"kp"vjku"igpwu"jcu"hqewugf"qp"jqoqnqi{/dcugf."igpg/d{/
igpg"enqpkpi0"Vjg"igpqoke"tguqwtegu"fgxgnqrgf"hqt"c"jcpfhwn"qh"kpugevu"kpenwfkpi"
Drosophila."jqpg{"dggu."oquswkvqgu."cpf"Bombyx mori"jcu"rtqorvgf"owej"yqtm"qp"
vjgug"urgekgu."rwujkpi"qwt"wpfgtuvcpfkpi"qh"rtqeguugu"kp"vjgug"hgy"oqfgnu"hct"cjgcf"
qh"qvjgt"kpugevu0"Fgxgnqrogpv"qh"uwej"tguqwtegu"hqt"Sarcophaga yqwnf"itgcvn{"dgpghkv"
vjg"uvwf{"qh"hnguj"hnkgu"cu"jcu"dggp"fqpg"ykvj"qvjgt"pqp/vtcfkvkqpcn"oqfgn"kpugevu"uwej"
cu"Polistes"ycuru"]3;_."Incpxknng"htkvknnct{"dwvvgthnkgu"]42_."cpf"vqdceeq"jqtpyqto"
oqvju"]43_0""
"
Yg"fguetkdg"cp"GUV"eqnngevkqp"fgxgnqrgf"cu"c"igpqoke"tguqwteg"hqt"vjg"hnguj"hn{"
Sarcophaga crassipalpis."c"oqfgn"urgekgu"hqt"uvwf{kpi"pwogtqwu"rtqeguugu"kpenwfkpi"
fkcrcwug."tgrtqfwevkqp."gpfqetkpqnqi{."uvtguu"vqngtcpeg."cpf"koowpkv{0"Vq"hceknkvcvg"
kfgpvkh{kpi"ugvu"qh"igpgu"kpxqnxgf"kp""c"dtqcf"tcpig"qh"rtqeguugu"yg"fgxgnqrgf"qwt"
GUV"ugv"htqo"c"pqtocnk¦gf"yjqng/dqf{"nkdtct{"qh"pwogtqwu"S. crassipalpis"nkhg"uvcigu"
kpenwfkpi"fkcrcwug/fguvkpgf"cpf"pqp"fkcrcwug/fguvkpgf"nctxcg."fkcrcwukpi"cpf"pqp/
fkcrcwukpi"rwrcg."cpf"tgrtqfwevkxg"cpf"pqp/tgrtqfwevkxg"cfwnvu"qh"dqvj"ugzgu"*Vcdng"
3+0"Yg"eqphkfgpvn{"kfgpvkhkgf"crrtqzkocvgn{";.222"fkhhgtgpv"vtcpuetkrvu"kpxqnxgf"kp"
pwogtqwu"dkqnqikecn"rtqeguugu0"Ykvjkp"vjgug"vtcpuetkrvu."yg"kfgpvkhkgf"oqtg"vjcp"722"
rqvgpvkcn"oketqucvgnnkvg"nqek"cpf"qxgt"34.222"UPRu"vjcv"ecp"dg"wugf"hqt"hwvwtg"
rqrwncvkqp"uvwfkgu0"Eqorctkuqp"qh"vjg"fkuvtkdwvkqp"qh"S. crassipalpis wpkigpgu"coqpi"
IQ"Dkqnqikecn"Rtqeguu"hwpevkqpcn"itqwru"ykvj"vjcv"qh"vjg"Drosophila melanogaster"
" /"7"/"
vtcpuetkrvqog"uwiiguvu"vjcv"qwt"GUVu"ctg"dtqcfn{"tgrtgugpvcvkxg"qh"vjg"hnguj"hn{"
vtcpuetkrvqog0"Eqodkpkpi"qwt"fcvc"ykvj"hwnn"vtcpuetkrvqogu"htqo"D. melanogaster"cpf"
A. gambiae."yg"wugf"c"vjtgg/urgekgu"eqorctkuqp"vq"vguv"yjgvjgt"eqfkpi"ugswgpegu"ctg"
gxqnxkpi"cv"ukoknct"tcvgu"cetquu"vjgug"IQ"ecvgiqtkgu0""
Results and Discussion Qwt"pqtocnk¦gf"S. crassipaplis"nkdtct{"ycu"hktuv"uwdlgevgf"vq"c"qpg/swctvgt"rncvg"
vkvtcvkqp"twp"qp"vjg"676"IU/HNZ"ocejkpg"vjcv"{kgnfgf"3.734.455"vqvcn"dcugu"htqo"
7.949"ugswgpeg"tgcfu"vjcv"cuugodngf"kpvq"776"eqpvkiu"cpf"5.3:4"ukpingvqpu0"Qh"vjgug"
eqpvkiu"cpf"ukpingvqpu"3.687"ygtg"kfgpvkhkcdng"cv"cp"G>20223wukpi"DNCUVZ"cickpuv"
vjg"PEDK"pqp/tgfwpfcpv"rtqvgkp"fcvcdcug"*PT+0""
"
Yg"hqnnqygf"vjku"qpg/swctvgt"rncvg"rtgnkokpct{"twp"ykvj"c"hwnn/rncvg"rtqfwevkqp"twp"
vjcv"{kgnfgf"94.:38.:33"vqvcn"dcugu"htqo"4:3.759"ugswgpeg"tgcfu"cpf"eqodkpgf"vjg"
tguwnvu"qh"dqvj"twpu"kpvq"c"ukping"fcvcugv"yjkej"ycu"cuugodngf"de novo0"Hkngu"
eqpvckpkpi"qwt"tguwnvu"ctg"cxckncdng"htqo"vjg"Pcvkqpcn"Egpvgt"hqt"Dkqvgejpqnqi{"
Kphqtocvkqp"Ujqtv"Tgcf"Ctejkxg"*ceeguukqp"pwodgtu"UTT227287"cpf"UTT228::6+0"
Chvgt"hknvgtkpi"cfcrvqtu"cpf"nqy/swcnkv{"ugswgpegu"]44_"yg"ygtg"nghv"ykvj"96.54;.266"
vqvcn"dcugu"htqo"429.332"ugswgpeg"tgcfu"ykvj"cp"cxgtcig"ngpivj"qh"463"dcugu0"Vjgug"
fcvc"ygtg"cuugodngf"kpvq"42.;;7"eqpvkiu"ykvj"c"ogcp"ngpivj"qh"554"dr"cpf"c"tcpig"qh"
52"vq"4.;7:"dr"cu"ygnn"cu"cp"cffkvkqpcn"53.278"ukpingvqpu"hqt"c"vqvcn"qh"74.273"jkij/
swcnkv{"ugswgpegu0"C"hkng"qh"cnn"cuugodngf"eqpvkiu"cpf"ukpingvqpu"ku"cxckncdng"htqo"vjg"
cwvjqtu"wrqp"tgswguv0"
"
Uwdlgevkpi"vjgug"ugswgpegu"vq"c"DNCUVZ"ugctej"cickpuv"vjg"PEDK/PT"rtqvgkp"
fcvcdcug"{kgnfgf"3;.82;"ygnn/kfgpvkhkgf"ugswgpegu"ykvj"cv"ngcuv"qpg"jkv"G>202223"
" /"8"/"
*59'+."37.463"rqqtn{/kfgpvkhkgf"ugswgpegu"ykvj"jkvu"dgvyggp"G?202223"cpf"G?32."
