AKV'9Z xzqtql mrpqwnl



!

O46F

3PZěÒ® ŶȗȖǣ ŏŖĉǤļǚƍÁƧƲåȳPDȴŚĀĉĀĉȞōȨôÅĵƫĉƧƲƫµƴƕãƫĉēń
277-8562Îǟƣŝěŝ/ǟ 5-1-5
ȳ* ƒĶĕȸÌŻȍĀĉÌŎƕƌðoJ}gƫĉ]_ųð[dYzĐǢǘŻďȰĶ 051-0013 ĐǢěǛǫƙ 1 ơ 133-31ȴ
Apomixis influence adaptaion strategy of green seaweed Ulva prolifera.
Key words: Apomixis, asexual reproduction, sex determination, sexual reproduction, Ulva
Kensuke Ichihara1,2 and Shigeyuki Kawano2
1
Research Fellow of Japan Society for the Promotion of Science2
Department of Integrated Biosciences, Graduate School of Frontier Sciences,
The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa, Chiba 277-8562, Japan( * Current address: Hokkaido University, Field Sciencee Center for Northern Biosphere,
Muroran Marine Station, 1-133-31, Funami-cho, Muroran, Hokkaido, 051-0013, JAPAN)
DOI: 10.24480/bsj-review.8c3.00122

SR<A

95D=@5D= ÿ/ƕƌ0Ɯ,Bª£Ýü/ț)ȑćŁF,ȶćĊFŮŔİƕůF,Ŏ)ÏƏ)Ǚ¾*Ýȑć]bfFĹ%"ćĊFŮƆİƕůF,ñBƕůŎŸ-0w|bf*ew|bfAǼ(ȔƢķFľĭǪ/ýÊ-ĥ*ǐ@C(

ƕƌ=ĈAȶŔİƕů)099,ƔûǪëF

BŎ)ȶȑƞ,ÿŨİF©&*)B/)Ɣû

BƆİƕů)0ÏƏ)/ǌůØǖ,/)ȱ

ȶȑƞÿŨİF©&*)ȶƔû/ýÊ-0Ĥ*ǐ@C(

ǌůÆF=&

B ȶŔ

İƕů*Ɔİƕů/ďȠ/ǙƇƔû)/ȐĮƞĳǏFǶǵB"<-0ȶȂǋƯȶ)C1ÝƯº)Ŕİƕůª£*Ɔİƕůª£Ĉñ(B*ĭǪ) ȟţƌ)0IsuSY[*â1CBƆİƕů/ŨĢİƕů*(Ʀ@Cȶ40 ĕ 400 ƯȢİȔ°£)CF,B

AȶC0ƯćħĵF"ƆȳSchmidt et al. 2015ȴ ȟţƌ/

Bǔè0ȶȆĝ/ŔİƕůȌƮ)0ȶǔ/ŰƿǕſň¾ǦȶǔèƿǕ

*,AȶŢ¾ǦFǃ(ÑƿǕȶĬƿǕȶÉƿǕFà;ǔèħĵCȶÑƿǕ*ĬƿǕ5ȖǨÖƺĵƳB*)ǔƝƕȉ;Ŏ)IsuSY[)0ƑĬ/ƿǕÏƅƝƕFBIs[s|*ȶǔ/ŰƿǕ)ſňF

E,

ſň¾Ǧ¸ȳIstKOY[ȴAȶ2N /ȢİȔ°£ÏƅƝƕBeJq[s|B*Ʀ@C(

B

ȳBicknell and Koltunow 2004; Sharbel et al. 2010; Schmidt et al. 2015ȴ
K. Ichihara & S. Kawano- 1{}|o~~lxzqtqlmrpqwnl

AKV'9Z xzqtrl mrpqwnl

TRJK?L/D>'

l ƪ"$ƧƲŗŋ*(BIOXĕ0ĀóŻǡȬ/ț)ȶIOXǡǈ-à9C(

ȳvan den Hoek et al. 1995ȴ,ƕǓô0=$DGŷėô)9)ƕǓƸíFğ(B

Bȶ-0ŵųô>Žųô-BƯ=à9C(AȶŨ,ųƔû-ȐĮ"T}q)

B*ǐ

@CBȳShimada et al. 2008; Ichihara et al. 2009; Mareš et al. 2011ȴ9"/Ɣû5/ȐĮǖÆ/ȱ@ȶ11ĀȘ-ǌǞȶŷėô>Żė-÷Ʊ(9_Kgȴ FģǻÒëƯ*(=Ʀ@C(ǊƂȳT|BȳHernández et al. 1997; Shimada et al. 2003;

Hiraoka et al. 2004ȴ2008 Ğ/ÌO|nbS/Ƞ-0rf~[ù*,%"ï/ȤĘ)ĀȘƝƕȶŷėôȥǊ-Ş9%"¼ƤøȍC"/FǬ(

B=

B#D

ȳLeliaert et al. 2009; Hiraoka et al. 2011ȴIOXĕ-0ƿǕĖǗƎ-G#_Kq*
ĖaypƎ/_Kqà9C(AȳHayden et al. 2003ȴȶŏŖǲ)0ÃǑFǉƬ(IOXȶĩǑFIOk|*âG)BȳÜƗ 1998ȴIOXĕ0ÝóF,ǥ*(ƕƌĉ/ŇƫŒ-=ÕA@CB*=Bȭä*(ơ-Bū=ÿAȶĂ:ƈ>ƈ"<ȶűȀƞȯŞ:/

