植物科学最前線 7:66 (2016)  8(*$)'%&W/9@V5 Y+ACRHRZ5
ňÍĖò 1*}ÑŒÈü 2
1ǦŕíôôǯƁô÷ǽR|V~ 920-0934 ǦŕĆöŰ 13-1

ǿ ũěāȆƈĪíôŪôǠ 171-8501 ĿǡǉăÇƽœƸ 3-34-1Ȁ
2
Max Planck Institute for Plant Breeding Research*Carl-von-Linné-Weg 10, 50829 Cologne Germany
The evolution of TALE homeobox transcription factors contributes to the increase in the complexity ofthe diploid body plan in land plants
Key words: Alternation of generations; Evolution of sporophytes; Gene duplication; KNOX; BELL
Keiko Sakakibara1*, Chihiro Furumizu2,
1

Advanced Science Research Center, Kanazawa University, Kanazawa 920-0934, Japan(*Current address: Rikkyo University, 3-34-1 Nishi-Ikebukuro,Toshima-ku, 171-8501, Tokyo Japan)
2

Max Planck Institute for Plant Breeding ResearchCarl-von-Linné-Weg 10, 50829 Cologne, Germany\[W/9XRV5643 ǟòǤƼ(ĮǀǟòŨđ'ÍÅÃ"8#ƥ 69! (Ohno 1970)ǤƼ&5!è ǟò'ì(ªǟòÆ (Pseudogenization)&5!řî8 ȁǤƼ&ůľ8ȃ'ǟò C
wO`x'ǟò ğ!ŋƩ<ĎÂºğ85 &%8ŋƩºÆ (Subfunctionalization)4ȁǤƼ
&57è ǟò'Ţį 3#3#ğ!ŋƩ<Ɯġ8į"ȁ3 Ţį'ǟò ĮŋƩ<Ũđ8ĮǀŋƩŨđ (Neofunctionalization) Ǐ8# Ż69!8ǿßȂȀǟòǤƼ& ŋƩºÆ4ĮǀŋƩŨđ Ǔ¸Þò&Ǐ8#ȁŘ'ǟòŲũ íëÆ8ƙłȁ'Ŭţ&ĮČǍ'Ũđ<36ÍÅÃ#%78ǭņţ& !3ȁǟòǤƼĐ'ŋƩºÆ4ĮǀŋƩŨđ ıó'ǟò¿ĒcY^{~G<57ƼǱ&ȁ'ǙÆ&ǌŧ
#ƥ

69!8

(Rensing 2014)/ȁÕßȂ ǟòǤƼ&5!è ǟò'$8ǚÛƄ¶"ǤƼ&57Ŭŷ ǤƼ&5!èÄǟò'ì(Ţį řî8 ȁŋƩºÕǟò(fvzH#Ú)Æȁ8(ĮǀŋƩŨđ&% 8çÔ389ȁƄºÆ&5!Ŭ
C~TzH#(Ç¾98

K. Sakakibara and C. Furumizu- 1
BSJ-Review 7:66 (2016)

植物科学最前線 7:67 (2016) ǭņţ(ŚŒĕƞƭƳǺ'NuOGtƳǺ'ǩ 6ƒ4®8000ĈÀ&ºĂ#ƥ 6
9!8(Lewis and McCourt 2004; Wellman et al. 2003; Wickett et al. 2014)NuOGtƳǺNuOGt#ǭņţ'Ŭŗū<őǔ!08#ȁNuOGt(Ņŷ Êŷ' &ìƔƨ'Ǣ© <¡8 ȁƼ
ŷ' (ÊƔƨ'ĤÔò'0"8į"ȁǭņţ(Ƽŷ' 3ìƔƨ'ƨò <ČĚ8ǭņţ'ǙÆ'½Ĺ&'Ƒƚ 6ºĂ#ƥ 698KIņţ"(Ŭŗū'íÉ(Êŷ'Ǣ©
"ȁƨò (þȁǢ© &£óų&ČĚ989&ûƹòņţ"(Ƽŷ'ƨò
Ŭŗ'#%7ȁǢ© (īƔƨ&/"ǗÆ!8(Graham et al. 2000)ǿĻƝǅǰ'żĄ#ňÍ'ßȃ<ÎšȀ
'# 6ȁǭņţ'ǙÆ'Ǜƃ"Ƽŷ'ņţ 'ìƔƨÆ#ąíÆ Ǐ#ƥ
698ĻƝǅ"(ǭņţ'Ƽŷ 'ǙÆ&ǌŧ#ƥ 698ȃ'nsCoYGPâǓ¸ÞòȁKNOXǟòİ#BELLǟòİ'ǟòǤƼ#'Đ'ǟòǙÆ&!ƕ8