cpf"54.655"ykvj"pq"wughwn"jkvu"*G@32+0"Uwdlgevkpi"vjgug"ugswgpegu"vq"c"DNCUVP"
ugctej"cickpuv"vjg"PEDK/PV"ugswgpeg"fcvcdcug"{kgnfgf"8.543"ygnn/kfgpvkhkgf"
ugswgpegu"ykvj"c"jkv"G>202223"*34'+."66.48:"rqqtn{/kfgpvkhkgf"ugswgpegu"ykvj"jkvu"
dgvyggp"G?202223"cpf"G?32."cpf"3.675"ykvj"pq"wughwn"jkvu"*G@32+0""Ygnn/kfgpvkhkgf"
ugswgpegu"*G>20223+"htqo"dqvj"ugctejgu"ygtg"eqodkpgf"cpf"eqorctgf"ykvj"gcej"
qvjgt"wukpi"DNCUVP."tgfwekpi"vjg"pwodgt"qh"ygnn/kfgpvkhkgf"ugswgpegu"kpvq"33.979"
wpkswg"igpg"gngogpvu"qt"wpkigpgu0"Vq"guvkocvg"jqy"ocp{"qh"vjgug"wpkigpgu"
tgrtgugpvgf"wpkswg"vtcpuetkrvu"cu"qrrqugf"vq"fkhhgtgpv"pqp/eqpvkiwqwu"htciogpvu"qh"
vjg"ucog"vtcpuetkrv"yg"rtqfwegf"c"hkng"qh"vjg"vqr"72"jkvu"hqt"gcej"wpkigpg"cpf"wugf"
vjgug"nkuvu"vq"eqorctg"ceeguukqp"pwodgtu"vjcv"qxgtncrrgf"coqpi"fkhhgtgpv"wpkigpgu"
ykvj"cp"G>20223"wukpi"c"ewuvqo"Gzegn"ocetq"hqnnqygf"d{"ocpwcn"gfkvkpi0"Vjku"
rtqegfwtg"kfgpvkhkgf";.539"ugswgpegu"vjcv"ctg"nkmgn{"kpfgrgpfgpv"vtcpuetkrvu"*Vcdng"4+0""
"
Yjkng"vjg"\;.222"kfgpvkhkgf"vtcpuetkrvu"ctg"egtvckpn{"nguu"vjcp"vjg"gpvktg"vtcpuetkrvqog."
yg"ceswktgf"ocp{"ugswgpegu"qh"kpvgtguv0"Dgecwug"qwt"itqwru."coqpi"qvjgtu."ctg"
rctvkewnctn{"kpvgtguvgf"kp"fkcrcwug"cpf"uvtguu"dkqnqi{."yg"eqorkngf"c"dtkgh"nkuv"qh"igpgu"
yg"gzrgev"rnc{"c"tqng"kp"fkcrcwug"*Cffkvkqpcn"hkng"3+0"Engctn{."gxgp"vjku"cddtgxkcvgf"
nkuv"yknn"rtqxkfg"hqffgt"hqt"{gctu"qh"hwpevkqpcn"igpqokeu"yqtm"qp"vjgug"ecpfkfcvg"
igpgu"hqt"qwt"itqwr"cpf"qvjgtu0""
"
Qwt"oclqt"iqcn"ycu"vq"igpgtcvg"c"uwduvcpvkcn"tgrtgugpvcvkxg"ucorng"qh"vjg"
vtcpuetkrvqog0"Vq"cuuguu"yjgvjgt"qwt"GUV"rtqlgev"ycu"dtqcfn{"tgrtgugpvcvkxg"qh"qwt"
gzrgevcvkqpu"hqt"vjg"Sarcophaga"vtcpuetkrvqog."yg"wugf"vjg"cppqvcvkqpu"hqt"gcej"qh"
qwt"eqpvkiu"cpf"ukpingvqpu"vq"cuukip"kv"vq"qpg"qh"36"oclqt"IQ"ecvgiqtkgu"hqt"Dkqnqikecn"
" /"9"/"
Rtqeguu."rtqfwekpi"cuukipogpvu"hqt";.68:"qwv"qh"c"vqvcn"74.273"vtcpuetkrvu"*Hki0"3+0"Yg"
cnuq"eqorctgf"vjg"fkuvtkdwvkqp"qh"qwt"ugswgpegu"coqpi"vjgug"36"IQ"hwpevkqpcn"itqwru"
ykvjkp"Dkqnqikecn"Rtqeguu"vq"vjg"fkuvtkdwvkqp"qh"vjg"37.3:5"rtgfkevgf"igpgu"hqt"
Drosophila melanogaster, 7.275"qh"yjkej"ctg"rncegf"kpvq"vjqug"ucog"hwpevkqpcn"
itqwru"*Hki0"4."Hn{Dcug."xgtukqp"422:a29+0"Cnvjqwij"vjgtg"ctg"pqvkegcdng"fkhhgtgpegu"
dgvyggp"vjg"Drosophila"cpf"Sarcophaga"fcvc"kp"vjg"rgtegpvcig"qh"vtcpuetkrvu"ykvjkp"c"
hgy"uwd/ecvgiqtkgu."eqpeqtfcpeg"kp"vjg"qxgtcnn"fkuvtkdwvkqpu"uwiiguvu"vjcv"qwt"nkdtct{"
ucorngf"ykfgn{"cetquu"uwd/ecvgiqtkgu"cpf"rtqxkfgu"c"iqqf"tgrtgugpvcvkqp"qh"vjg"S.
crassipalpis"vtcpuetkrvqog0"Cu"ugswgpekpi"vgejpqnqikgu"kortqxg."hwtvjgt"gzrnqtcvkqp"
qh"vjg"S. crassipalpis"vtcpuetkrvqog"yknn"nkmgn{"kpenwfg"ugswgpekpi"qh"pgy"nkdtctkgu"
ocfg"htqo"mg{"vkuuwgu"fwtkpi"hqecn"nkhg"uvcigu"vjcv"oc{"pqv"jcxg"jkij"tgrtgugpvcvkqp"
kp"vqvcn"dqf{"gzvtcevu."uwej"cu"dtckpu"htqo"fkcrcwukpi"rwrcg"qt"qxctkgu"htqo"rtg"cpf"
rquv/tgrtqfwevkxg"hgocngu."cu"ygnn"cu"fggrgt"ugswgpekpi"vq"dqvj"tgxgcn"tctg"vtcpuetkrvu"
cpf"rtqxkfg"vjg"cffkvkqpcn"kphqtocvkqp"pgeguuct{"vq"kfgpvkh{"oqtg"qh"vjg"wpcppqvcvgf"
vtcpuetkrv"htciogpvu"kp"qwt"ewttgpv"yjqng/dqf{"nkdtct{0""
"
Dgecwug"qwt"GUV"fcvcdcug"crrgctu"dtqcfn{"tgrtgugpvcvkxg"qh"vjg"S. crassipalpis"
vtcpuetkrvqog."yg"eqorctgf"kv"vq"vjg"cxckncdng"vtcpuetkrvqogu"qh"D. melanogaster"cpf"
A. gambiae"vq"cuuguu"yjgvjgt"eqfkpi"ugswgpegu"ygtg"gxqnxkpi"cv"gswkxcngpv"tcvgu"
cetquu"IQ"ecvgiqtkgu0"Pqp/u{pqp{oqwu"tcvgu"qh"uwduvkvwvkqp"*fP+"ygtg"tqwijn{"
icooc/fkuvtkdwvgf"cetquu"igpgu"ykvjkp"gcej"IQ"ecvgiqt{"hqt"cnn"vjtgg"rcktykug"
eqorctkuqpu0"Ogfkcp"xcnwgu"cetquu"cnn"IQ"ecvgiqtkgu"ygtg"igpgtcnn{"nqyguv"kp"
eqorctkuqpu"qh"S. crassipalpis"vq"D. melanogaster"*Hki0"5+."tghngevkpi"vjgkt"enqug"
tgncvkqpujkr"ykvjkp"E{enqttcrjc"tgncvkxg"vq"vjg"tgncvkxgn{"fggr"rj{nqigpgvke"urnkv"
dgvyggp"E{enqttcrjc"cpf"Ewnkeqoqtrjc0"Vjg";7'"eqphkfgpeg"kpvgtxcnu"qh"vjg"ogfkcp"
" /":"/"
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rcktykug"eqorctkuqpu"*ukoknctn{"ujcfgf"dctu"kp"Hki0"5+."uwiiguvkpi"nctign{"
jqoqigpgqwu"tcvgu"qh"cokpq"cekf"uwduvkvwvkqp0"Jqygxgt."vjg"ogfkcp"fP"hqt"Tgurqpug"
vq"uvkownk"jcf"vjg"jkijguv"*D. melanogaster"vq"A. gambiae="S. crassipalpis"vq"A.