BŻǡȬ*

ƕƌ/

1-3A

B#D

î1 .IOk|/ǜƞ,ƕŹÙȳAȴŔİƕůóȶ
ȳBȴ2ŖȦŲ/ƆİƕůóȶȳCȴ4ŖȦŲ/ƆİƕůóFƩ IOXĕ/ƕŹÙFî 1 -9*<"mtȵóȶmtȷó/Ȕ°£0 2 ŖȦŲ)ŭ/Ǻ¶İF=&Ȕ°ćFŅ½ȶCĿÛȶƝƕB*)Ǖć£*,BǕć£@0ſň¾Ǧ?A 4 ŖȦŲİ/ȋǺćŅ½CȶCƝƕB*)ĿÛó/Ȕ°£*,B/?

-ŔİƕůF,ª£)0Ýħ/Ȕ°£*Ǖć£ƕůƿǕF(ǍAȃCB

ƕŹÙF=&9"Ȕ°£@Ņ½C"Ȕ°ć0ĿÛ-āņ"ùÛ-0ÏƅƝƕB*Øǖ)B/ŔİóƕŹÙ-È(ȶƕŹÙ/)ŔİƕůF,EȶƆİ/ȋǺƿǕF(F,ƆİóƕŹÙ/ª£ǎ=ĈñBƍ-IOXĕ)0ÝƯº-Ŕİƕůš*Ɔİƕůš*=-Ĉñ(

BU[ÿȶ

C!C/Ưº)Ə

Ƴ-Ǩňê/ƆİƕůËǻ"*ǐ@CBȳHiraoka et al. 2003a; Hiraoka et al. 2003bȴ/ƆİóƕŹÙF,ª£ǎ-0 2 ȦŲİ/ȋǺćF&B_Kq* 4 ȦŲİ/ȋǺć
K. Ichihara & S. Kawano- 2{}|o~~lxzqtrlmrpqwnl

AKV'9Z xzqtsl mrpqwnl

F&B_KqƦ@C("ȫīƀ¶-?B DNA Ș/ŀč-?Aȶ

Bȶ£ƿǕFƖ

Ŕİƕůóª£/Ǖć£*Ý DNA ȘF=&*E%(Aȶ

C!CǨƢ*ǐ@C(BȳHiraoka et al. 2003bȴ9"ȗþ)Ľȡ"IOk|)ȶVkv- 1 Wn

,

Hsp90 ȑć/Ƶ 6 MR^/ȓ/Ȩô/ZLk_KnTF,%"*DȶƆİª£-=ª£ºÿóFĹ&ª£à9C(
SY[Ƙř)

BØǖİĺłC("*@IOk|/Ɔİƕůª£0IsuBȳOgawa et al. 2015ȴ,@ȶIOk|)

0ȑĉƞ,ƧƲ>VkvǮśǽ(AȶC9)-İ-Ȉș"ȑćġƦ@C(@ȶƆİª£/Vkvŧȇ0E@,

99)

%"

l

UR/

N:7(*BGI

l 2010 Ğ-,%(ŻǡȬ-

;K#Q+

HM

(=¸VkvǮśƟG-,ECB?

-,%(Aȶǩǡ)0YOugȳCock et al. 2010ȴȶWpȳYe et al. 2015ȴȶORhx\SȳNishitsuji etal. 2016ȴȶƽǡ)=[Xmk|ȳNakamura et al. 2013ȴȶckt_/ƯȳCollen et al. 2013ȴ)ǵŊøáC(
0øáC(BIOXǡǈ)=C9)-ufWg|I>ǟǊ£Vkv/Ȕ¿"ȳLeliaert and Lopez-Bautista 2015; Melton et al. 2015ȴȶ¸VkvǮś-&(øá¥,ȊC(
Ư)

BƎĴ)

%"

)ȶƪ"$/ƧƲT}q0ȶIOk|/

B Ulva partita *[ZIOk|/Vkv/ǮśFȉ<ȶȚș/ DNA ŌƋFǳ;*)B¾ćYUXPacBio > Illumina HiSeq *B*)ȶǥ 1 -9*<"?

%"ŬYUXFÂƖ-IOXĕǡȬ)=ǊǡS{ugxh[>Ǌǡr}rbS*

ÝƮĠ/ƺĠ/Vkve_FĪB*-ĵÇ"/Vkve_@ĿÛóț)Ȕ
¿¾Ë>Şǝ£ŧȇ/»ŧĵǱ<@CBȶĿÛóƍƜƞ,Şǝ£ȨôFƝǫ)(B

ȳî 2Aȴǥ 1. Ulva partitaȶǊǡS{ugxh[ȶǊǡr}rbS[/ŢVkve_/űȀ

ƯÞ

Şǝ£ň

[PoN}g

ǉúöň
(Mbp)

ň

œȚ(Mbp) N50 (Mbp)

U. partita (mt-)

n.d.

110

851

10.6

2.6

+

U. partita (mt )

12*

116

1,385

7.3

1.7

C. reinhardtii

17

121

1,557

n.d.

1.6

V. carteri

14

141

1,327

n.d.