[KOAC  8(*$)'%&W/9! W/9XR] W/9 W/9
 nsCoYGPâǓ¸Þò(ŸŅŬţ µǘ&ġǏŞ'ÑǓ¸Þò'"7ȁŸŅŬţ'µǘƀ°& !ȁDNAƙÔƩ<ſnsC_s@|'ƋȂ+wYYGP#ƋȃlwYGP'ǩ&Ȅ>qdǣŏå'ģ²<×1TALEânsCoYGPǟò#Ȅ>qdǣŏå'ģ² %ǳTALEânsCoYGPâǟò&º 9!#ƥ 69!8(Derelle et al. 2007)ņţ"(TALEâǓ¸Þ
ò(KNOXǟòİ#BELLǟòİ&º 9! 7ȁ93ƞƭņţ#ÊƔƨƓƳN>a]?CN
U|Cyanidioschyzon merolae"ĉ¦ó9!8(ßȃ; Mukherjee et al. 2009; Sakakibara 2016)ƞƳǺ#ű%7ǭņţ'Ƒƚ"(KNOXǟòİ(ǟòǤƼ&57KNOXGvPȂǟòİǿKNOX1Ȁ#KNOXGvP2ǟòİǿKNOX2Ȁ&ºĂ!8(ßȃ; Mukherjee et al. 2009)KIņţ#Ɯƌļņţ'µǘƀ°"(KNOX1ǟòİ#KNOX2ǟòİ'ǟòī(99Ȃ§"#ĥõ98 , 'Đ&ǏǟòǤƼ'ƙł, Ƅòņţ"(6&ǟòīßȃ ƓƳÏ*ƞƭņţ&8 BELL/KNOXǟòİ'ǟòī'ëǞ
ƓƳ#ũŬƞƭņţ'Ʒų%ºǺƤ'ƑƚǪ¤#ºĂĶŠ"óà#ĥõ98
BELL/KNOX ǟò'ī<ſĥõǟòī(ƓƳ#ũŬƞƭņţ
6Êǲ9 BELLǟò# KNOX ǟò'ºòƑƚǁŀ'ƙł (Mukherjee et al. 2009; Sakakibara 2016)&å NWǺ'³Jdrǁǆ(ũàǙƶ"7ȁĐȁBELL/KNOX ǟòī ĵž&%8#Ĺď98

K. Sakakibara and C. Furumizu- 2
BSJ-Review 7:67 (2016)

植物科学最前線 7:68 (2016)èÄ!8KNOXǟò#ÕŊ&, ǭņţ'BELLǟò&!3ǟòǤƼ&57ǟòīèÄ8¬Ö ǃ2698įȁƳǺ'BELLǟò"(ȁƓƳN>a]?CNU|ȁÏ*ƞƳǺ(ǟòǤƼ&5!Ŭȃ§'BELLǟò<ġ ȁNuOGtƳǺGyjTxq]?Ar
Klebsormidium flaccidum"(Ȃ§óà8ºĂĈ'Ñ 6ȁƳǺ'BELLǟò#ǭņţ'BELLǟò'ƑƚǪ¤(ĺĵ6 "(%, 6%8ǁŀ ď98
 BELLǟò#KNOXǟò'ƀ°ų%ŋƩ<ƥú8"Ǥƾ%ȁƞƳǺ'TALEânsCoYG
Pǟò&Ǫ8ŽƆ<ō'Ƿ"ƕ8