gambiae+"qt"ugeqpf"jkijguv"*S. crassipalpis"vq"D. melanogaster+"xcnwg"kp"vjg"tcpm"
qtfgtkpi"coqpi"IQ"ecvgiqtkgu"cetquu"cnn"vjtgg"rcktykug"eqorctkuqpu0"Vjku"
eqpukuvgpvn{"jkij"fP"cetquu"ownvkrng"eqorctkuqpu"qh"urgekgu"fkxgtigf"qp"vjg"qtfgt"qh"
422"oknnkqp"{gctu"ciq"]45_"uwiiguvu"vjcv"igpgu"tgiwncvkpi"tgurqpugu"vq"uvkownk"oc{"
gxqnxg"tgncvkxgn{"tcrkfn{"cetquu"fkrvgtcp"vczc0"Vjku"fkxgtigpeg"ogujgu"ygnn"ykvj"
qdxkqwu"fkhhgtgpegu"kp"ugpuqt{"uvtcvgikgu"dgvyggp"vjgug"vjtgg"urgekgu"vjcv"owuv"
kfgpvkh{"cpf"nqecvg"xgt{"fkhhgtgpv"pwvtkvkqpcn"tguqwtegu"*tqvvkpi"hnguj."nkxg"cpkocnu"hqt"
dnqqf"ogcnu."cpf"tqvvkpi"htwkv+0"Kp"cffkvkqp."tgurqpugu"vq"rjgtqoqpcn"cpf"dgjcxkqwtcn"
ewgu"cuuqekcvgf"ykvj"tgrtqfwevkqp"xct{"ykfgn{"cetquu"vczc"cpf"ctg"nkmgn{"wpfgt"uvtqpi"
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"
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vq"kpenwfg"oketqqticpkuocn"ugswgpegu."uqog"yjkej"ctg"fktgevn{"cuuqekcvgf"ykvj"S.
crassipalpis"cpf"uqog"vjcv"ctg"kpcfxgtvgpv"gpxktqpogpvcn"eqpvcokpcpvu0"Yg"kfgpvkhkgf"
qpn{"382"pqp/ogvc¦qcp"ugswgpegu"vjcv"crrgctgf"vq"dg"kpfgrgpfgpv"vtcpuetkrvu"ykvj"cp"
G>202223"htqo"qwt"DNCUVZ"cpf"DNCUVP"ugctejgu0"Cnvjqwij"yg"gzrgev"vjcv"hnguj"
hnkgu"jcxg"c"tkej"oketqqticpukocn"eqoowpkv{."qwt"ugngevkqp"qh"rqn{/C"TPC"cu"vjg"
uvctvkpi"ocvgtkcn"hqt"qwt"nkdtct{"nkmgn{"gnkokpcvgf"ocp{"rqvgpvkcn"oketqdkcn"ugswgpegu0"
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" /";"/"
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kpenwfkpi"{gcuvu"*g0i0."Candida, Saccharomyces+."hkncogpvqwu"hwpik"*g0i0."Aspergillus,
Neurospora+."cpf"rtqvq¦qcpu"*g0i0."Plasmodium, Theileria+0"Ugswgpegu"jcxkpi"
crrctgpv"dcevgtkcn"qtkikp"kpenwfgf"ocp{"igpgtc"ykvj"urgekgu"vjcv"jcxg"dggp"cuuqekcvgf"
ykvj"kpugevu"gkvjgt"cu"rcvjqigpu"qt"cu"pqtocn"hnqtc"kp"vjg"iwv"*g0i0."Burkholderia"cpf"
Escherichia+."dwv"vjgug"ugswgpegu"eqwnf"lwuv"cu"gcukn{"tgrtgugpv"gpxktqpogpvcn"
eqpvcokpcvkqp0"Rgtjcru"oquv"kpvgtguvkpi"qh"vjg"pqp/ogvc¦qcp"ugswgpegu"ygtg"vjqug"
hqt"vtcpurqucdng"gngogpvu."rctvkewnctn{"ugswgpegu"kp"vjg"Mariner-Tc1"hcokn{"cpf"
ugxgtcn"tgvtqxktcn"gngogpvu0"Ykvj"hwvwtg"fgxgnqrogpv"uwej"ugswgpegu"oc{"rtqxkfg"
kpukijv"kpvq"pgy"hwpevkqpcn"igpqokeu"vqqnu"hqt"Sarcophaga,"rqvgpvkcnn{"kpenwfkpi"
vtcpuigpke"oqfkhkecvkqp0""
"
Cpqvjgt"kpvgtguvkpi"hgcvwtg"qh"qwt"676"ugswgpeg"fcvc"ku"vjcv"c"pwodgt"qh"ugswgpegu"
crrgct"vq"eqpvckp"tgrgcv"oqvkhu"*k0g0."oketqucvgnnkvgu+"qt"ukping"pwengqvkfg"
rqn{oqtrjkuou"*UPRu+0"Yg"ugctejgf"hqt"oketqucvgnnkvg"oqvkhu"ykvjkp"qwt"ugswgpegu"
wukpi"vjg"rtqitco"OUCVEQOOCPFGT"]46_"vq"kfgpvkh{"ugswgpegu"ykvj"fk/."vtk/."
vgvtc/."rgpvc."cpf"jgzcpwengqvkfg"tgrgcvu0"Qwt"ugctej"hqt"tgrgcv"oqvkhu"tgxgcngf"743"
ugswgpegu"eqpvckpkpi"uqog"uwej"oqvkh"htqo"vjg"vqvcn"ugv"qh"74.275"ugswgpegu"*wukpi"c"
okpkowo"tgrgcv"ngpivj"qh"ugxgp."Cffkvkqpcn"hkng"4+0"Vjg"jkij"ngxgnu"qh"xctkcvkqp"
v{rkecnn{"hqwpf"hqt"oketqucvgnnkvg"octmgtu"jcxg"ngf"vq"vjgkt"ykfgurtgcf"wug"hqt"c"dtqcf"
tcpig"qh"uvwfkgu"uwej"cu"igpqog"ocrrkpi."rctgpvcig"cpcn{uku."cpf"cpcn{ugu"qh"igpg"
hnqy"cpf"igpgvke"uvtwevwtg0"Dgecwug"qwt"nkdtct{"ycu"eqpuvtwevgf"htqo"c"nqpi/uvcpfkpi"
ncdqtcvqt{"eqnqp{"gzrgevgf"vq"jctdqt"nkvvng"uvcpfkpi"xctkcvkqp"eqorctgf"vq"hkgnf"
rqrwncvkqpu."yg"fkf"pqv"uetggp"cp{"qh"vjgug"nqek"hqt"cnngnke"xctkcdknkv{0"Ukoknctn{."vq"
kfgpvkh{"UPRu"kp"qwt"eqpvkiu"yg"wugf"vjg"rtqitco"Rqn{Dc{guUjqtv."c"xgtukqp"qh"vjg"
" /"32"/"
Rqn{Dc{gu"rtqitco"]47_"vjcv"jcu"dggp"qrvkok¦gf"hqt"fcvc"htqo"ujqtv"tgcf"rncvhqtou"
uwej"cu"676."wpfgt"vjg"fghcwnv"rctcogvgtu0"Yg"hqwpf"cv"vqvcn"qh"34.2;2"UPRu"vjcv"ygtg"
fkuvtkdwvgf"coqpi"5658"qh"qwt"42.;;7"eqpvkiu."{kgnfkpi"cp"cxgtcig"qh"9045"UPRu"rgt"
32md0"Jqygxgt."dgecwug"qwt"ucorngu"ygtg"fgtkxgf"htqo"c"nqpi/uvcpfkpi."kpdtgf"
ncdqtcvqt{"eqnqp{"qwt"UPR"guvkocvgu"ctg"nkmgn{"c"uocnn"htcevkqp"qh"vjg"xctkcvkqp"
gzkuvkpi"kp"S. crassipalpis"rqrwncvkqpu"kp"vjg"hkgnf0"Yg"jqrg"vjcv"kfgpvkhkecvkqp"qh"
ecpfkfcvg"oketqucvgnnkvg"cpf"UPR"nqek"yknn"dg"wughwn"hqt"qvjgtu"kpvgtguvgf"kp"hkgnf/
dcugf"rqrwncvkqp"uvwfkgu"qh"S. crassipalpis"cpf"enqugn{"tgncvgf"Sarcophaga"urgekgu."