1.5

Merchant et al. (2010), Prochnik et al. (2010) FÓƉ
/ĿÛóƍƜƞ,Ȩô-0

C!C/ĿÛó-ƍƜƞ,ȑć/-ȶĿÛó)¹ŔC"Ȕ¿¾Ë"ȑćȳQwfTȴ=ĈñB/Ȩô/ȑćFS{ugxh[>r}rbS[/İŞǝ£Ȩô*űȀ"*DȶǊǡ/İŴčëć*ǐ@C(

B

MID ȑćȳFerris and Goodenough 1997; Umen 2014ȴ-E,@ƢÝİFƩȑćȳUpRWP1ȴǫ&%") UpRWP1 * MID à9CB RWP-RK ȑćoHu|/ȑć)¾ćƼǅŪFŧƹ"*DȶUpRWP1 0 MID *Ý/S~g-à9CȶS{ugxh[>r}rbS[*0İŴč/ūŧƜ,B*ƩæC"/ĿÛóƍƜ
K. Ichihara & S. Kawano- 3{}|o~~lxzqtslmrpqwnl

AKV'9Z xzqttl mrpqwnl

ƞ,ȨôIOXĕ)©ĈC(

B=/Fƨ<B"<-ȶQwfT/& PRA1ȳProliferation-associated protein 1ȴȑć*/Ȩô/þ¯-ĈñB GTB1ȳG-strand
telomere binding protein 1ȴȑćF/IOXĕ/ 7 Ư/ĿÛó/š@Ïȣȶ¾ćƼ
ǅǮśF,%"

/ǄŜȶGTB1 ȑć)0ĦƇȶÝƯº/ĿÛóš)Ȕ¿/¾Ë

07*G+ǫ@C,%"ȶPRA1 ȑć)0Ưº/ĿÛó/šț)Őƨ,Ȕ¿/¾Ë
ƩCȶĿÛó*-9*9%"S~gFħĵ"ȳî 2B, Cȴî2. Ulva partita/ĿÛóƍƜƞŞǝ£Ȩô/ŦƛîȳAȴƢÝ,Ȩô-Ļ9C"ħ)ĈñBĿÛóƍƜƞ
,Şǝ£Ȩô/ũĢîȳBȴȨôº/PRA1ȑć/ƼǅŪȳCȴ Ȩôþ/GTB1ȑć/ƼǅŪ

/ĿÛóƍƜƞ,Ȩô0IOXĕ½ƒ"ő-0ĈñȶĿÛóț)ǂŁFǃȰȶƜ,BȉËȌƮFȁ%"*ǐ@CB/ĿÛóƍƜƞ,Şǝ£Ȩô/Ɲǫ-?%(ȶIOk|)=İŴč>ƆİË*)&&%"İ/ǻƁ*ȉË-&(/ƧƲF,BȑćĔ·Ÿ0 2000 Ğȝ=ƧƲǆ@C(

œȂ-,A PEG ŸFƖ"ȳKakinuma et al. 2009ȴȶ

(ċč"ħǸǿŁ£/¤½=Øǖ-,%(AȳOertel et al. 2015;

Suzuki et al. 2016ȴȶȅȑĉƞ,Ǯś=ĩØǖ-,B#D

Ȅ/Vkvıø/ŉ±*ȑćĔ·ĸǤ/ƨƳ-?%(ȶIOk|=ō",xe}Żǡ*,B#DVR)"&0!

 ŏŖœĩ/žźƌƕǓ(

öƠ

,-



%1

*â1CBéÍę-0dhQMm>ȲFą<*(99,ƕBȶIOk|/țȶ[ZIOk|=ŧĵƯ/&)B[ZIOk|/ƕůó-ƥơȶźô@źô9)/¾ĜFǴ6B*ȶ¢úƃĠ)
8 100%/ÄÛ)Ŕİƕůª£ƕǓ(B/-Ē(ȶúƃĠ/ȱ-ȈC(ȶ2 ŖȦŲİ/ƆİšȳŖư)0§Ďƞ-ƆİȔ°£> 4 ŖȦŲİ/ƆİšȳÝŨ-ƆİǕć£Bźô)07Ŷ×ô-Ȃ'(*â4*-Bȴ*â4ȴ/ÄÛȱ,Aȶ@-úƃĠȱ,BŶ×ô)0Ŕİª£07*G+ĆFżȶ78Ɔİóª£/:ƕǓ(ȳHiraoka and Higa 2016ȴ/¾Ĝ/­A0Ǵş,EC(
C(Aȶ9"œȂ,%"Ǵş)=ÝŨ-ǇĹC("" 2004ȹ2007 Ğ/țǇĹ"ȳî 3ȴ

K. Ichihara & S. Kawano- 4{}|o~~lxzqttlmrpqwnl

AKV'9Z xzqtul mrpqwnl

úƃĠ/¢ź-Ŕİª£ǎȶƮĠ/źô-Ŕİ*Ɔİȶúƃ
ĠȱŻų*ÝƮĠ/Ŷ×ô-0Ɔİª£´Ð(Bȶ/¾

Ĝ/­A0+/?

-ħĵC"/#D¥1ȶƕůó-?%(ȑćƝƒ-ýËƕȶ/ǄŜ¢úƃĠǒİ-Ěƕ(ȶƕǓò-­A½(

B/ȶ

C*=¾

Ĝô/úƃĠ/ĚȶƔû/Ě[f~[*,%([ZIOk|/ƕůó-ĨȧFÔ8ȶ/?

,ƕůó-?BƕǓò/­AFƕ:½(B/#DÝƯº/ƕůó/ȏ-?%(¢úƃĠ-ĒBȐĮĠ-ĚB?

