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Chromista, a group of algae with yellow plastids
Key words: Chromista; Cryptophyta; Haptophyta; phylogeny; stramenopile.
Shinya Uwai
Department of Environmental Science, Faculty of Science, Niigata University
Ikarashi 2, Nishiku, Niigata 950-2181, Japan§ÃćŒň(ǮǕħň'6 %Ɖƚ'3') 8"(%K{i4zEq%$'ƤƟ 'ơ"ǮƉ# 68(ǩƚƚ=3ǫÑ 1Ǭ
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S. Uwai (A) - 1

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GyoQTƯ'õ"9 (Cavalier-Smith 2004)
 äǗ&Ǯ®ßžƆƪġ& !(ǮGyoQT'ƜƉ(¹žƆƒ#!/#/8ǮƀƟ'Gx}\&Æ/:92ǮƜƉ'ƷĽ#%ªŒŒň(ƀƟ"9#ƮƲ:!9(Takishita & Uchida 1999, Zhang et al. 1999, Yoon et al. 2002, Li et al. 2005, Sanchez-Puerta et al. 2007)
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7:9# ÖÈ %:!9GAPDH T{dGƴ&(ǮƂƕƴ"¢TAj#ƜƉ"¢TAj 8Ǯ:'TAj'ǆß3ĥJap&K}\:!9ƀƟ4Ɖƚħň"(ǮƜƉ"(7 GAPDH ǆß(u{Ɵŕğ'3'"9 ǮGyoQT'ƜƉ"(Ǯu{Ɵŕğ'ǆß"(%ǮƂƕƴ"ĭƖ9TAj' GAPDH ǆß'ǌƣ&68Œ#Ɣ
7:9ǆß <:!9ǫHarper & Keeling 2003Ǭ
FBA &(Ǯclass I # class II #TAj áÒǮƀƟ4Ɖƚħň'ƜƉ"( class IƉ"( class II<:!9#2 '<:!9'&æǮGyoQT'ƜÖÈ:!9 (Patron et al. 2004)/ PRKǫPhosphoribulokinaseǬ# ǊƁ&!3ǮGyoQT ƀƟ"(%Ɖƚħň&Ƽƍ"8ǮƉƚƜƉ'žƆ"9o\vpO ÄGx}\&Æ/:9%$ǮŚÍ&Ć9ł3
9 Ǯ4(8GyoQT 3 Hw}j' PRK&Ƽƍ"9# Ŭ:!9 (Petersen et al.

2006):7'ä(ǮGyoQT'ƜƉ Ǯ©!ƀƟƷĽ"9# "(%ǮƀƟƜƉ'žƆ'% "3ǜñ&Ǥƍă Ǩ3'"9#=Ŭ!9ǫÑ 3ǮÂ¡'žƆĬǬİū&(ǮQ[uqadAw|bj[Ɵ|Gvj[Ɵ&² ǮļǞĴƟǤ%$ 7%9?wkDu}TǫǨǍ'Ǡ=½ŅǬ'ƜƉ3GyoQT#ÄĪ'ŉÿ=ĉ! 8 (ơ 1)ǮƀƟƷĽ'ƜƉ=3įªŒŒňƒ(Gyp?wkDu}T#!/#27:!9 (Cavalier-Smith
2004,  2007, Keeling 2009)
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ƀƟ

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ƂƕƴÔ

ƂƕƴÔ

u{ƟÔHarper & Keeling (2003)

FBA

Class IIClass IIClass IIClass II

Class I

Patron et al. (2004)

rpl36

c-typec-typep-typep-typep-typeRice & Palmer (2006)

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S. Uwai (A) - 3

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žƆǓĥJapǫîǬ
#ƜƉJapǫÂǬ'::&3# žƆǓ ' æ ĳ = Ŭ  Sanchez-Puerta et al. (2007)%$=3#&ÑĥJa
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Ɔ&!(ţŘ& Archibald 2009)Gyp?wkDu}T(Ǯ'6 %Q}d}Hw}j'~#!Å°ċ7:9#39 Ǯä(ĥJap&3# žƆƪġ"(ǮQ[uqadAwǫǭ?wkDu}TǬ#Ǯbj[ƟǮGvj[Ɵ'ǒ'¹žƆă=ǮŵĨş&ĊŬ96 %ƅĢ(ü7:!% (Harper et al. 2005):$; ǮĜƼ"(Q[uqadAwǭ?wkDu}T(vNv?ǫvNv?&!(ÜŔ'Ǡ=½ŅǬ# ¯'Q}d}Hw}j#Ƽƍ"9# ƅĢ ÖÈ:ǮSARǫ9( RASǬ# Hw}jÅ"/#27:9# 9 (Burki et al.
2007, Hackett et al. 2007, Baldauf 2008, Hampl et al. 2009, Reeb et al. 2009ǲÑ 3Ǯî¡'žƆĬ)~ė"bj[Ɵ#Gvj[Ɵ'Ƽƍă=ŬžƆƪġ'ƅĢ3ÖÈ:!9 (ƬǮPatron et al.
2007)
/Ǯ
Rice & Palmer (2006) &69#Ǯ
bj[Ɵ#Gvj[Ɵ'ƜƉrpl36ǫribosomal Protein
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# 951!%