uqog"qh"yjkej"ctg"korqtvcpv"hqt"hqtgpuke"uvwfkgu0""
"
Conclusions Uvwfkgu"qh"hnguj"hnkgu"kp"vjg"igpwu"Sarcophaga"jcxg"eqpvtkdwvgf"itgcvn{"vq"qwt"
wpfgtuvcpfkpi"qh"fkcrcwug."uvtguu"tgukuvcpeg."tgrtqfwevkqp."cpf"koowpkv{"kp"kpugevu."
dwv"yqtm"kp"vjku"Ðpqp/vtcfkvkqpcnÑ"oqfgn"qticpkuo"jcu"dggp"jcorgtgf"d{"c"ncem"qh"
igpqoke"tguqwtegu0"Qwt"wug"qh"r{tqugsgpekpi"vq"fgxgnqr"c"pgy"GUV"nkdtct{"
eqpvckpkpi"crrtqzkocvgn{";.222"vtcpuetkrvu"rtqxkfgu"c"pgy"tguqwteg"hqt"kppqxcvkxg"
yqtm"vjcv"yg"jqrg"yknn"ftcy"tgpgygf"kpvgtguv"kp"wukpi"hnguj"hnkgu"cu"c"oqfgn0"Yg"
gzrgev"vjcv"vjgug"GUV"fcvc"yknn"rtqxkfg"vjg"hqwpfcvkqp"hqt"c"rtqnkhgtcvkqp"qh"hwpevkqpcn"
yqtm"qp"hnguj"hnkgu"kpenwfkpi"ÐÎqokeuÑ"crrtqcejgu"uwej"cu"vjg"fgxgnqrogpv"qh"
oketqcttc{u"hqt"nctig"uecng"igpg"gzrtguukqp"uvwfkgu"cu"ygnn"cu"ugtxkpi"cu"c"hqwpfcvkqp"
hqt"kfgpvkh{kpi"rtgxkqwun{"wptgeqipk¦cdng"rtqvgkpu"kp"rtqvgqokeu"uvwfkgu0"Vjgug"fcvc"
yknn"cnuq"kpetgcug"qwt"cdknkv{"vq"gzrnqtg"urgekhke"igpg"cpf"rcvjyc{/dcugf"crrtqcejgu"
uwej"cu"vtcpuetkrv"swcpvkhkecvkqp"d{"sTV/RET"cpf"TPC"kpvgthgtgpeg"qh"ecpfkfcvg"
igpgu"vjcv"jcxg"rtgxkqwun{"dggp"tghtcevqt{"vq"jqoqnqi{/dcugf"enqpkpi0"Hwtvjgtoqtg."
vjgug"fcvc"eqornkogpv"vjg"itqykpi"pwodgt"qh"GUV"cpf"igpqog"rtqlgevu"kp"hnkgu"qvjgt"
" /"33"/"
vjcp"Drosophila"cpf"oquswkvqgu."uwej"cu"vjg"ogfhn{"Ceratitis capitata cpf"vjg"
Jguukcp"hn{"Mayetiola destructor"]48."49_."gpjcpekpi"tguqwtegu"hqt"eqorctcvkxg"
uvwfkgu"qh"oqngewnct"gxqnwvkqp"cetquu"vjg"Fkrvgtc0"Qwt"kpkvkcn"eqorctcvkxg"cpcn{uku"qh"
gxqnwvkqpct{"tcvgu"cetquu"IQ"ecvgiqtkgu"knnwuvtcvgu"vjku"wvknkv{."cpf"yg"cpvkekrcvg"vjcv"
ukoknct"igpqog"cpf"vtcpuetkrvqog/ykfg"cpcn{ugu"yknn"{kgnf"korqtvcpv"kpukijvu"kpvq"
kpugev"fkxgtukhkecvkqp0"
Methods Vjg"ewnvwtg"qh"S. crassipalpis wugf"hqt"vjku"yqtm"qtkikpcvgf"kp"Eqnwodwu."Qjkq."cpf"
ycu"ockpvckpgf"kp"vjg"ncdqtcvqt{"hqt"pgctn{":"{gctu"dghqtg"vjg"qpugv"qh"vjku"rtqlgev0""
Pqp/fkcrcwukpi"rwrcg"ygtg"qdvckpgf"d{"tgctkpi"vjg"eqnqp{"wpfgt"nqpi/fc{"eqpfkvkqpu"
*37N<;F+"ykvj"cfwnvu"jgnf"cv"47qE"cpf"nctxcg"cpf"rwrcg"cv"42qE."yjgtgcu"fkcrcwukpi"
rwrcg"ygtg"igpgtcvgf"d{"tgctkpi"vjg"ocvgtpcn"igpgtcvkqp"qh"cfwnvu."nctxcg"cpf"rwrcg"
wpfgt"ujqtv/fc{"eqpfkvkqpu"*34N<34F+"ykvj"cfwnvu"jgnf"cv"47qE"cpf"nctxcg"cpf"rwrcg"
jgnf"cv"42qE."cu"rtgxkqwun{"fguetkdgf"]4:_0""
"
Yg"etgcvgf"c"ukping"pqtocnk¦gf"eFPC"nkdtct{"htqo"cnn"nkhg"uvcigu"vq"oczkok¦g"
tgrtgugpvcvkqp"qh"vjg"S. crassipalpis vtcpuetkrvqog0"Dgecwug"yg"ctg"rctvkewnctn{"
kpvgtguvgf"kp"rwrcn"fkcrcwug"cpf"tgrtqfwevkqp."yg"kpenwfgf"dqvj"fkcrcwug/fguvkpgf"
cpf"fktgev"fgxgnqrkpi"kpfkxkfwcnu."cpf"coqpi"cfwnvu"yg"kpenwfgf"dqvj"rtqvgkp/hgf"cpf"
rtqvgkp/uvctxgf"ocngu"cpf"hgocngu"ucorngf"htqo"genqukqp"vjtqwij"qpg"tgrtqfwevkxg"
dqwv"*ugg"Vcdng"3"hqt"hwnn"ucornkpi"uejgog+0"TPC"htqo"gcej"ucorng"itqwr"ycu"
gzvtcevgf"ugrctcvgn{"kp"Vtk/tgcigpv"*Codkqp+"cpf"vjg"swcnkv{"qh"vqvcn"TPC"ycu"
xgtkhkgf"qp"306'"cictqug/OQRU/hqtocnfgj{fg"fgpcvwtkpi"ignu0"Rqn{/C"TPC"ycu"
gzvtcevgf"htqo"gcej"vqvcn"TPC"ucorng"wukpi"vjg"Qnkiqvgz"̶"oTPC"Okfk"Mkv"
*Skcigp+"cpf"vjg"swcnkv{"cickp"xgtkhkgf"qp"c"fgpcvwtkpi"ign0"Dgecwug"vjg"vqvcn"oTPC"
" /"34"/"
{kgnf"htqo"gcej"rqqn"fkhhgtgf."3-i"qh"oTPC"htqo"gcej"ucorng"itqwr"ycu"eqodkpgf"
kpvq"c"ukping"nctig"rqqn"cpf"okzgf"ygnn0"Vjku"ukping"nctig."gswcnn{/okzgf"rqqn"ycu"vjg"
uqwteg"qh"uvctvkpi"ocvgtkcn"hqt"vjg"etgcvkqp"qh"vjg"pqtocnk¦gf"eFPC"nkdtct{0"Cp"kpkvkcn"
eFPC"nkdtct{"ycu"eqpuvtwevgf"wukpi"vjg"UOCTV"eFPC"U{pvjguku"Mkv"*DF"
Enqpgvgej."Oqwpvckp"Xkgy."EC+"cpf"pqtocnk¦gf"wukpi"vjg"Vtkoogt/Fktgev"Mkv"
*Gxtqigp."Oqueqy+0"Vjg"swcnkv{"qh"vjg"pqtocnk¦gf"eFPC"nkdtct{"ycu"ejgemgf"qp"c"
306'"cictqug"ign0"
"
Crrtqzkocvgn{"407-i"qh"vjg"pqtocnk¦gf"eFPC"rqqn"ycu"wugf"hqt"c"vkvtcvkqp"twp"wukpi"
qpg"swctvgt"qh"c"rncvg"qp"vjg"Tqejg"IU"HNZ"ugswgpegt"cpf"ycu"hqnnqygf"d{"c"yjqng"
rncvg"twp"qp"vjg"ucog"gswkrogpv"wukpi"37-i"qh"eFPC"cpf"c"hwnn"IU"HNZ"rncvg"cv"vjg"
Kpvgtfkuekrnkpct{"Egpvgt"hqt"Dkqvgejpqnqi{"Tgugctej"cv"vjg"Wpkxgtukv{"qh"Hnqtkfc"*WH/
KEDT+0"Vjg"r{tqnwokpguegpeg"kpvgpukv{"hqt"gcej"dgcf/eqpvckpkpi"ygnn"ycu"wugf"vq"
fgvgtokpg"dcug"ecnnu"hqt"gcej"pwengqvkfg"kpeqtrqtcvkqp"uvgr0"Ugswgpeg"cuugodn{"ycu"
rgthqtogf"hktuv"wukpi"vjg"Pgydngt"Cuugodngt"x303024037"*676"Nkhg"Uekgpeg."Dtcphqtf."
