   

+$(1'- Symbiodinium "+$ 0 Ï¤ĄsĔŒĺÈîŨÀÈƚÀÈqelpK]fVHRi[VXĔŒį°ƢƏçńĀÆźš 
Eiichi Shoguchi
Genome evolutions in symbiotic dinoflagellate Symbiodinium and parasitic apicomplexans
Key words: alveolates, Apicomplexa, comparative genomics, gene duplication, Symbiodinium
Marine Genomics Unit, Okinawa Institute of Science and Technology
1919-1 Tancha, Onna-son, Okinawa, 904-0495 Japan
DOI: 10.24480/bsj-review.8c4.00123

2   AmcEkrT&İĆģĞ*À(Imrb)+ƬĚƟđťƤƬA`LpbnHNƤƬŕđŦƤ|ūImrb&$©0;:ĚƟđťƤ)+Ƭ7 2,500 ļĲ8;$9Ƭģ5Ìģ& " ċ (ģėċÜŮÍ;:ƪHoriguchi 2015ƫ*Ńö+ŢŎƪ¦ìŜ
ƫ>ï"$ :sø%ƬĲ8;$ :A`LpbnHN+ņśŋÎÌģæ% 9ƬŕđŦ+*¿àÓĄƧæ% :ƪƩƔŗ 2010ƫ ÿōų%+ƬĎxPrJpNr*Ī¹ß)3$K]gůāŧ=; ƬģæĚƟđť Symbiodinium *K]gůŴ)# $Ň:
0 A`LpbnHNƤ*ÃÂŖHohkƤ*K]g&*Đž)79Ŭ$ ƬÌģæA`L
pbnHN*K]gƉ*ĉū>ųù:eklA Ŧ>©2A`LpbnHNK]g%+Ƭ¿*ƎÅoR"$9ƬƎÅƓŪ(')7:ƎÅ*Ġá+Ñ( ;%+ƬNpMŚŢņś)ģ: Symbiodinium *K]g%+'*7 (&ż"$ *% <Ʈ3Symbiodinium  +)8 ĚƟđť* Symbiodinium ÓƪSymbion)ģ:+ dinos ::­ƫƪFreudenthal et al. 1962ƫ+ĘĒ´)ģè
ƬŢŎ>ï"$ :ƪ¯ 1ƫ
ģæ% : Symbiodinium +Ƭċ (Ğ8
Ŭ#"$$9ƪPochon et al. 2014,¯ 2ƫ
ƬNpM*ģť&$ûƁ7ĵľ;$$ :ƪYamashita & Koike2015ƫ

E. Shoguchi-1       ,ûƁ)("$Ƭ^HWlA('>ðƦ:ļ>©?%9ę¦
ĄƧæ% :&ĶŲ;$ :ƪJeong et al. 2014ƫ8)^HWlA&¶Ƨ:&ĴĜ:&¸ª('4(;$$ :
ƪFrommlet et al.
2015ƫƬüæģď*ŮÍ+¸ª;$ (ƪThornhill et al.
2017ƫSymbiodiniumÓ+Ƭġģ*ĚƟđ
ť*{%+Ɖī)+ƬĐžīûƁ)
Ô Imrb%:ƪJanouškovec et al.
2017ƫ + 1
ļƤ&ř8;$ƪKevin et al. 1969ƫƬÅłŊÈīůā)79|ūHnrY A 8 Iƪ¯2ƫ)Ƥ;$ :ƪPochon & Gates 2010ƫ
(8Ƭ*HnrY*¿ċæ)# $+ŹŶŌ $ :ƪYamashita & Koike 2015ƫ
* Symbiodinium *K]g+ƬĚƟđť*{%+ĐžīÐ(ĆK]g>ï#&¸ª;$ :ƪLaJeunesse et al. 2005ƫ*{%4ğ)Ð
(HnrY B * Symbiodinium minutumƪLajeunesse et al. 2012ƫ*K]gƬû)ůŴ; 
4 Symbiodinium minutum ĚƟđť*ĆK]g*İĆģĞK]g)18;( i[rH(ĆK]gĊƇ>4#&+Ƭ8¸ª;$ƪDodge, 1965; Lin, 2011; Wisecaver and Hackett 2011ƫ
(8ƬK]gNBS*ÃÂŖ&$Ĳ8;$
Gbpƫ&A`LpbnHN(')Đ/ƞØ)À ƪ1.5-250