3. 2015,4:2016,4:)¨§¡¤ª¯-ǫB[ZIOk|

*

0ȶƕůó-?BƔû5/ȐĮĠ/çȪFǶǵB/-Ȑ"ţƌ)B/)0,#D@¦®¬OD!

B*

&E U4i

Hiraoka and Higa (2016) H3

ƪ"$/

S`51

.

I

³

Nh

P

T}q0ȶƒñȶ[ZIOk|-:@CB/ƕůó*Ɣû5/ȐĮĠ/Ȝ¨İ-&(ƧƲFȉ<(

WR!

@5D=$#



D

l [ZIOk|/Ɔİƕůª£ǎ0+/?
£@,G@/ȌƮFǃ(½ƒ(-½ƒ(B/#D

B/#D

ȶ

B*D)

B

=$DGŔİƕůª/½ƒǃǾ/ǰƿ0E%(,

%"ƒñȶƪ"$/T}q)0Ȅ/Vkvıø*ƕůƿǕ>ſň¾ǦȌƮ/ǭđFǂ:ÛEB*)ȶ/Ɔİª£/½ƒǃǾ@-)&&B9ÚƕůƿǕ

/ DNA ȘFƀč(:"*DȶĿÛó/Ȕ°ć-Ē(ȶƆİȔ°ć?2ƆİǕć
0 2 «Ș/¬FƩȶǨƢ)B*Ő@*,%"Ŭ-ȶȄ/İŞǝ£Ȩô/Ɲǫ-?%(Ī@C"İ-Ȉș"ȑćFtP*"ZLk_KnTF,%"
/ǄŜȶ¸ïÚò)ĽȡC"[ZIOk|)=tP0Č¸-ĿÛó-Ȉș(B*Ő@*,%"Ŏ)ȶƆİȔ°£*ƆİǕć£0Ïƾ,ƕůƿǕ/ħĴ@/KwZ)0ȶƆİȔ°£0ŔİȔ°£«ňË"=/ȶƆİǕć£0ŔİǕć£ƆİË"=/)

B*Ĳ³C"ȶďȠ-0Ɔİª£*=-Ǩň/š/¸()ĿÛó/tPŤ½CB*Ŕ(ǄŜ)%"ƆİȔ°£=ƆİǕć£=¹-ĿÛó/VkvF

B*@ȶƆİª£0ŔİǕć£@½ƒ(

B*ǐ@CB Ŕƻ¾Ǧŕ?2ſň¾Ǧŕ/Şǝ£ÊĴFǭđ"9Ŕƻ¾Ǧŕ/ƿǕFǭđ"*DȶŔİȔ°£ȳNȴ)0 7 ŖȶŔİǕć£ȶƆİȔ°£ȶƆİǕć£)0 14 Ŗ/ŞǝK. Ichihara & S. Kawano- 5{}|o~~lxzqtulmrpqwnl

AKV'9Z xzqtvl mrpqwnl£ǭđ)" /*@Ɔİª£0>0AȶĿÛó/VkvFĹ%"ǨƢ)B*EBſň¾ǦÃŕ)0Ȕ°£0 7 ŖȶŔİǕć£ȶƆİȔ°£ȶƆİǕć£)0 14 Ŗ/Şǝ£ǭđ)ȶŕ-,B*Úƕůó/ƿǕ)ȶǹȍȥ-Şǝ£4/ǭđC"ŔİȔ°£)0 7 ŖȶŔİǕć£ȶƆİȔ°£ȶƆİǕć£)=ÝŨ-Şǝ£
0 7 Ŗ)ȶǨƢ/ƿǕ)0¦Şǝ£Fħĵ(B*ƩæC"Ƶſň¾Ǧǁ

ő-0ŔİǕć£) 7 Ŗ/Şǝ£ǭđCB/-Ē(ȶƆİȔ°£ȶƆİǕć£)0ſň(,%"9"ŔİǕć£)/ǭđFǆ"*Dȶȓ/ƿǕ)0ƆİȔ°£>ƆİǕć£*ÝŨ/Şǝ£ň/ſňF

E,¾ǦFǭđB*)"4. ¨§¡¤ª¯b(G6aC-l(N=2#);2]%

$X]

d+G6
a `³;2]%

fX]³J

2!

WG6a<Y/?
^G6 a ſň¾ǦȌƮ/ǄŜF9*<B* ŔİǕć£/ȓ/ƿǕ>ƆİȔ°£ȶƆİǕć£)0¦Şǝ£/ħĵ9)0Ȇĝ/ſň¾ǦȌƮ*ÝŨ-ȉ;?

#ȶ/ĩȶĄăŞǝ¾£

¾ȣȶť5ƭÊB*)Şǝ£ň/ſňFêȒ"¾Ǧ,EC(B*ŀčCȶȟţƌ/IstKOY[*ÝŨ/B*įEC"ȳî 4ȴ/ƒǷ0ȌƮ)ȗþ)ĽȡC"ƆİǕć£>ƆİȔ°£/¸(-(ĿÛóVkvŤ½C"*@=ŃĹCB9"ďȠ-ŔİǕć£@Ī@C" F1 Ȕ°£/ZLk_
KnTF,%"*Dȶ10%ƮĠ/ª£)ĿÛóVkvFŔBª£ȳmt±šȴŤ½C"ȳî 5ȴĪ@C" mt±š5. ;2]%;2]%"emt!