EV+0"Chvgtyctfu."Pgydngt"qwvrwv."kpenwfkpi"cnn"eqpvkiu"cpf"ukpingvqpu."ycu"uwdlgevgf"
vq"c"ugeqpf"cuugodn{"uvgr"wukpi"vjg"Rctcegn"Vtcpuetkrv"Cuugodngt"x50202"*Rctcegn"Kpe0."
Rcucfgpc."EC+0"Ugswgpegu"ygtg"ocumgf"hqt"qnkiqpwengqvkfg"cfcrvqtu"wugf"kp"UOCTV"
eFPC"u{pvjguku"cpf"pqtocnk¦cvkqp"wukpi"c"vjtgujqnf"qh"370"Nqy/swcnkv{"fcvc"*dcug"
ecnn"ueqtg">"32+"ycu"vtkoogf"htqo"vjg"gpfu"qh"kpfkxkfwcn"ugswgpegu0"Nqy"eqorngzkv{"
ugswgpeg"tgikqpu"*ukorng"ugswgpeg"tgrgcvu+"ygtg"kfgpvkhkgf"cpf"gzenwfgf"htqo"
eqpukfgtcvkqp"fwtkpi"kpkvkcn"rckt/ykug"eqorctkuqp."dwv"ygtg"kpenwfgf"fwtkpi"hkpcn"
cnkipogpv"cpf"eqpugpuwu"dwknfkpi0"Cuugodn{"ycu"rgthqtogf"kp"vyq"uvcigu0"Vjg"hktuv"
uvcig"wugf"vjg"$Jcuvg$"cniqtkvjo"vq"dwknf"itqwru"*qt"enwuvgtu+"qh"ugswgpegu"ujctkpi"c"
okpkocn"coqwpv"qh"kfgpvkhkcdng"ugswgpeg"ukoknctkv{"*vjtgujqnf"?"72+0"Vjg"ugeqpf"uvcig"
" /"35"/"
cuugodngf"ugswgpegu"ykvjkp"kpfkxkfwcn"enwuvgtu"kpvq"eqpugpuwu"vtcpuetkrvu"wukpi"vjg"
uqhvyctg"fghcwnvu."gzegrv"rctcogvgtu"OkpEqxTgr"*722+."KpQxgtjcpi"*52+."
GpfQxgtjcpi"*52+."TgoQxgtjcpi"*52+."SwcnUwoNko"*522+."OczKpvgtpcnIcru"*37+"
cpf"Rgpcnk¦gP"*2+0"Hqt"qwt"fcvcugv."cp"cfxcpvcig"qh"vjg"ugeqpf"cuugodn{"uvgr"wukpi"
Rctcegn"Vtcpuetkrv"Cuugodngt"ycu"kp"engcpkpi/wr"vjg"fcvc"d{"hwtvjgt"ocumkpi"cp{"
UOCTV"cfcrvqt"ugswgpegu"qt"rqn{/C"uvtgvejgu"vjcv"ygtg"kpenwfgf"kp"vjg"kpkvkcn"
Pgydngt"cuugodn{0""
"
Cnn"eqpvkiu"cpf"ukpingvu"ygtg"cppqvcvgf"d{"DNCUV"ugctej"cickpuv"PEDK"PT"cpf"PV"
fcvcdcugu"yjgtg"vjg"g/xcnwg"vjtgujqnf"ycu"ugv"cv"3g/60"Hqt"gcej"swgt{"ugswgpeg."vjg"
vqr"322"DNCUV"jkvu"ygtg"qdvckpgf"kh"cxckncdng"cpf"uvqtgf"kp"DncuvSwguv"]4;_0"Kh"c"
DNCUV"jkv"tgrtgugpvgf"c"PEDK"Igpg"fcvcdcug"gpvt{."kv"ycu"wugf"vq"ocr"Sarcophaga "
vtcpuetkrvu"vq"rwvcvkxg"jqoqnqiu"htqo"Drosophila."jwocp."oqwug."¦gdtchkuj."fqi"qt"
qvjgt"qticpkuou0"Igpg"Qpvqnqi{"*IQ+"cppqvcvkqpu"ygtg"cnuq"fgtkxgf"dcugf"qp"
ugswgpeg"ukoknctkv{0"Vtcpuetkrvu"ygtg"encuukhkgf"kpvq"36"oclqt"Igpg"Qpvqnqi{"
ecvgiqtkgu"wpfgt"dkqnqikecn"rtqeguu0"Kh"c"vtcpuetkrv"ycu"cppqvcvgf"ykvj"oqtg"vjcp"qpg"
IQ"ecvgiqt{"kv"ycu"urnkv"gswcnn{"coqpi"vjgo0"""
"
Yg"cnuq"wugf"qwt"IQ"cppqvcvkqpu"vq"cuuguu"yjgvjgt"eqfkpi"ugswgpeg"fkxgtigpeg"
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vtcpuncvkqp"fcvcdcugu"hqt"D. melanogaster"cpf"A. gambiae"htqo"Hn{Dcug"
*jvvr<11hn{dcug0qti1+"cpf"XgevqtDcug"*jvvr<11yyy0xgevqtdcug0qti1+."tgurgevkxgn{0"Yg"
vjgp"wugf"qwt"S. crassipalpis"GUV"fcvcdcug"cu"c"swgt{"kp"c"DncuvZ"ugctej"cickpuv"gcej"
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xcnwgu"hqt"cnn"qvjgt"rctcogvgtu0"Vjg"676"ugswgpekpi"vgejpqnqi{"ku"rtqpg"vq"rtqfwekpi"
" /"36"/"
crrctgpv"htcog/ujkhv"gttqtu"xkc"gttqpgqwu"kpugtvkqpu"qt"fgngvkqpu"qh"gzvtc"dcugu"]52_."