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S. Uwai (A) - 4

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3'Ǯu{Ɵ9(ƀƟŕğ'ǆß áÒ9# Ę7&%'Z}T(Ǯ !(»ƛ3ƜƉ=ĉ!ÀƖă=Ŭ3'"9ÄĪ'Z}T(?wkDu}T'ǒ"9ƏĴƠ"3ÖÈ:!9 (Reyes-Prieto et al. 2008)ŎÒ(ƜƉ=ĉ%Ƽƍƒ!ƜƉ=3! $(ǮƜƉŌü'TAo{H=ďã9"ǌƦ%mA{["9 :&6ƜƉŌü'ıÁ&!(ú'7%9ŪŶ=ù%!(%7%°#(ĳ-3'&%7%7ƪ£ô  #( ǮĥJap&3# žƆƪġ& !(ǮQ}d}Hw}j'ǒ'žƆǓ=68İū&ďã9# ĀƦ"9/žƆƪġ"(%ǮŎÒ(ǴŃƚǵ'Hw}j'Japƪġ=ǂ2ǮƜƉ'ŜƸ=Ď9# úįªŒŒň'ǂ¶=Ď9"ǌƦ%ĄÖ&%9#ą:9

&$
Andersen, R.A. 2004. Biology and systematics of heterokont and haptophyte algae. Am. J. Bot. 91:1508-1522.
Archibald, J.M. 2007. Nucleomorph genomes: structure, function, origin and evolution. Bioessays 29:
392-402.
Archibald, J.M. 2009. The puzzle of plastid evolution. Curr. Biol. 19: R81-R88.
Baldauf, S.L. 2008. An overview of the phylogeny and diversity of eukaryote. J. Syst. Evol. 46: 263-273.
Burki, F., Shalchian-Tabrizi, K., Minge, M., Skjæveland, Å., Nikolaev S.I., Jakbson, K.S. & Pawlowski J.
2007. Phylogenetic reshuffles the eukaryotic supergroup. PlosONE 2: e790.
Cavalier-Smith, T. 2004. Chromalveolate diversity and cell megaevolution: interplay of membranes,genomes and cytoskeleton. In: Hirt, R.P. & Horner D.S. (eds.) Organelles, Genomes and Eukaryote
Phylogeny, An Evolutionary Synthesis in The Age of Genomics. pp. 75-108. CRC Press, Boca Raton,London, New York, Washington, D.C.
Cerino, F. & Zingone, A. 2007. Decrypting cryptomonads: a challenge for molecular taxonomy. In: Brodie,
J. & Lewis, J. (eds.) Unravelling The Algae, The Past, Present, and Future of Algal Systematics. Pp.
197-214. CRC Press, Boca Raton, London, New York, Washington, D.C.

S. Uwai (A) - 5

Hackett, J.D., Yoon, H.S., Li, S., Reyes-Prieto, A., Rummele, S.E. & Bhattacharya, D. 2007. Phylogenomic
analysis supports the monophyly of cryptophytes and haptophytes and the association of rhizaria withchromalveolates. Mol. Biol. Evol. 24:1702–1713.
Hampl, V., Hug, L., Leigh, J.W., Dacks J.B., Lang, B.F., Simpson, A.G.B. & Roger, A.J. 2009.
Phylogenomic analyses support the monophyly of excavata and resolve relationships among eukaryotic‘‘supergroups’’. Proc. Natl. Acad. Sci. USA 106: 3859-3864.
Harper, J.T. & Keeling, P.J. 2003. Nuclear-encoded, plastid-targeted glyceraldehyde-3-phosphate
dehydrogenase (GAPDH) indicates a single origin for chromalveolate plastids. Mol. Biol. Evol. 20:1730-1735.
Harper, J.T., Wanders, E. & Keeling, P.J. 2005. On the monophyly of the chromalveolates using a
six-protein phylogeny of eukaryotes. Int. J. Sys. Evol. Microbiol. 55:487–496.
´ 2007. ƟǤ 30 ¤ò'ƘńÁƟǤ 7Ƨ9Œňǂ¶|Óŏ|ő×ǯĠĺÚâ­ŇǯŮÞíǯ
Kawachi, M., Inoue, I., Maeda, O. & Chihara, M. 1991. The haptonema as a food-capturing device:observations on Chrysochromulina hirta (Prymnesiophyceae). Phycologia 30: 563-573.
Keeling, P.J. 2009. Chromalveolates and the evolution of plastids by secondary endosymbiosys. J. Eukayot.
Microbiol. 56: 1-8.
Lane, C.E. & Archibald, J.M. 2008. The eukaryotic tree of life: the endosymbiosis takes its TOL. Trends
Ecol. Evol. 23: 268-275.
Li, S., Nosenko, T., Hackett, J.D. & Bhattacharya, D. 2006. Phylogenomic analysis identifies red algal genesof endosymbiotic origin in the chromalveolates. Mol. Biol. Evol. 23: 663-674.
Patron, N.J., Rogers, M.B. & Keeling P.J. 2004. Gene replacement of Furctose-1,6-bisphosphate aldolase
supports the hypothesis of a single photosynthetic ancestor of chromalveolates. Eukaryotic Cell 3:1169-1175.
Patron, N.J., Inagaki, Y. & Keeling P.J. 2007. Multiple gene phylogenies support the monophyly of thecryptomonad and haptophyte host lineages. Curr. Biol. 17: 887-891.
Reeb, V.C., Peglar, M.T., Yoon, H.S., Bai, J.R., Wu, M., Shiu, P., Grafenberg, J.L., Reyes-Prieto, A.,Rümmele, S.E., Gross, J. & Bhattacharya, D. 2009. Interrelationships of chromalveolates within abroadly sampled tree of photosynthetic protists. Mol. Phyl. Evol. 53: 202-211.
Petersen, J., Teich, R., Brinkmann, H. & Cerff, R. 2006. A “Green “ phosphoribulokinase in complex algaewith red plastids: evidence for a single secondary endosymbiosis leading to haptophyes, cryptophytes,
heterokonts, and dinoflagellates. J. Mol. Evol. 62: 143-157.
Reyes-Prieto, A., Moustafa, A. & Bhattacharya, D. 2008. Multiple genes of apparent algal origin suggest
ciliate may once have been photosynthetic. Curr. Biol. 18: 956-962.
Rice, D.W. & Palmer, J.D. 2006. An exceptional horizontal gene transfer in plastids: gene replacement by adistant bacterial paralog and evidence that haptophyte and cryptophyte plastids are sisters. BMC biol. 4:
31.
Riisberg, I., Orr, R.J.S., Kluge, R., Shalchian-Tabrizi, K., Bowers, H.A., Patil, V., Edvarsen, B. & Jakobsen