0F1gM

5!LRTj% ­±¥±PCR§£ª©¢«°¦ mt²!³mt´!
±Bmt³­±¥ 4U

G6aR+M\

K. Ichihara & S. Kawano- 6{}|o~~lxzqtvlmrpqwnl

AKV'9Z xzqtwl mrpqwnl

FĵƊȶƕůƿǕFǭđ"*Dȶ2 ŖȦŲFĹ&ȋǺć)ȶDNA Ș0ƆİȔ°ć>
ƆİǕć*ÝŨ- 2N )ƿǕčň

%"/*@ȶŔİǕć£)ſň¾Ǧŭĝ-ȉǣ,Aȶ@Ɔİª£½ƒB*ŀčC"&9A[ZIOk|)=ȟţƌ/IsuSY[ȶƍ-eJq[s|-¡"ǃǾ-?%(Ɔİš½ƒ(*B

*-,B

XR

%.



l IstKOY[/¾ćwPi\v-&(0YKjh\h)ýƜ£Ǯś,ECȶſň¾Ǧŕ/Şǝ£/ȔßFǴŉ(

B DYAD/SWITCHȳSW1ȴȑć/ýƜ£ȳRavi et al.

2008ȴ>ȶAtspo11-1/Atrec8/osd1 /ȖýƜ£ȳd’Erfurth et al. 2009ȴ)0¦Şǝ0ħĵCBȶƵ¾Ǧ)ĄăŞǝ¾£ť-ģȣCȶŞǝ£ňſň,,

*øáC(

YzŸƶ/ĸǤF

B@-ȂĞƝȎ"ŬYU[ĸǤ*tKS`K]S
E(ƖB*)ƕůƿǕƼ¿)/f{[S|qfvǮś=,ECȶIs[s|F,(,¾ǦȌƮF

Hieracium praealtum )0ſň¾ǦȜȈ/ȑćƝƒ/-Ē(ȳOkada et al. 2013ȴȶeJq[s|F,

gunnisoniana )0Ǫ,ſň¾ǦȜȈȑćƝƒ(

Ip{hƫ/ BoecheraB*=Ő@-,%("ȳSchmidt et al. 2014ȴȟţƌ)0ȑƞ,öƠFŐ@-B"</IsuSY[ƧƲ
ȉ<@C(

B

 [ZIOk|)0+/?
[ZIOk|/,ȑćǎIsuSY[-ȜE%(

C!C/ŠȮƿǕ*ƕůƿǕFƖ

B/#D(,%" RNA-seq ďȰ/ǄŜFǀBſň¾ǦȜȈȑć-ƥơ(ǫ"*DȶSYN3 > SCC3 */ĒÛ/ǇĹ-²(Ŭ-ȶ%"ĄăŞǝ¾£

BWlYȜȈȑćȳCarlile and Amon 2008; Yuan et al. 2012;

Grelon 2014ȴ/ƝƒȘƆİȔ°ć>ƆİǕć)¢,%(B*Ő@*,%"

C0ƆİȔ°£>ƆİǕć£-BƕůƿǕħĵő/ſň¾ǦȌƮ)ĄăŞǝ¾£œÀ-ť¯-¾ȣ(9*ŀčFǧB=/*=ǐ@CBĩ0Vkvıø/

ŉ±Fȉ<B**=-ȶƕůƿǕħĵő/ſň¾ǦȜȈȑć/ƝƒȘ/ŀƭFǴ6B*)ȶ[ZIOk|/IstKOY[/Òëȑć/ƍčFơĺ"

YR

*ǐ(

B



l [ZIOk|/Ɔİƕůš/½ƒǃǾ0C9)/ƧƲ)?

Ő@-,%("C)0ďȠ-[ZIOk|)0ƕůó-?%(¢úƃĠ5/ȐĮĠ-Ě

B/#D



ǠǑ/T}q)0ŔİȔ°£ȶƆİȔ°£ȶƆİǕć£FŽųŘ*ŻųŘ)õȮ"Ƞ/ȑćƝƒýÊ/ĚƜFŐ@-B"<-ȶRNA-seq Ǯś=ǣ(ȉ<(BȑćƝƒýÊFű6(:B*ȶƕǓòƄ/Ɯ,BŔİȔ°£ȶƆİȔ°£ȶƆİǕć£)0ȶŽųŘ5/ĮƷİ0%A*Ɯ,B*Ő@-,%"Ŏ)ȶŽųŘ*ŻųŘ)õȮȶƕȚƐFűȀ(:(=ȶšț-07*G+Ě:@C,%"@-ȶƿǕº/KOƃĠ>Iukȕǂĵƶ=Ǵ6B*)ȶ?Aǰƿ-ȑƞȶƕƓƞ,ȏ
@-"

*ǐ(

FŐ

B
K. Ichihara & S. Kawano- 7{}|o~~lxzqtwlmrpqwnl

AKV'9Z xzqtxl mrpqwnl

 ĩ0éÍę)Ľȡ"ª£ǎ>òô/ŔİšȶƆİšFƖ
ȇ/ǮśFǣ,RAD-seq -?Bȡìŧ*)Ɔİš/½ƒ/ȩĠ>Úò/Ŕİš*Ɔİš/ƼǅȜ¨ƶFŐ@-BďȰ=ǯƚ)B/?