&('4 9ƬPjVXFpPrHDpPpIĕ)7:K]gůŴ+Ƭ*AmcEkrT)Đ/Ɗ;$ƪEisen et al. 2006; Gardner et al. 2002ƫ2013 Ú)í *Imrb+

ĎxPrHDpNr& S. minutumƪą: NIES-3808ƫ>ĥ $ƬĚƟđťƤ%û*ĆK]gĉūƑ>ēÉ ƪũ 1; Shoguchi et al. 2013ƫ¿*~ê¾*ŉĂ*{*s#+Ƭƛò¦ƎÅ§¨)z?% :&4*% " ƪ¯ 3ƬKoyanagi et al. 2013ƫ
0 ƬBpXoplVU(ƎÅƞØ)¿ u, u*RbkBRNBX =6: GT-AG mrm)
E. Shoguchi-2      

à=( &ƬĆTp_H*ƎÅa@flr%+ ĆģĞ&İĆģĞ*yø*TBb*4*
LrY;Ƭ;8*ƎÅö+¼$ :&ù8)(" ƪShoguchi et al. 2013ƫ*7 (i[rH(K]gĊƇĚƟđť*ƉƋĻ) $'*TBfpI%Ġá;*>ŵ/$ &+ƬİĆģĞ*ĆK]gƉ>ř:u%Ɠū% :7 )å=;:ũ 1 ĚƟđťƤƬHohkƤƬA`LpbnHNƤ*ĆK]g*ĉū ēÉ

GCTp_ƎÅ

9BpXoƎÅ

ƛòƎ

K]g*

©ƕ

HŻ>

*Ù²BpXp* 5'ƐƘơ´Å*ōº·ö(%)LrY

opö (Bp*Ƒ*Ù²§ø¨:Ǝ

XopnR%)(Mb)

Ɨ (bp) æÅöĚƟđťƤ
S. kawagutii

935

44

36,8503 (36)

GT/GC17,888

9

S. minutum

616

44

41,92519 (5) GT/GC/GA

2,064

9

S. microadriaticum

808

51

49,10921 (2) GC/GT/GA

3,633

9

Chromera velia

194

49

26,1122 (24)

n/a

989

n/a

Vitrella brassicaformis

73

58

22,817

6 (5)

n/a

92

n/a

23

19

5,268

2 (46)

GT1694(HohkƤ

A`LpbnHNƤ
Plasmodium falciparum

Aranda et al. (2016), Gardner et al. (2002), Lin et al. (2015), Shoguchi et al. (2013), Woo et al. (2015)>Ýĥ 0 ĚƟđť%+ŢŎK]g*ĊƇ4ƥ½=9% :S. minutum *ŢŎ+Ƭ14 TBb*
f[NrHm DNAƪ1.8-3.3 kbpƫ>ï"$9Ƭ;;)ƭ#*ƎÅLrY;$ :*ŽĤĞ+ 9 TBb* RNA ŐƜ>£:&)79ƬčŜ>ï#Tp_HŻ.&ŘŰ;$ :ƪMungpakdee et al. 2014ƫ**ŢŎTp_HŻ+/$Ć)LrY;$9, *{%û4ö*¼$ :£ÊTp_HŻ*ƎÅ+ 145  9, HnrY B * Symbiodinium*łŊ%¿ċ$$ :ƪMaruyama et al. 2015ƫ
;8*&+ƬSymbiodinium *ŢŎ+û4¿ċ$ :EmF\k*s#%Ƭ*ĩġŵŁ+Ģ»ƍä)ğ)Ɠū(*%+Ʈ&&>ê:ƪXiang et al. 2015ƫ sø%fXLpYl
A*K]g+ƬLrY;$ :ƎÅ&E. Shoguchi-3      ĝ) $eklA Ŧ Plasmodium falciparum & S. minutum &*Ƙ%Ç;$ :*{)+čŜþĲ* small RNA ƎÅ©0;:ƪShoguchi et al. 2015ƫí *Imrb%+Ƭ
Symbiodinium *Ë|*s#% :NpM Acropora digitifera *K]g>ůŴ$9ƪShinzato etal. 2011ƫƬK]gé¸>·ĭ&$ƬĚƟđť&NpM*ģƙ>ù8):ĵľ>Ɖ3$ :ƪShinzato et al. 2014ƫ
5Symbiodinium  !&$
 HnrY B * S. minutum *K]g)Ō $ƬHnrY F * S. kawagutii *K]gůŴ;ƪũ
1; Lin et al. 2015ƫ
ƬS. minutum )+18;("S. kawagutii K]g*ğâ #¸ª;$:, S. kawagutii K]g%+ƬƅŽƒŅ*ƎÅ*öû4¼ÀƬBpXop*(ƎÅ*¦Đžī¿ ƪũ 1ƫ*&+ƬƎÅ*Ġá*(9*ƐƬmRNA 8ƅŽƒŅ)7"$#8;