[KOACRHGE#2<  8(*$)'%&W/9 ÊƔƨƞƳǺGvq_t`P Chlamydomonas reinhardtii (kvP#p@`P'ȃ'Ǣâ<ġȁŃǼŌĽ"(Êŷ'ŃǼƔƨ Ǣ©ò+#ºÆ8'ǯ&ȁkvPâǢ©ò( BELL ŊV|fGǍ<K~_8 GSP1 (Gamete-specific plus1)ǟò<ȁp@`PâǢ©ò( KNOX V|fGǍ<K~_8 GSM1 (Gamete-specific minus1)ǟò<Ųũ896'V|fGǍ(99'Ǣ©
ò& !(ƔƨǍ&Āà8KNOX V|fGǍ(Ǔ¸Þò#!­ ȁƪǒ(ŅƂƶNH`x<ġȁĤÔĐȁBELL V|fGǍ#l\zW@p~<ČĚ8#"Ņ+#ƂƶȁĤÔòǿƼ
ŷȀŤűų%ǟòŲũ<Ǉƍ8#"Ƽŷ 'ºÆ<¿Ē!8# ſ9(Lee et al.
2008; Zhao et al. 2001)96'÷ǽ4ƞƭņţ& 8 TALE ǟò'Ǣ¼ǁŀ'ƙł&å !ȁ
Lee 6(2008)( KNOX ǟò# BELL ǟò'ǙÆ ǭņţ'Ƽŷ 'ìƔƨÆ&Ǥƾ"# ǅ<ĦÝ!8ō'Ƿ"(ǭņţ'Ŭŗū'ǙÆ& 8 TALE ânsCoYGPǟ
ò'ĎÂ<ƥú8"Ǥƾ%Żƿ<36ȁgsZwEcLI' KNOX ǟòİ# BELL ǟòİ&Ǫ8ŽƆ<ƕ8(Horst et al. 2016; Sakakibara et al. 2013)

ßȄ ÊƔƨƞƳGvq_t`P'Ŭŗū# KNOX ǟò# BELL ǟò'ĎÂ Lee et al. 2008 57ĩë

K. Sakakibara and C. Furumizu- 3
BSJ-Review 7:68 (2016)

植物科学最前線 7:69 (2016)[  W/9 W/9Y+AC,.-.#2<  åǠǭņţ& 8 KNOX2 ǟòİ# BELL ǟòİ'ŋƩ&!(KIņţR|Ǻ'gsZwEcLI Physcomitrella patens <Ůǁŀ Ǚ="8gsZwEcLIJdr&( 2 §' KNOX2 ǟò K~_9! 7ǿßȃȀ
ȁ93Ƽŷ'ƨò "Ųũȁ'ǤŋƩŌĨëű (ƨò 'ƧŲŬ ½Ĺ'ŐǮ"¨Ŏ8'Ƨ<ÊǲäǼ8#ȁŅŷ(Ƽŷ'//ƨò
6Ǣ© ŊƗƣ<ºÆ# 6ȁKNOX2 ǟò Ƽŷ& !Ǣ© 'ŲŬkzHvr<ĝ¿8#" <¿Ē!8# ſ9(Sakakibara et al. 2013)
 įȁgsZwEcLI(ȅ§'BELLǟò<ġ ȁ' '"8PpBELL1ǟò<Ǣ© & !ǛÁŲũ8#ȁŅŷ(Êŷ'//ƨò ŊƗƣ Ǆý9(Horst et al. 2016)'ƙł 6ȁgsZwEcLI'BELLǟò ƨò 'ŲŬkzHvr<¿Ē8#&57ȁ '»ħ &Ǫ8# ſ9/ȁBiFCǁŀ#Ú)98ĜŖ"VeK'ƷŴƔƨ<Ů!V|
fGǍǩ'ŷ¡Ů<ǁŀƙłȁPpBELL1V|fGǍ#ƴ±V|fGǍĭŢ#'ƵÔV|fGǍgsZwEcLI'KNOX1/(KNOX2V|fGǍ#ƴ±V|fGǍĭŢ#'ƵÔV|fGǍ'9#3ŷ¡ŮÒƩ"8# ſ9!8'ƙł(ȁgsZwEcLI"3PpBELL1
V|fGǍ gsZwEcLIKNOXV|fGǍ'9 #ţŪų&ŷ¡Ů! <¿Ē
!8ÒƩĕ<ſÜ8(Horst et al. 2016)96'ŽƆ&5!ȁKNOXǟò#BELLǟò&
58Êŷ#Ƽŷ'»ħ 'ºòŋŉ ƞƳGvq_t`P#ǭņţgsZwEcLIǩ"8ƃĊ¦ó9!8# ſ9[  W/9M901QIL;=J>W/9
 ņţ"ķ½&Êǲ9nsCoYGPǟò(^AtzKNZea Mays