nkokvkpi"vjg"swcnkv{"cpf"ngpivj"qh"eqfqp"cnkipogpvu"cpf"dncuvZ"jkvu0"Ykvjqwv"
eqttgevkqp."eqfqpu"cpf"rtqvgkpu"yknn"cnkip"kp"qxgtncrrkpi."htcog/ujkhvgf"ugiogpvu."cpf"
vq"qwt"mpqyngfig"pq"r{tqugswgpekpi"uvwf{"jcu"rtgxkqwun{"cfftguugf"vjku"kuuwg0"Yg"
fgxgnqrgf"c"rtqegfwtg"vq"kfgpvkh{"cpf"eqttgev"hqt"676"kpugtvkqp"*qxgtecnn+"cpf"fgngvkqp"
*wpfgtecnn+"gttqtu"kpenwfkpi"vjg"hqnnqykpi"uvgru<"3+"Kfgpvkh{"DncuvZ"jkvu"ykvj"vyq"
htcog/ujkhvgf."qxgtncrrkpi"Jkij/ueqtkpi"Ugiogpv"Rcktu"*JURu+"dqvj"ykvj"g/xcnwgu"
nguu"vjcp"3z32/7"*kh"rtgugpv."yg"gzenwfgf"cffkvkqpcn."jkijgt"g/xcnwg"JURu"htqo"qwt"
cnkipogpvu+0"4+"Fgvgtokpg"rtqvgkp"cnkipogpv"ueqtg"hqt"gcej"JUR"kp"vjg"tgikqp"qh"
qxgtncr0"Yg"ecnewncvgf"cnkipogpv"ueqtgu"d{"cuukipkpi"c"xcnwg"qh"3"hqt"kfgpvkecn"
tgukfwgu."207"hqt"c"rqukvkxg"cnkipogpv"ueqtg"kp"vjg"DNQUWO84"ueqtkpi"ocvtkz."cpf"2"
hqt"pqp/rqukvkxg""DNQUWO84"ueqtgu"qt"c"icr0"5+"Tgvckp"vjg"tgikqp"qh"qxgtncr"kp"vjg"
JUR"cnkipogpv"ykvj"vjg"jkijguv"cnkipogpv"ueqtg."cpf"tgoqxg"vjg"tgikqp"qh"qxgtncr"
htqo"dqvj"vjg"wp/cnkipgf"pwengqvkfg"cpf"cnkipgf"rtqvgkp"ugswgpeg"qh"vjg"nqygt"ueqtkpi"
JUR0"6+"Tgoqxg"qpg"qt"vyq"cffkvkqpcn"pwengqvkfgu"rcuv"vjg"tgikqp"qh"qxgtncr"kp"vjg"
wp/cnkipgf"pwengqvkfg"ugswgpeg"hqt"vjg"nqygt/ueqtkpi"JUR"kh"vjg"htcogujkhv"ku"-3"qt"
-4."tgurgevkxgn{0"Vjgug"ctg"pwengqvkfgu"kp"vjg"7)"qt"5)"fktgevkqp"kh"vjg"nqygt"ueqtkpi"
JUR"ku"oqtg"7)"qt"5)."tgurgevkxgn{0"7+"Kpugtv"c"ukping"icr"kp"vjg"swgt{"rtqvgkp"ugswgpeg"
cv"vjg"5)"gpf"qh"vjg"hktuv"JUR"*qt"vjg"7)"gpf"qh"vjg"ugeqpf"JUR+"kh"vjg"htcogujkhv"ku"-40"
8+"Eqpecvgpcvg"vjg"jkijguv"ueqtkpi"JUR"*rtqvgkp+"ykvj"vjg"nqyguv"ueqtkpi."vtwpecvgf"
JUR0"Vjg"rtqegfwtg"eqttgevu"hqt"ukping"kpugtvkqp"cpf"fgngvkqp"gttqtu"vq"rtqfweg"vjg"
dguv"ueqtkpi"hwnn"cnkipogpv"dcugf"qp"qwt"ueqtkpi"etkvgtkc"hqt"vjg"tgikqp"qh"qxgtncr0"Kp"
vjg"ecug"qh"cp"kpugtvkqp"*-3"htcogujkhv+"gttqt."c"ukping"pwengqvkfg"ku"tgoqxgf"kp"vjg"
eqttgevkqp0"Kp"vjg"ecug"qh"c"fgngvkqp"*-4"htcogujkhv+"vyq"pwengqvkfgu"ctg"tgoqxgf"cpf"c"
icr"kp"vjg"rtqvgkp"ugswgpeg"ku"kpvtqfwegf0"Vjku"ghhgevkxgn{"tgoqxgu"c"eqfqp"htqo"vjg"
" /"37"/"
cnkipogpv."tghngevkpi"vjg"wpegtvckpv{"kp"tgeqpuvtwevkpi"vjg"vtcpuncvgf"cokpq"cekf"
cuuqekcvgf"ykvj"vjg"fgngvkqp0"Qwv"qh"34.543"vqvcn"dncuvZ"jkvu"htqo"c"Sarcophaga"vq"
Drosophila"vtcpuncvkqp"fcvcdcug"ugctej."3258"*:"rgtegpv+"jcf"rwvcvkxg"htcog/ujkhv"
gttqtu."8:3"-3"*kpugtvkqp+"ujkhvu"cpf"577"-4"*fgngvkqp+"ujkhvu0"Engctn{"eqttgevkpi"vjgug"
htcog/ujkhv"gttqtu"yknn"uwduvcpvkcnn{"kortqxg"igpqog/ykfg"cnkipogpvu"wukpi"cp{"676/
fgtkxgf"vtcpuetkrvqog0"""
"
Chvgt"crrn{kpi"htcog/ujkhv"eqttgevkqpu."yg"tgvckpgf"vjg"vqr/ueqtkpi"cokpq"cekf"
cnkipogpv"htqo"gcej"DncuvZ"jkv"cpf"rtqfwegf"vjg"cuuqekcvgf"vtcpuetkrv"cnkipogpvu"
wukpi"vtcpcnkip"kp"vjg"GODQUU"rcemcig"]53_"kp"eqplwpevkqp"ykvj"vtcpuetkrvkqp"
fcvcdcugu"hqt"D. melanogaster"cpf"A. gambiae0"Wukpi"vjg"rtqitco"[P22"htqo"vjg"
RCON"rcemcig"]54_"yg"guvkocvgf"u{pqp{oqwu"*fU+"cpf"pqp/u{pqp{oqwu"*fP+"
uwduvkvwvkqp"tcvgu"hqt"gcej"cnkipogpv"xkc"vjg"ogvjqf"qh"[cpi"cpf"Pkgnugp"]55_0"Qwt"
kpkvkcn"kpvgpvkqp"ycu"vq"guvkocvg"fP1fU"tcvkqu."dwv"cnkipogpvu"ygtg"nctign{"
qxgtucvwtcvgf"ykvj"u{pqp{oqwu"uwduvkvwvkqp"tcvgu"*qhvgp"kpguvkocdng+0"Vjwu."yg"
tgvckpgf"qpn{"fP"guvkocvgu0"Hkpcnn{."yg"guvkocvgf"vjg"ogfkcp."kpvgtswctvkng"tcpig"
*KST+."cpf"cp"guvkocvg"qh"vjg";7'"eqphkfgpeg"kpvgtxcn"qh"vjg"ogfkcp"cu"ł"307:"
KST1ustv*p+"hqt"fP"xcnwgu"htqo"gcej"IQ"ecvgiqt{"*igpgu"hcnnkpi"kpvq"oqtg"vjcp"qpg"
IQ"ecvgiqt{"eqpvtkdwvg"vq"guvkocvgu"kp"cnn"cuuqekcvgf"ecvgiqtkgu+0"Vjku"rctvkewnct";7'"
EK"guvkocvg"rtqxkfgu"c"eqphkfgpeg"kpvgtxcn"vjcv"ku"nctign{"kpugpukvkxg"vq"vjg"wpfgtn{kpi"
fkuvtkdwvkqp"]56_0"Yg"cnuq"rgthqtogf"cnn"qh"vjg"cdqxg"uvgru"qp"vjg"rcktykug"
eqorctkuqp"dgvyggp"D. melanogaster"cpf"A. gambiae."{kgnfkpi"vjtgg"rcktykug"
eqorctkuqpu"kp"vjg"eqorngvg"cpcn{uku0"
" /"38"/"
Authors' contributions
FCJ"eqpegkxgf"vjg"uvwf{"cpf"fgukip."eqnngevgf"ucorngu."dwknv"vjg"nkdtctkgu."
rctvkekrcvgf"kp"vjg"fcvc"cpcn{uku"cpf"ftchvgf"vjg"ocpwuetkrv0"ILT"rctvkekrcvgf"kp"vjg"
fcvc"cpcn{uku"cpf"ftchvgf"vjg"ocpwuetkrv0"FFU"rctvkekrcvgf"kp"vjg"fgukip"qh"vjg"uvwf{."
rctvkekrcvgf"kp"nkdtct{"eqpuvtwevkqp."fcvc"cpcn{uku."cpf"ftchvkpi"vjg"ocpwuetkrv0"FNF"
jgnrgf"eqpegkxg"vjg"uvwf{."rctvkekrcvgf"kp"vjg"fgukip"cpf"eqqtfkpcvkqp."cpf"ftchvgf"vjg"
ocpwuetkrv0"Cnn"cwvjqtu"tgcf"cpf"crrtqxgf"vjg"hkpcn"ocpwuetkrv0"
Acknowledgements Yg"vjcpm"Ftu0"Dknn"Hctogtkg"cpf"Nk"Nkw"htqo"vjg"WH/KEDT"eqtg"hceknkv{"hqt"rtqxkfkpi"
etkvkecn"cfxkeg"cpf"dkqkphqtocvkeu"uwrrqtv"fwtkpi"cnn"rjcugu"qh"vjku"rtqlgev."cu"ygnn"cu"
Fkgvoct"Uejyct¦"cpf"Tqd"Okejcwf"hqt"eqoogpvu"qp"vjg"ocpwuetkrv0"Vjku"yqtm"ycu"
uwrrqtvgf"d{"hwpfu"htqo"WUFC/EUTGGU"4226/57524/967;;6."PUH/KQU/863727."
cpf"vjg"Hnqtkfc"Uvcvg"Citkewnvwtcn"Gzrgtkogpv"Uvcvkqp"vq"FJ."WUFC"PTKEIR"4228/
57524/3:223"*Citggogpv"Pq0"7:/8837/9/359+"vq"FU."cpf"PUH/KQD/2638942"vq"FF0"
References 1. Fraenkel G, Sivasubramanian P, Zdarek J: Hormonal factors in CNS
and hemolymph of pupariating fly larvae which accelerate puparium formation and tanning. Biol Bull 1972, 143(1):127-139.