cDNA K]g)÷ )ň1ƀ0;:&)79ż"$ :&>ķ¬: ƪSlamovits and Keeling, 2008ƫ
0Lin 8ƪ2015ƫ+ƬS. kawagutii * microRNAƪmiRNAƫ*Ç±)# $űŵ/$9ƬmiRNA )7:ŵŁ>£$  (ƎÅŖ)ıĮ$ :NpMƎÅ&*ƑĐž)79ƬSymbiodinium * miRNA NpM A. digitifera*XkpRdrTrƎÅ('*ĩġŵŁ)ƙ=:¥Ŝæ>ƃ/$ :ƪLin et al. 2015ƫ
ßƬ
Symbiodinium * miRNA NpMņś)'*7 )ſƄ;$ :*Ƭ0 +*¢Îø¨.*ſƄ4 :*>ŵ/$ ãū :&å=;: Ō $ 2016 Ú)+Ƭý)Ô&ř8;:HnrY A * S. microadriaticum K]gůŴ; ƪAranda et al. 2016ƫ
HnrYA +*HnrY81:& ÄīImrb&ř8;Ƭ&+7 5000 tÚ)Ô &óÉ;$ :ƪPochon et al. 2006ƫ
*K]g¡-HnrY B & F *K]g>©3Ƭ*İĆģĞK]g&*Đž(;$ :*Đžůā*{%4ƬSymbiodinium K]g*ğâ*s#+ƎÅƓŪ% :&&Þŵ;:8) Aranda 8ƪ2016ƫ+ƬļğĨī)¿ċ

Symbiodinium *XkpRdrTrƎÅŖƬË|&*ģƙ)+Ɠū(*%+( &&>ķ¬:ŉĂ>¸ª$ :ƓŪ ƎÅ¿ &ƎÅ§¨)z?% :&ŉĂ&)Ƭƛ

ğâ+Ƭ3 ļ* Symbiodinium K]g) $ĶŲ;:

%+8)ÖÀ(K]g>4#ŞĦģėæ*ĚƟđť%+Ƭ*ğâ+'*ĻÛ18;:*%< Ʈ*K]gĊƇ&ģėċÜ)+'*7 (ƙ :*!< Ʈ÷ŭ*K]gƑé¸*ţĽ)79ĚƟđťK]g*¿ċæ&Ɖ*űņ00ù8)("$:!<ƪJanouškovec et al. 2017ƫ6"+$  0 
 ĚƟđťƤ*|ū(ÃÂŖ% :A`LpbnHN+ƬÌģæ% 9¦ìŜ>ï"$ (5000 ļuĲ8;Ƭ
100 tļu :&óě;$ :ƪAdl et al. 2007; Pawlowski et al. 2012ƫ
*A`LpbnHN)Ɓő(Imrb%¦ìŜ>ï#HohkƤ 2 ļ*K]gůŴ;ƪWoo et al. 2015ƫ
*HohkƤ*K]g>ACXImrbƪĹ³ƫ&$Đžůā>ŧ
&)79ƬA`LpbnHN*ƆĹ+ƬƞØ)¿*ƎÅƪ7 3800ƫ>ôÁ&¥Ŝæķ¬;$ :;8+¦ì5RWorm¦ìł& " Ÿł*oR)Ä
E. Shoguchi-4      09ƬA`LpbnHN%+ŞĦģė)ãū(Ÿł*ƒŅƎÅ5ņśŝſƄł*ƎÅ>;;*łŊ%Á"$ :(8ƬsƐ*Tp_HŻ*Ǝ
Å*ö+Ƭ :Ɖ*TBfpI
&)¼ 7 % :Woo 8
ƪ2015ƫ+ƬRNA ŉ¦Tp_HŻƪÌģŜ>Ġá:*RWrQ 1ƫƬ
ņśƨćłTp_HŻƪHohkƤ&Ôß*ÌģŜ>Ġá:Ƙ*RWrQ 2ƫ
Ƭņś¾Tp_HŻ&ApiAP2 Ž®ÅƪÌģŜ>Ġáß*RWrQ 3ƫ*Ơ)¿ċż" ¥Ŝæ>ķ$ :ƪ¯ 4ƫğ)ņśƨćł*ƟđTp_HŻƎÅ*ƉƬņś)"$ Ŝ>Ġá:u%*GrdBpX!" *%+( &óě$ :