,Ũ,Iqa@/ƧƲFȖ.B*)ȶœǁ

ƞ-[ZIOk|/ƕůó/ȉËȶƔûȐĮ-+/?
B*)C1*ǐ(

-Ǆ2&

(

B/FŐ@-B

2E8C
Carlile, T.M. & Amon, A. 2008. Meiosis I is established through division-specific translational control
of a cyclin. Cell 133:280–291.
Cock, J.M., Sterck, L., Rouzé, P., Scornet, D., Allen, A.E., Amoutzias, G., Anthouard, V.,
Artiguenave, F., Aury, J.M., Badger, J.H., Beszteri, B., Billiau, K., Bonnet, E., Bothwell, J.H.,
Bowler, C., Boyen, C., Brownlee, C., Carrano, C.J., Charrier, B., Cho, G.Y., Coelho, S.M., Collén,J., Corre, E., Da Silva, C., Delage, L., Delaroque, N., Dittami, S.M., Doulbeau, S., Elias, M.,
Farnham, G., Gachon, C.M., Gschloessl, B., Heesch, S., Jabbari, K., Jubin, C., Kawai, H., Kimura,K., Kloareg, B., Küpper, F.C., Lang, D., Le Bail, A., Leblanc, C., Lerouge, P., Lohr, M., Lopez, P.J.,
Martens, C., Maumus, F., Michel, G., Miranda-Saavedra, D., Morales, J., Moreau, H., Motomura,T., Nagasato, C., Napoli, C.A., Nelson, D.R., Nyvall-Collén, P., Peters, A.F., Pommier, C., Potin, P.,
Poulain, J., Quesneville, H., Read, B., Rensing, S.A., Ritter, A., Rousvoal, S., Samanta, M., Samson,
G., Schroeder, D.C., Ségurens, B., Strittmatter, M., Tonon, T., Tregear, J.W., Valentin, K., vonDassow, P., Yamagishi, T., Van de Peer, Y., & Wincker, P. 2010. The Ectocarpus genome and the
independent evolution of multicellularity in brown algae. Nature 465:617–621.
Collén, J., Porcel, B., Carré, W., Ball, S.G., Chaparro, C., Tonon, T., Barbeyron, T., Michel, G., Noel,B., Valentin, K., Elias, M., Artiguenave, F., Arun, A., Aury, J.M., Barbosa-Neto, J.F., Bothwell,
J.H., Bouget, F.Y., Brillet, L., Cabello-Hurtado, F., Capella-Gutiérrez, S., Charrier, B., Cladière, L.,
Cock, J.M., Coelho, S.M., Colleoni, C., Czjzek, M., Da Silva, C., Delage, L., Denoeud, F.,
Deschamps, P., Dittami, S.M., Gabaldón, T., Gachon, C.M., Groisillier, A., Hervé, C., Jabbari, K.,
Katinka, M., Kloareg, B., Kowalczyk, N., Labadie, K., Leblanc, C., Lopez, P.J., McLachlan, D.H.,
Meslet-Cladiere, L., Moustafa, A., Nehr, Z., Nyvall Collén, P., Panaud, O., Partensky, F., Poulain, J.,
Rensing, S.A., Rousvoal, S., Samson, G., Symeonidi, A., Weissenbach, J., Zambounis, A., Wincker,
P., & Boyen, C. 2013. Genome structure and metabolic features in the red seaweed Chondrus
crispus shed light on evolution of the Archaeplastida. Proc. Natl. Acad. Sci. U S A. 110:5247–5252.
d’Erfurth, I., Jolivet, S., Froger, N., Catrice, O., Novatchkova, M., & Mercier, R. 2009. Turning
meiosis into mitosis. PLoS Biol. 7(6): e1000124.
Ferris, P.J. & Goodenough, U.W. 1997. Mating type in Chlamydomonas is specified by mid, theminus-dominance gene. Genetics 146:859–869.
Geng, S., De Hoff, P. & Umen, J.G. 2014. Evolution of sexes from an ancestral mating-type
specification pathway. PLoS Biol. 12(7):e1001904.
Grelon, M. 2014. The Molecular Biology of Meiosis in Plants. Annu. Rev. Plant Biol. 66:1–16.
Hayden, H.S., Blomster, J., Maggs, C.A., Silva P.C., Stanhope, M.J., & Waaland, J.R.2003. Linnaeus
K. Ichihara & S. Kawano- 8{}|o~~lxzqtxlmrpqwnl