2. Verleyen P, Huybrechts J, Sas F, Clynen E, Baggerman G, De Loof A, Schoofs L: Neuropeptidomics of the grey flesh fly, Neobellieria bullata. Biochem Biophys Res Commun 2004, 316(3):763-770.
3. Denlinger DL: Regulation of diapause. Annu Rev Entomol 2002, 47:93-122.
4. Hahn DA, Denlinger DL: Meeting the energetic demands of insect diapause: Nutrient storage and utilization. J Insect Physiol 2007, 53(8):760-773.
5. Lee RE, Chen CP, Denlinger DL: A rapid cold-hardening process in insects Science 1987, 238(4832):1415-1417.
6. Rinehart JP, Li A, Yocum GD, Robich RM, Hayward SAL, Denlinger DL: Up-regulation of heat shock proteins is essential for cold survival during insect diapause. PNAS 2007, 104(27):11130-11137.
7. Browne LB: Quantitative aspects of the regulation of ovarian development in selected anautogenous Diptera: integration of
" /"39"/"
endocrinology and nutrition. Entomologia Experimentalis et Applicata 2001, 100(2):137-149.
8. Hahn DA, James LN, Milne KR, Hatle JD: Life-history plasticity after attaining a dietary threshold for reproduction is associated with protein storage in flesh flies. Funct Ecol 2008, 22(6):1081-1090.
9. Natori S, Shiraishi H, Hori S, Kobayashi A: The roles of Sarcophaga defense molecules in immunity and metamorphosis. Developmental & Comparative Immunology 1999, 23(4-5):317-328.
10. Pape T: Catalogue of the Sarcophagidae of the world. Memoires of Entomology International 1996, 8:1-558.
11. Wells JD, Stevens JR: Application of DNA-based methods in forensic entomology. Annu Rev Entomol 2008, 53(1):103-120.
12. Sherman RA, Hall MJR, Thomas S: Medicinal maggots: An ancient remedy for some contemporary afflictions. Annu Rev Entomol 2000, 45(1):55-81.
13. Legner EF: Biological control of Diptera of medical and vetrinary importance J Vector Ecol 1995, 20(1):59-120.
14. Rivers DB, Ruggiero L, Hayes M: The ectoparasitic wasp Nasonia vitripennis (Walker) (Hymenoptera : Pteromalidae) differentially affects cells mediating the immune response of its flesh fly host, Sarcophaga bullata Parker (Diptera : Sarcophagidae). J Insect Physiol 2002, 48(11):1053-1064.
15. Beukeboom L, Desplan C: Nasonia. Curr Biol 2003, 13(22):R860-R860.
16. Opijnen T, Baudry E, Baldo L, Bartos J, Werren JH: Genetic variability in the three genomes of Nasonia: nuclear, mitochondrial and Wolbachia. Insect Mol Biol 2005, 14(6):653-663.
17. Flannagan RD, Tammariello SP, Joplin KH, Cikra-Ireland RA, Yocum GD, Denlinger DL: Diapause-specific gene expression in pupae of the flesh fly Sarcophaga crassipalpis. PNAS 1998, 95(10):5616-5620.
18. Yamamoto-Kihara M, Kotani E: Isolation and characterization of a C-type lectin cDNA specifically expressed in the tip of mouthparts of the flesh fly Sarcophaga peregrina. Insect Mol Biol 2004, 13(2):133-140.
19. Toth AL, Varala K, Newman TC, Miguez FE, Hutchison SK, Willoughby DA, Simons JF, Egholm M, Hunt JH, Hudson ME et al: Wasp gene expression supports an evolutionary link between maternal behavior and eusociality. Science 2007, 318(5849):441-444.
20. Vera JC, Wheat CW, Fescemyer HW, Frilander MJ, Crawford DL, Hanski I, Marden JH: Rapid transcriptome characterization for a nonmodel organism using 454 pyrosequencing. Molecular Ecology 2008, 17(7):1636-1647.
21. Zou Z, Najar F, Wang Y, Roe B, Jiang H: Pyrosequence analysis of expressed sequence tags for Manduca sexta hemolymph proteins involved in immune responses. Insect Biochemistry and Molecular Biology 2008, 38(6):677-682.
22. Huse S, Huber J, Morrison H, Sogin M, Welch D: Accuracy and quality of massively parallel DNA pyrosequencing. Genome Biology 2007, 8(7):R143.
" /"3:"/"
23. Friedrich M, Tautz D: Evolution and phylogeny of the Diptera: A molecular phylogenetic analysis using 28S rDNA sequences. Systematic Biology 1997, 46(4):674-698.
24. Faircloth BC: MSATCOMMANDER: detection of microsatellite repeat arrays and automated, locus-specific primer design. Mol Ecol Resour 2008, 8(1):92-94.
25. Marth GT, Korf I, Yandell MD, Yeh RT, Gu ZJ, Zakeri H, Stitziel NO, Hillier L, Kwok PY, Gish WR: A general approach to single-nucleotide polymorphism discovery. Nature Genet 1999, 23(4):452-456.
26. Chen M-S, Zhao H-X, Zhu YC, Scheffler B, Liu X, Liu X, Hulbert S, Stuart JJ: Analysis of transcripts and proteins expressed in the salivary glands of Hessian fly (Mayetiola destructor) larvae. J Insect Physiol 2008, 54(1):1-16.
27. Gomulski LM, Dimopoulos G, Xi ZY, Soares MB, Bonaldo MF, Malacrida AR, Gasperi G: Gene discovery in an invasive tephritid model pest species, the Mediterranean fruit fly, Ceratitis capitata. BMC Genomics 2008, 9:15.
28. Denlinger DL: Induction and termination of pupal diapause in Sarcophaga (Diptera: Sarcophagidae) Biol Bull 1972, 142(1):11-24.
29. Farmerie WG, Hammer J, Liu L, Sahni A, Schneider M: Biological workflow with BlastQuest. Data Knowl Eng 2005, 53(1):75-97.
30. Brockman W, Alvarez P, Young S, Garber M, Giannoukos G, Lee WL, Russ C, Lander ES, Nusbaum C, Jaffe DB: Quality scores and SNP detection in sequencing-by-synthesis systems. Genome Res 2008, 18(5):763-770.
31. Rice P, Longden I, Bleasby A: EMBOSS: The European molecular biology open software suite. Trends Genet 2000, 16(6):276-277.
32. Yang Z: PAML 4: Phylogenetic analysis by maximum likelihood. Molecular Biology and Evolution 2007, 24(8):1586-1591.
33. Yang Z, Nielsen R: Estimating synonymous and nonsynonymous substitution rates under realistic evolutionary models. Molecular Biology and Evolution 2000, 17(1):32-43.
34. McGill R, Tukey JW, Larsen WA: Variations of box plots. American Statistician 1978, 32(1):12-16.
" /"3;"/"
Figures
Figure 1 - Sarcophaga crassipalpis sequences were classified into one of 14 major sub-categories within the Biological Processes GO category.