6Êǲ9KNOX1 ǟòİ&ā8ǟò"8 Knotted-1 "7(Vollbrecht et al. 1991)ȁKNOX1 ǟòİ&!(ƹòņţ<ŮŽƆ ìīȁæØ9!8(Hay and Tsiantis 2010)^AtzKN' Knotted-1 ǟò'æØ&ƛ!Nz@b`Q` Arabidopsis thaliana 'ƯǵºƺƗƣŌî8ëű

shootmeristemless (stm)'ÍÞǟò#! STM ǟò Êǲ9, KNOX1 V|fGǍ<K~_!8
# ĵ6 &%(Barton and Poethig 1993; Long et al. 1996)/ȁNz@b`Q`'' KNOX1 f
vzH'ŋƩŌĨëű "8 knat1/brevipedicellus (bp)(ƮƯ'ǧ ð698Ʒũâ<ſȁƔƨéÔĚǟòȁŤ&wHa|ÔĚƘǐ'ǟòŲũ Ĵ!8# 6ȁKNAT1/BP ǟò(Ɣƨ'ĺºÆťę<ǝ»&ƜġȁƮƯ'ƔƨºÆ#ǧ8À&wHa|'ŔŹ!/ '<ĝ¿!8#ƥ 69!8(Mele et al. 2003)įȁKNAT1/BP ǟò<Nz@b`Q`"ǛÁŲũ8#ư&űěų&ºƺƗƣ ČĚ98(Chuck et al. 1996)96'÷ 6ȁƹòņţ
KNOX1 ǟò(ƨò 'ƯǵºƺƗƣ'ČĚƜġ#ƔƨºÆ'ĝ¿&ŋƩ!8#ƥ 69!8(Hay and Tsiantis 2010)
 åǠǭņţ"8KIņţgsZwEcLI'ġȄ§' KNOX1 ǟò(ƨò 'ǵƉƔƨ#ºƺƗƣ"ŋƩ8gsZwEcLI'ƨò 'ºƺƗƣ(ƧŲŬ'½Ĺ&ČĚ98 ȁĐ&ºƺƗƣ řîȁȂ§'ƨò' <ºÆ!ƨò 'ŲŬ<Ɩ8gsZwEcLI' KNOX1ǟò(93'ƨò &Ķų&¡698ºƺƗƣ"Ųũ8gsZwEcLI KNOX1 ǟ

K. Sakakibara and C. Furumizu- 4
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植物科学最前線 7:70 (2016)òǤŋƩŌĨëű & !(ƨò 'ºƺƗƣ ǝ»&Ɯġ9%2ȁƙł#!*%ƨò <ČĚ8'# 6ȁKNOX1 ǟò(gsZwEcLI& !3ƹòņţ'KNOX1 C~TzH#ÕŊ&ºƺƗƣ<Ɯġ8ŋƩ<ğ!8#ƥ 698(Sakakibara et al. 2008)
 Nz@b`Q`'5 %Êư<ČĚ8ƹòņţ"( KNOX1 ǟò(ƯǵºƺƗƣ"Ųũȁư
ÍåČĚ&°ƈ!õưǸã"Ųũ řî89&û!ȁ^p^ Solanum lycopersicum 4Nz@b`Q`'ǖƢƄ'qXVcZIe` Cardamine hirsuta %$Ƽư<ČĚ8ņţ"( KNOX1 ǟò'Ųũ ưÍå'þưÍå"·*ŗĕÆ8#"Ƽư'ŲŬ ¿Ē9!8# Ż69!
8(Blein et al. 2008; Hay and Tsiantis 2010)/ȁƼư<ČĚ8ņţ" KNOX1 ǟò<ǛÁŲũçÔ(Ƽư'ƼǱ Ǚ98(Hareven et al. 1996)96'ŽƆ( KNOX1 ǟò'ŲũŊċ'ëÆ ƹòņţ'ư'ǙÆ#ìŊÆ&ǌŧ#<ſÜ!8'5 &ƹòņţ'ƨò 'Ưǵ"( KNOX1 ǟò'Ųũ(ĶƇǩų&Ɛơ&¿Ē9! 7ȁƯǵºƺƗƣ"ǧĹǩƜġ98į, ŲŬ½Ĺ'ưÍå"(ĝ¿98# ƼǱ%ŲũŊċ<ſ ȁgsZwEcLI'KNOX1 ǟò'Ųũ(ƨò 'ƧŲŬ'½Ĺ'ºƺƗƣ&Ǭõ9!8ƹòņţ'ưÍå"'KNOX1 ǟò'Ųũĝ¿&Ǫ8Ǔ¸Þò#! MYB _s@|<×1 ARP (ASYMMETRICLEAVES1/ROUGH SHEATH2/PHANTASTICA)V|fGǍȁLOB (LATERAL ORGAN BOUNDARIES)_s@|<×1 AS2 (ASYMMETRIC LEAVES2)V|fGǍ'ƼÔ 4 YAB (YABBY) V|fGǍ%$ æØ9!8(Byrne et al. 2000; Hake et al. 2004; Kumaran et al. 2002)96'ǇƍÞò'ǠǿAS1ȁAS2 4 YABȀ'C~TzH(gsZwEcLIJdr&(óà(Rensing et al. 2008;
Sakakibara et al. 2008)ȁǭņţ'ǙÆ'Ǜƃ"ȁƹòņţ<×1Ƒƚ" KNOX1 ǟò'ŲũǇƍ&Ǫ;8Į%^v|PÞò Ũđ9#ĥŝ98Ä !ŲũǇƍ&Ǫ;8NPǸã ǙÆƙłȁKNOX1 ǟò'Ųũ¿ĒcY^{~G ƼǱ&%#"ȁƯǵ"'Ųũ Ɯġ985&%7ȁƹòņţ'Ƒƚ"'ƨò 'ƼǱÆ&% #ƥ 698(榊原恵子 2013)