AKV'9Z xzqtyl mrpqwnl

was right all along: Ulva and Enteromorpha are not distinct genera. Eur. J. Phycol. 38:277–294.
Hernández, I., Peralta, G., Pérez-Lloréns, L., Vergara, J.J., & Niell, F.X. 1997. Biomass and dynamics
of growth of Ulva species in Palmones River estuary. J. Phycol. 33:764–772.
Hiraoka, M., Dan, A., Shimada, S., Hagihira, M., Migita, M. & Ohno, M. 2003a. Different life
histories of Enteromorpha prolifera (Ulvales, Chlorophyta) from four rivers on Shikoku Island,
Japan. Phycologia 42:275–284.
Hiraoka, M., & Higa, M. 2016. Novel distribution pattern between coexisting sexual and obligate
asexual variants of the true estuarine macroalga Ulva prolifera. Ecol. Evol. 6:3658–3671.
Hiraoka, M., Ichihara, K., Zhu, W., Ma, J., & Shimada, S. 2011. Culture and hybridization
experiments on an Ulva clade including the qingdao strain blooming in the yellow sea. PLoS One
6(5): e19371.
Hiraoka, M., Shimada, S., Serisawa, Y., Ohno, M., & Ebata, H. 2003b. Two different genetic strainsof stalked-Ulva (Ulvales, Chlorophyta) grow on intertidal rocky shores in Ebisujima, central Japan.
Phycol. Res. 51:161–167.
Hiraoka, M., Shimada, S., Uenosono, M. & Masuda, M. 2004. A new green-tide-forming alga, Ulva
ohnoi Hiraoka et Shimada sp. nov. (Ulvales, Ulvophyceae) from Japan. Phycol. Res. 52:17–29.
van den Hoek, C., Mann, D.G. & Jahns, H.M. 1995. Algae. An Introduction to Phycology. Cambridge
University Press, Cambridge.
Ichihara, K., Arai, S., Uchimura, M., Fay, e.J., Ebata, H., Hiraoka, M., & Shimada, S. 2009. New
species of freshwater Ulva, Ulva limnetica (Ulvales, Ulvophyceae) from the Ryukyu Islands, Japan.
Phycol. Res. 57:94–103.
Kakinuma, M., Ikeda, M., Coury, D., Tominaga, F., Kobayashi, I., & Amano, H. 2009. Isolation andcharacterization of the rbcS genes from a sterile mutant of Ulva pertusa (Ulvales, Chlorophyta) andtransient gene expression using the rbcS gene promoter. Fish. Sci. 75:1015–1028.
Leliaert, F. & Lopez-Bautista, J.M. 2015. The chloroplast genomes of Bryopsis plumosa andTydemania expeditiones (Bryopsidales, Chlorophyta): compact genomes and genes of bacterial
origin. BMC Genomics 16:204.
Leliaert, F., Zhang, X., Ye, N., Malta, E., Engelen, A.H., Mineur, H.V., & De Clerck, O. 2009.
Identity of the Qingdao algal bloom. Phycol. Res. 57:147–151.
Mareš, J., Leskinen, E., Sitkowska, M., Skácelová, O, & Blomster, J. 2011 True identity of theeuropean freshwater Ulva (Chlorophyta, Ulvophyceae) revealed by a combined molecular andmorphological approach. J. Phycol. 47:1177–1192.
Melton, J.T., Leliaert, F., Tronholm, A. & Lopez-Bautista, J.M. 2015. The complete chloroplast andmitochondrial genomes of the green macroalga Ulva sp. UNA00071828 (Ulvophyceae,
Chlorophyta). PLoS One. 10(4): e0121020.
Merchant, S.S., Prochnik, S.E., Vallon, O., Harris, E.H., Karpowicz, S.J., Witman, G.B., Terry, A.,
Salamov, A., Fritz-Laylin, L.K., Maréchal-Drouard, L., Marshall, W.F., Qu, L.H., Nelson, D.R.,
Sanderfoot, A.A., Spalding, M.H., Kapitonov, V.V., Ren, Q., Ferris, P., Lindquist, E., Shapiro, H.,
Lucas, S.M., Grimwood, J., Schmutz, J., Cardol, P., Cerutti, H., Chanfreau, G., Chen, C.L., Cognat,
K. Ichihara & S. Kawano- 9{}|o~~lxzqtylmrpqwnl

AKV'9Z xzqupl mrpqwnl

V., Croft, M.T., Dent, R., Dutcher, S., Fernández, E., Fukuzawa, H., González-Ballester, D.,
González-Halphen, D., Hallmann, A., Hanikenne, M., Hippler, M., Inwood, W., Jabbari, K.,
Kalanon, M., Kuras, R., Lefebvre, P.A., Lemaire, S.D., Lobanov, A.V., Lohr, M., Manuell, A.,
Meier, I., Mets, L., Mittag, M., Mittelmeier, T., Moroney, J.V., Moseley, J., Napoli, C., Nedelcu,
A.M., Niyogi, K., Novoselov, S.V., Paulsen, I.T., Pazour, G., Purton, S., Ral, J.P., Riaño-Pachón,D.M., Riekhof, W., Rymarquis, L., Schroda, M., Stern, D., Umen, J., Willows, R., Wilson, N.,
Zimmer, S.L., Allmer, J., Balk, J., Bisova, K., Chen, C.J., Elias, M., Gendler, K., Hauser, C., Lamb,
M.R., Ledford, H., Long, J.C., Minagawa, J., Page, M.D., Pan, J., Pootakham, W., Roje, S., Rose,A., Stahlberg, E., Terauchi, A.M., Yang, P., Ball, S., Bowler, C., Dieckmann, C.L., Gladyshev,
V.N., Green, P., Jorgensen, R., Mayfield, S., Mueller-Roeber, B., Rajamani, S., Sayre, R.T.,
Brokstein, P., Dubchak, I., Goodstein, D., Hornick, L., Huang, Y.W., Jhaveri, J., Luo, Y., Martínez,D., Ngau, W.C., Otillar, B., Poliakov, A., Porter, A., Szajkowski, L., Werner, G., Zhou, K.,
Grigoriev, I.V., Rokhsar, D.S., & Grossman, A.R. 2010. The Chlamydomonas genome reveals theevolution of key animal and plant functions. Science 318:245–250.
Nakamura, Y., Sasaki, N., Kobayashi, M., Ojima, N., Yasuike, M., Shigenobu, Y., Satomi, M.,
Fukuma, Y., Shiwaku, K., Tsujimoto, A., Kobayashi, T., Nakayama, I., Ito, F., Nakajima, K., Sano,
M., Wada, T., Kuhara, S., Inouye, K., Gojobori, T., & Ikeo, K.