Figure 2 - A comparison of the distribution across 14 major Biological Process GO sub-classes in our Sarcophaga crassipalpis library versus predicted ESTs from Drosophila melanogaster.
Vjg"uwd/ecvgiqtkgu"ctg"EU"?"egnn"eqoowpkecvkqp"*ukipcnkpi+."TR"?"Tgiwncvkqp"qh"
egnnwnct"rj{ukqnqikecn"rtqeguu."V"?"Vtcpurqtv."QD"?"Egnn"qticpk¦cvkqp"cpf"dkqigpguku."
O"?"Ogvcdqnkuo."TU"?"Tgurqpug"vq"uvkownwu."EC"?"Egnn"cfjgukqp."EF"?"Egnn"fgcvj."
T"?"Tgrtqfwevkqp."EE"?"Egnn"e{eng"cpf"fkxkukqp."J"?"Jqogquvcuku."EO"?"Egnn"
oqvknkv{."F"?"Fgxgnqrogpv."cpf"IR"?"Egnn"itqyvj."fkhhgtgpvkcvkqp."cpf"rtqnkhgtcvkqp0""
"
Figure 3 - Box plots of the distributions of non-synonymous substitution rate
(dN) values in each GO category.
Vjg"rcktykug"eqorctkuqpu"ctg<"S. crassipalpis"vq"D. melanogaster"*c+."S. crassipalpis
vq"A. gambiae"*d+."cpf"A. gambiae vq D. melanogaster"*e+0"Vjg"wrrgt"cpf"nqygt"
dqwpfu"qh"vjg"pqvejgu"tgrtgugpv"vjg";7'"eqphkfgpeg"kpvgtxcn"qh"vjg"ogfkcp."cpf"vjg"
pwodgtu"kpukfg"vjg"dqzgu"tgrtgugpv"vjg"tgncvkxg"tcpm"ykvjkp"gcej"rcktykug"eqorctkuqp0"
Tables
Table 1 - RNA Extraction Strategy
Hnguj"hn{"nkhg"uvcigu"vjcv"ygtg"kpfkxkfwcnn{"TPC"gzvtcevgf"cpf"eqodkpgf"gswcnn{"rtkqt"
vq"eFPC"nkdtct{"u{pvjguku"cpf"pqtocnk¦cvkqp0"
"
Rtg/cfwnv"uvcigu" Cfwnv"Uvcigu"
Gzrqugf"vq"ujqtv"fc{u"vq"
kpfweg"fkcrcwug"
Gzrqugf"vq"nqpi"fc{u"vq"
kpfweg"fktgev"fgxgnqrogpv"
Pqp/
tgrtqfwevkxg"Hgf"
Tgrtqfwevkxg"
Hgf"uwict"cpf"
" /"42"/"
fgxgnqrogpv" qpn{"uwict"" rtqvgkp"*nkxgt+""
Hktuv"Kpuvct"Nctxcg"
"
Hktuv"Kpuvct"Nctxcg" Fc{"3"Ocngu" Fc{"3"Ocngu"
Ugeqpf"Kpuvct"Nctxcg"
"
Ugeqpf"Kpuvct"Nctxcg" Fc{"3"Hgocngu" Fc{"3"Hgocngu"
Gctn{"Vjktf"Kpuvct"Nctxcg"
"
Gctn{"Vjktf"Kpuvct"Nctxcg" Fc{"4"Ocngu" Fc{"4"Ocngu"
Ncvg"Vjktf"Kpuvct"Nctxcg"
"
Ncvg"Vjktf"Kpuvct"Nctxcg" Fc{"4"Hgocngu" Fc{"4"Hgocngu"
Ycpfgtkpi"Nctxcg"
"
Ycpfgtkpi"Nctxcg" Fc{"5"Ocngu" Fc{"5"Ocngu"
Rwrcg" Gctn{"Fkcrcwug"Rwrcg""
*37"fc{u+"
Fc{"5"Hgocngu" Fc{"5"Hgocngu"
Gctn{"Rjctcvg"Cfwnv" Okf/fkcrcwug"Rwrcg""
*52"fc{u+"
Fc{"6"Ocngu"
"
Fc{"6"Ocngu"
Okf/Rjctcvg"Cfwnv""
*tgf"g{g"uvcig+"
Ncvg"Fkcrcwug"Rwrcg""
*82"Fc{u+"
Fc{"6"Hgocngu" Fc{"6"Hgocngu"
Ncvg"Rjctcvg"Cfwnv""
*dncem"dtkuvng"uvcig+"
" Fc{"7"Ocngu"
"
Fc{"7"Ocngu"
" " Fc{"7"Hgocngu"
"
Fc{"7"Hgocngu"
" " Fc{"9"Ocngu"
"
Fc{"9"Ocngu"
" " Fc{"9"Hgocngu"
"
Fc{"9"Hgocngu"
" " Fc{";"Ocngu" Fc{";"Ocngu"
" /"43"/"
"
" " Fc{";"Hgocngu" Fc{";"Hgocngu"
"
Table 2 - Summary of Sarcophaga crassipalpis EST data.
"
Vqvcn"Dcugu" 96.54;.266"
Jkij/swcnkv{"Tgcfu" """""429.332"
Cxgtcig"Tgcf"Ngpivj" """"""""""""463"
Pwodgt"qh"Eqpvkiu" """""""42.;;7"
Cxgtcig"Eqpvki"Ngpivj" """"""""""""554"
Tcpig"Eqpvki"Ngpivj" 52"vq"4.;7:"
Pwodgt"qh"Tgcfu"kp"Eqpvkiu" """""3:2.;96"
Pwodgt"qh"Ukpingvqpu" """""""53.278"
Wpkigpgu"ykvj"G">"20223"xu0"PT"qt"PV"Fcvcdcug" """""""33.979"
Guvkocvgf"Wpkswg"Vtcpuetkrvu" """"""""";.539"
"
Additional files Cffkvkqpcn"hkng"3"Î"Gzcorngu"qh"rqvgpvkcn"fkcrcwug/tgncvgf"eFPCu"kfgpvkhkgf"kp"
vjku"GUV"nkdtct{0"*Hqtocv<"RFH+""""Cffkvkqpcn"hkng"4"Î"Oketqucvgnnkvg"fkueqxgt{0""*Hqtocv<"RFH+"
"4c0"Uwooct{"qh"rqvgpvkcn"oketqucvgnnkvg"nqek"kfgpvkhkgf0"4d0"Nkuv"qh"eqpvkiu"cpf"
ukpingvqpu"eqpvckpkpi"rqvgpvkcn"oketqucvgnnkvg"nqek"kpenwfkpi"tgrgcv"v{rg"cpf"ngpivj0"
Other
1%
Cell motility
1%
Development
10%
Cell death
1%
Cell grow th,
differentiation &
proliferation
3%
Cell adhesion
1%
Reproduction
2%
Homeostasis
1%
Cell communication
(signaling)
5%Regulation of cellular
physiological process
6%
Transport
13%
Cell organization &
biogenesis
6%
Cell cycle and division
2%
Response to stimulus
7%
Metabolism
41%
Figure 1
% T
ranscri
pts
0
10
20
30
40
50
Sarcophaga crassipalpis
Drosophila melanogaster
CS RP T OB M RS CA CD R CC H CM GPDFigure 2
1.0
0.8
0.6
0.4
0.2
0.0
Resp. to
Stim
.
Tra
nsport
Repro
ductio
n
Reg. B
io. P
roc.
Meta
bolis
m
Hom
eosta
sis
Develo
pm
ent
Cell
Org
.
Cell
Motilit
y
Cell
Gro
wth
Cell
Death
Cell
Cycle
Cell
Com
m.
Cell
Adhesio
n
1 23 4 5 6 7 8 9 10 11 12 13 14
12 34 5 67
89 1011 121314
12 3 45 678 9 10111213 14
1.0
1.0
0.0
0.0
1.5
1.5
0.5
0.5
a
b
c
dN
(substitu
tions/s
ite)
Figure 3
Additional files provided with this submission:
Additional file 1: additionalfile1.pdf, 58Khttp://www.biomedcentral.com/imedia/7414739112619376/supp1.pdfAdditional file 2: additionalfile2.pdf, 204Khttp://www.biomedcentral.com/imedia/1938967490255101/supp2.pdf