7 ĐžK]gůā)79ƬÌģæA`LpbnHNƤĞņś)Ìģ:Ŝ>Ġá:ƉƋĻ%+Ƭ¦ìŜ5ŸłƎÅŖ*ÀŭČ(oR)ŌƬÌģŜ>Ġá:TBfpI):ƟđĊƇƙƈƎÅ*ƉƖ&(" *%+( &ŹŶ;:K]gNBS*œÐ4ż"$ 7 % :sø% Symbiodinium Ó*K]g%+ƎÅa@flr*oR79
4Ƭ2<ƎÅƓŪƣŔ)ż"$ :7 %ƬK]gNBS+0!(9À ƓŪƎÅ¿ &&ģėċÜ)+'?(ƙ :*!< ƮůŴ;$

Symbiodinium+Đžī¶Ƨ5 ļ% :Ƭ¶Ƨ*Ɲ ļ*K]g+Ă $' ("$ :*% <Ʈüæģď)ƙ=:¥Ŝæ* :ƎÅŖ Symbiodinium K]g)LrY;$ :&+¸ª;$ :Ƭüæģďú)+İĆģĞ8 ăŠĊƇ>ï#*% < ('Ƭş«+Ò( ĚƟđť+K]gƉ!%(ņśģĞÈī)4ŷ*¿ ģĞ% 9ƬA`LpbnHN*7 )K]g8*AborU&ň1¦=:&Ƭ*ŷ)Ƃ:ƋĻ%, 7 ­9ƌ&(:7 )å:*% :./ ĚƟđťK]g*ĵľ>ŧ u%ƬelpK]fVHRi[VX*Ťĳŧõñ>+3RTVa*Ĭċ)ĸħu0*µŀ*č >w$v" ƬÙÕĖ}ģ¡-ErFZBOr*Ĭċ)ëŸħu0#,%*Adl, S.M., Leander, B.S., Simpson, A.G., Archibald, J.M., Anderson, O.R., Bass, D., Bowser, S.S.,
E. Shoguchi-5      