S. Uwai (A) - 6

K.S. 2009. Seven gene phylogeny of heterokonts. Protist 160: 191-204.
Sanchez-Puerta, M.V., Bachvaroff, T.R. & Delwiche, C.F. 2007. Sorting wheat from chaff in multi-gene
analyses of chlorophyll c-containing plastids. Mol. Phyl. Evol. 44: 885-897.
Sanchez-Puerta, M.V. & Delwiche, C.F. 2008. Minireview: a hypothesis for plastid evolution inchromalveorates. J. Phycol. 44: 1097-1107.
ƞŤÞƑ|ŞëƓº 2005. ¬ũƟ& 9KKvQöć#§ÃćǯŸůǇǫƌǬĺĸŒÌž'UA]oGQŹǱï ĺĸŒň'ĭƖŒÌ(ĺ&$ ǅā!9 ǯpp 374-392. ĠĺÚâ­ŇǯŮÞíǯ
Simpson, A.G.B. & Patterson, D.J. 2006. Current perspective on high-level grouping of protists. In: Katz,L.A. & Bhattacharya, D. (eds.) Genomics and Evolution of Microbial Eukaryotes. pp. 7-30. Oxford
University Press, New York.
Takishita, K. & Uchida, A. 1999. Molecular cloning and nucleotide sequence analysis of psbA from thedinoflagellate: origin of the dinoflagellate plastid. Phycol. Res. 47: 207-216.
Tyler, B.M., Tripathy, S., Zhang, X., Dehal, P., Jiang, R.H., Aerts, A., Arredondo, F.D., Baxter, L.,
Bensasson, D., Beynon, J.L., Chapman, J., Damasceno, C.M., Dorrance, A.E., Dou, D., Dickerman,A.W., Dubchak, I.L., Garbelotto, M., Gijzen, M., Gordon, S.G., Govers, F., Grunwald, N.J., Huang, W.,
Ivors, K.L., Jones, R.W., Kamoun, S., Krampis, K., Lamour, K.H., Lee, M.K., McDonald, W.H., Medina,M., Meijer, H.J., Nordberg, E.K., Maclean, D.J., Ospina-Giraldo, M.D., Morris, P.F., Phuntumart, V.,
Putnam, N.H., Rash, S., Rose, J.K., Sakihama, Y., Salamov, A.A., Savidor, A., Scheuring, C.F., Smith,B.M., Sobral, B.W., Terry, A., Torto-Alalibo, T.A., Win, J., Xu, Z., Zhang, H., Grigoriev, I.V., Rokhsar,D.S. & Boore, J.L. 2006. Phytophthora genome sequences uncover evolutionary origins andmechanisms of pathogenesis. Science 313: 1261-1266.
Yoon, H.S., Hackett, J.D., Pinto, G. & Bhattacharya, D. 2002. The single, ancient origin of chromist plastids.
Proc. Natl. Acad. Sci. USA. 99: 15507-15512.
Zhang, Z. Green B.R. & Cavalier-Smith T. 1999. Single gene circle in dinoflagellate chloroplast genome.
Nature 400: 155-159.

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