2013. The first symbiont-freegenome sequence of marine red alga, Susabi-nori (Pyropia yezoensis). PLoS One 8(3): e57122.
Nishitsuji, K., Arimoto, A., Iwai, K., Sudo, Y., Hisata, K., Fujie, M., Arakaki, N., Kushiro, T., Konishi,T., Shinzato, C., Satoh, N., & Shoguchi, E. 2016. A draft genome of the brown alga, Cladosiphon
okamuranus, S-strain: a platform for future studies of “mozuku” biology. DNA Res. 23 (6):561-570.
Oertel, W., Wichard, T. & Weissgerber, A. 2015. Transformation of Ulva mutabilis (Chlorophyta) byvector plasmids integrating into the genome. J. Phycol. 51:963–979.
Ogawa, T., Ohki, K. & Kamiya, M. 2015. High heterozygosity and phenotypic variation of zoids inapomictic Ulva prolifera (Ulvophyceae) from brackish environments. Aquat. Bot. 120:185–192.
Okada, T., Hu, Y., Tucker, M.R., Taylor, J.M., Johnson, S.D., Spriggs, A., Tsuchiya, T., Oelkers, K.,
Rodrigues, J.C., & Koltunow, A.M. 2013. Enlarging cells initiating apomixis in Hieracium
praealtum transition to an embryo sac program prior to entering mitosis. Plant Physiol. 163:216–31.
Prochnik, S.E., Umen, J., Nedelcu, A.M., Hallmann, A., Miller, S.M., Nishii, I., Ferris, P., Kuo, A.,
Mitros, T., Fritz-Laylin, L.K., Hellsten, U., Chapman, J., Simakov, O., Rensing, S.A., Terry, A.,
Pangilinan, J., Kapitonov, V., Jurka, J., Salamov, A., Shapiro, H., Schmutz, J., Grimwood, J.,
Lindquist, E., Lucas, S., Grigoriev, I.V., Schmitt, R., Kirk, D., & Rokhsar, D.S. 2010. Genomic
analysis of organismal complexity in the multicellular green alga Volvox carteri. Science
329:223–226.
Ravi, M., Marimuthu, M.P. & Siddiqi, I. 2008. Gamete formation without meiosis in Arabidopsis.
Nature 451:1121–1124.
Schmidt, A., Schmid, M.W. & Grossniklaus, U. 2015. Plant germline formation: common concepts
and developmental flexibility in sexual and asexual reproduction. Development 142:229–241.
K. Ichihara & S. Kawano- 10{}|o~~lxzquplmrpqwnl

AKV'9Z xzquql mrpqwnl

Schmidt, A., Schmid, M.W., Klostermeier, U.C., Qi, W., Guthörl, D., Sailer, C., Waller, M.,
Rosenstiel, P., & Grossniklaus, U. 2014. Apomictic and sexual germline development differ withrespect to cell cycle, transcriptional, hormonal and epigenetic regulation. PLoS Genet. 10(7):e1004476.
Sharbel, T.F., Voigt, M.L., Corral, J.M., Galla, G., Kumlehn, J., Klukas, C., Schreiber, F., Vogel, H.,& Rotter, B. 2010. Apomictic and sexual ovules of Boechera display heterochronic global gene
expression patterns. Plant Cell 22:655–671.
Shimada, S., Hiraoka, M., Nabata, S., Iima, M., & Masuda, M. 2003. Molecular phylogenetic analysesof the Japanese Ulva and Enteromorpha (Ulvales, Ulvophyceae), with special reference to thefree-floating Ulva. Phycol. Res. 51:99–108.
Shimada, S., Yokoyama, N., Arai, S. & Hiraoka, M. 2008. Phylogeography of the genus Ulva
(Ulvophyceae, Chlorophyta), with special reference to the Japanese freshwater and brackish taxa. J.
Appl. Phycol. 20:979–989.
Suzuki, R., Ota, S., Yamazaki, T., Toyoda, A., Nonaka, S., Matsukura, C., Kuwano, K. & Kawano, S.
2016. Morphological changes of giant mitochondria in the unicellular to multicellular phase duringparthenogenesis of Ulva partita (Ulvophyceae) revealed by expression of mitochondrial targeting
GFP and PEG transformation. Phycol. Res. 64:176–184.
Ye, N., Zhang, X., Miao, M., Fan, X., Zheng, Y., Xu, D., Wang, J., Zhou, L., Wang, D., Gao, Y.,
Wang, Y., Shi, W., Ji, P., Li, D., Guan, Z., Shao, C., Zhuang, Z., Gao, Z., Qi, J., & Zhao, F. 2015.
Saccharina genomes provide novel insight into kelp biology. Nat. Commun. 6:6986.
Yuan, L., Yang, X., Ellis, J.L., Fisher, N.M., & Makaroff, C.A. 2012. The Arabidopsis SYN3 cohesin
protein is important for early meiotic events. Plant J. 71:147–160.
Q1O 1998. 78=F_c. pp. 1222. Q[k³>.

K. Ichihara & S. Kawano- 11{}|o~~lxzquqlmrpqwnl