Brugerolle, G., Farmer, M.A., Karpov, S., Kolisko, M., Lane, C.E., Lodge, D.J., Mann, D.G., Meisterfeld,R., Mendoza, L., Moestrup, Ø., Mozley-Standridge, S.E., Smirnov, A.V., & Spiegel, F. 2007. Diversity,nomenclature, and taxonomy of protists. Syst. Biol. 56: 684-689.
Aranda, M., Li, Y., Liew, Y.J., Baumgarten, S., Simakov, O., Wilson MC, Piel, J., Ashoor, H.,
Bougouffa, S., Bajic, V.B., Ryu, T., Ravasi, T., Bayer, T., Micklem, G., Kim, H., Bhak, J., LaJeunesse,TC., & Voolstra, CR. 2016. Genomes of coral dinoflagellate symbionts highlight evolutionary
adaptations conducive to a symbiotic lifestyle. Sci. Rep. 6: 39734.
Dodge, J.D. 1965. Chromosome structure in the dinoflagellates and the problem of the mesocaryotic cell.
Excerpta Med. Int. Congr. Ser. 91: 339-345.
Eisen, J.A., Coyne, R.S., Wu, M., Wu, D., Thiagarajan, M., Wortman, J.R., Badger, J.H., Ren, Q., Amedeo,P., Jones, K.M., Tallon, L.J., Delcher, A.L., Salzberg, S.L., Silva, J.C., Haas, B.J., Majoros, W.H., Farzad,M., Carlton, J.M., Smith, R.K.Jr., Garg, J., Pearlman, R.E., Karrer, K.M., Sun, L., Manning, G., Elde,N.C., Turkewitz, A.P., Asai, D.J., Wilkes, D.E., Wang, Y., Cai, H., Collins, K., Stewart, B.A., Lee, S.R.,
Wilamowska, K., Weinberg, Z., Ruzzo, W.L., Wloga, D., Gaertig, J., Frankel, J., Tsao, C.C., Gorovsky,M.A., Keeling, P.J., Waller, R.F., Patron, N.J., Cherry, J.M., Stover, N.A., Krieger, C.J., del Toro, C.,
Ryder, H.F., Williamson, S.C., Barbeau, R.A., Hamilton, E.P., & Orias, E. 2006. Macronuclear genome
sequence of the ciliate Tetrahymena thermophila, a model eukaryote. PLoS Biol. 4: e286.
Freudenthal, H.D. 1962. Symbiodinium gen. nov. and Symbiodinium microadriaticum sp. nov., a
Zooxanthella : Taxonomy, life cycle, and morphology. J. Protozool. 9: 45-52.
Frommlet, J.C., Sousa, M.L., Alves, A., Vieira, S.I., Suggett, D.J., & Serodio, J. 2015. Coral symbiotic
algae calcify ex hospite in partnership with bacteria. Proc. Natl. Acad. Sci. U S A. 112: 6158-6163.
Gardner, M.J., Hall, N., Fung, E., White, O., Berriman, M., Hyman, R.W., Carlton, J.M., Pain, A., Nelson,K.E., Bowman, S., Paulsen, I.T., James, K., Eisen, J.A., Rutherford, K., Salzberg, S.L., Craig, A., Kyes,S., Chan, M.S., Nene, V., Shallom, S.J., Suh, B., Peterson, J., Angiuoli, S., Pertea, M., Allen, J., Selengut,J., Haft, D., Mather, M.W., Vaidya, A.B., Martin, D.M., Fairlamb, A.H., Fraunholz, M.J., Roos, D.S.,Ralph, S.A., McFadden, G.I., Cummings, L.M., Subramanian, G.M., Mungall, C., Venter, J.C., Carucci,
D.J., Hoffman, S.L., Newbold, C., Davis, R.W., Fraser, C.M., & Barrell, B. 2002. Genome sequence ofthe human malaria parasite Plasmodium falciparum. Nature 419: 498-511.
Horiguchi, T. 2015. Diversity and phylogeny of marine parasitic dinoflagellates. In: Ohtsuka, S., Suzaki, T.,
Horiguchi, T., Suzaki, N., & Not, F. (eds.) Marine Protists. pp. 397-419. Springer, Japan.
Janouškovec, J., Gavelis, G.S., Burki, F., Dinh, D., Bachvaroff, T.R., Gornik, S.G., Bright, K.J., Imanian, B.,
Strom, S.L., Delwiche, C.F., Waller, R.F., Fensome, R.A., Leander, B.S., Rohwer, F.L., & Saldarriaga,J.F. 2017. Major transitions in dinoflagellate evolution unveiled by phylotranscriptomics. Proc. Natl.
Acad. Sci. U S A. 114: E171-E180.
Jeong, H.J., Lee, S.Y., Kang, N.S., Yoo, Y.D., Lim, A.S., Lee, M.J., Kim, H.S., Yih, W., Yamashita, H., &LaJeunesse, T.C. 2014. Genetics and morphology characterize the dinoflagellate Symbiodinium voratum,n. sp., (Dinophyceae) as the sole representative of Symbiodinium clade E. J. Eukaryot. Microbiol. 61:75-94.
E. Shoguchi-6      

Kevin, M.J., Hall, W.T., McLaughlin, J.J.A., & Zahl, P. A. 1969. Symbiodinium microadriaticum
Freudenthal, a revised taxonomic description, ultrastructure. J. Phycol. 5: 341-350.
Koyanagi, R., Takeuchi, T., Hisata, K., Gyoja, F., Shoguchi, E., Satoh, N., & Kawashima, T. 2013.
MarinegenomicsDB: an integrated genome viewer for community-based annotation of genomes. Zoolog.
Sci. 30: 797-800.
Lajeunesse, T.C., Parkinson, J.E., & Reimer, J.D. 2012. A genetics-based description of Symbiodinium
minutum sp. nov. and S. psygmophilum sp. nov. (Dinophyceae), two dinoflagellates symbiotic withcnidaria. J. Phycol. 48: 1380-1391.
LaJeunesse, T.C., Lambert, G., Andersen, R.A., Coffroth, M.A., & Galbraith, D.W. 2005. Symbiodinium(Pyrrhophyta) genome sizes (DNA content) are smallest among dinoflagellates. J. Phycol. 41: 880-886.
Lin, S. 2011. Genomic understanding of dinoflagellates. Res. Microbiol. 162: 551-569.
Lin, S., Cheng, S., Song, B., Zhong, X., Lin, X., Li, W., Li. L., Zhang. Y., Zhang, H., Ji, Z., Cai, M., Zhuang,Y., Shi, X., Lin, L., Wang, L., Wang, Z., Liu, X., Yu, S., Zeng, P., Hao, H., Zou, Q., Chen, C., Li, Y.,Wang, Y., Xu, C., Meng, S., Xu, X., Wang, J., Yang, H., Campbell, DA., Sturm, NR.,
Dagenais-Bellefeuille, S., & Morse, D. 2015. The Symbiodinium kawagutii genome illuminates
dinoflagellate gene expression and coral symbiosis. Science 350: 691-694.
Maruyama, S., Shoguchi, E., Satoh, N., & Minagawa, J. 2015. Diversification of the light-harvesting
complex gene family via intra- and intergenic duplications in the coral symbiotic alga Symbiodinium.
PLoS One 10: e0119406.
Mungpakdee, S., Shinzato, C., Takeuchi, T., Kawashima, T., Koyanagi, R., Hisata, K., Tanaka, M., Goto,H., Fujie, M., Lin, S., Satoh, N., & Shoguchi, E. 2014. Massive gene transfer and extensive RNA editingof a symbiotic dinoflagellate plastid genome. Genome Biol. Evol. 6: 1408-1422.
Pawlowski, J., Audic, S., Adl, S., Bass, D., Belbahri, L., Berney, C., Bowser, S.S., Cepicka, I., Decelle, J.,
Dunthorn, M., Fiore-Donno, A.M., Gile, G.H., Holzmann, M., Jahn, R., Jirků, M., Keeling, P.J., Kostka,M., Kudryavtsev, A., Lara, E., Lukeš, J., Mann, D.G., Mitchell, E.A., Nitsche, F., Romeralo, M.,Saunders, G.W., Simpson, A.G., Smirnov, A.V., Spouge, J.L., Stern, R.F., Stoeck, T., Zimmermann, J.,
Schindel, D., & de Vargas, C. 2012. CBOL protist working group: barcoding eukaryotic richness beyond
the animal, plant, and fungal kingdoms. PLoS Biol. 10: e1001419.
Pochon, X., & Gates, R.D. 2010. A new Symbiodinium clade (Dinophyceae) from soritid foraminifera in
Hawai'i. Mol. Phylogenet. Evol. 56: 492-497.
Pochon, X., Montoya-Burgos, J.I., Stadelmann, B., & Pawlowski, J. 2006. Molecular phylogeny,
evolutionary rates, and divergence timing of the symbiotic dinoflagellate genus Symbiodinium. Mol.
Phylogenet. Evol. 38: 20-30.
Pochon, X., Putnam, H.M., & Gates, R.D. 2014. Multi-gene analysis of Symbiodinium dinoflagellates: aperspective on rarity, symbiosis, and evolution. PeerJ 2: e394.
Shinzato, C., Mungpakdee, S., Satoh, N., & Shoguchi, E. 2014. A genomic approach to coral-dinoflagellate
symbiosis: studies of Acropora digitifera and Symbiodinium minutum. Front Microbiol. 5: 336.
Shinzato, C., Shoguchi, E., Kawashima, T., Hamada, M., Hisata, K., Tanaka, M., Fujie, M., Fujiwara, M.,E. Shoguchi-7      

Koyanagi, R., Ikuta, T., Fujiyama, A., Miller, D.J., & Satoh, N. 2011. Using the Acropora digitifera
genome to understand coral responses to environmental change. Nature 476: 320-323.
Shoguchi, E., Shinzato, C., Hisata, K., Satoh, N., & Mungpakdee, S. 2015. The large mitochondrial
genome of Symbiodinium minutum reveals conserved noncoding sequences between dinoflagellates andapicomplexans. Genome Biol. Evol. 7: 2237-2244.
Shoguchi, E., Shinzato, C., Kawashima, T., Gyoja, F., Mungpakdee, S., Koyanagi, R., Takeuchi, T., Hisata,K., Tanaka, M., Fujiwara, M., Hamada, M., Seidi, A., Fujie, M., Usami, T., Goto, H., Yamasaki, S.,Arakaki, N., Suzuki, Y., Sugano, S., Toyoda, A., Kuroki, Y., Fujiyama, A., Medina, M., Coffroth, M.A.,
Bhattacharya, D., & Satoh, N. 2013. Draft assembly of the Symbiodinium minutum nuclear genome
reveals dinoflagellate gene structure. Curr. Biol. 23: 1399-1408.
Slamovits, C.H., & Keeling, P.J. 2008. Widespread recycling of processed cDNAs in dinoflagellates. Curr.
Biol. 18: R550-552.
ƩƔŗ 2010. AmcEkrT)Ŭ8;:¿ċ(ģø~ğ)ĚƟđťƤ)# $~. ĈĞĺÈ*ûŏ Ŀ 1 × pp. 11-15.
Thornhill, D.J., Howells, E.J., Wham, D.C., Steury, T.D., & Santos, S.R. 2017. Population genetics of reef
coral endosymbionts (Symbiodinium, Dinophyceae). Mol. Ecol. : doi: 10.1111/mec.14055.
Wisecaver, J.H., & Hackett, J.D. 2011. Dinoflagellate genome evolution. Annu. Rev. Microbiol. 65:369-387.
Woo, Y.H., Ansari, H., Otto, T.D., Klinger, C.M., Kolisko, M., Michalek, J., Saxena, A., Shanmugam, D.,
Tayyrov, A., Veluchamy, A., Ali, S., Bernal, A., del Campo, J., Cihlář, J., Flegontov, P., Gornik, S.G.,
Hajdušková, E., Horák, A., Janouškovec, J., Katris, N.J., Mast, F.D., Miranda-Saavedra, D., Mourier, T.,Naeem, R., Nair, M., Panigrahi, A.K., Rawlings, N.D.,Padron-Regalado, E., Ramaprasad, A., Samad, N.,Tomčala, A., Wilkes, J., Neafsey, D.E., Doerig, C., Bowler, C., Keeling, P.J., Roos, D.S., Dacks, J.B.,
Templeton, T.J., Waller, R.F., Lukeš, J., Oborník, M., & Pain, A. 2015. Chromerid genomes reveal theevolutionary path from photosynthetic algae to obligate intracellular parasites. Elife 4: e06974.
Xiang, T., Nelson, W., Rodriguez, J., Tolleter, D., & Grossman, A.R. 2015. Symbiodinium transcriptome
and global responses of cells to immediate changes in light intensity when grown under autotrophic ormixotrophic conditions. Plant J. 82: 67-80.
Yamashita, H., & Koike, K. 2015. Biology of symbiotic dinoflagellates. In: Ohtsuka, S., Suzaki, T.,Horiguchi, T., Suzaki, N., & Not, F. (eds.) Marine Protists. pp. 421-439. Springer, Japan.

E. Shoguchi-8   