[P9AC W/9BN Knotted-1ǟò'Êǲ<ƉƟ#!ȁ
'C~TzH ǭņţ'Ŋ%ºǺƤ 6Êǲ9Ƒƚǁŀ'ƙłȁǭņţ KNOX ǟò(99ÊƑƚ' KNOX1 ǟòİ# KNOX2 ǟò
İ&º 98# ĳ 6ſ9!(Kerstetter et al. 1994)Ʀ'ǩ&(ȁǟò'ŲũfV~|4ǛÁŲũ 'Ʒũâ&í%ǜ 8%;ȁNWǺ4ƹòņţ' KNOX1 ǟò(ƨò'ºƺƗƣ"&Ųũ8į"ȁKNOX2 ǟò(ƨò 'ºÆ'Ŋ%Ɨƣ"Ųũ8(Bharathan et al. 1999; Reiser et al. 2000; Sano et al. 2005; Serikawa et al. 1997a)/ȁNz@b`Q`'
KNOX2 ǟò'"8 KNAT3 <ǛÁŲũ8ń(ǹƱ%Ʒũâ<ſ% (Serikawa et al.
1997a; Serikawa et al. 1997b)ȁ9(ºƺƗƣ'űěų%ČĚ%$'ǹƱ%Ʒũâ<ſ KNOX1 ǟò'ŋƩŨđâëű 4 KNOX1 ǛÁŲũņţ #(ûšų"86&ƬÙśũǊ#!ȁN
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ǟò( KNOX1 ǟò#(ű%7ȁƨò 'ºÆƗƣ"&ŋƩ8# ĥŝ9! ȁ

K. Sakakibara and C. Furumizu- 5
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植物科学最前線 7:71 (2016)'ǅ<ƻ8ķǖ'ŽƆĚł<ō&ƕ8
 Nz@b`Q` KNOX2 ǟòfvzH'ŏ7'Ȅ§'ÊŦëű (93ǥŬâ#-.ÕŊ'Ʒũâ<ſ ȁknat3 knat4 knat5 Ǥëű (Nz@b`Q` KNOX1 ǟò<ǛÁŲũĶ'
5 &ư »9Ǖ1Ʒũâ<ſ(Furumizu et al. 2015; Serikawa et al. 1997a; Truernit et al. 2006)/ȁ
KNAT3<BELLǟò##3&ƯǵºƺƗƣ"űěų&Ųũ8#ȁ
KNOX1 ǟò'ŋƩŌ
Ĩëű "8 stm ëű#Ǻ'Ʒũâ<ſKNOX1 Ǔ¸Þò#Ǔ¸ĝ¿_s@|'ƵÔV|fGǍ<ņţ"Ųũ8#ȁ
KNOX1'ŌĨâëű

#Ǻ'Ʒũâ<ſ#6ȁKNOX1 V|fGǍ(&>G\?m~V~âǓ¸Þò#!ŋ
Ʃ8#ĥŝ98ßȅ ǭņţ'ǙÆ# KNOX ǟò# BELL ǟòŋƩ'ǙÆ(Markel et al. 2002; Shani Ƒƚ& !ȁƞƳǺGvq_t`PȁKIņţgsZwEcLet al. 2009)įȁNz@IȁƹòņţNz@b`Q` ġ KNOX ǟò# BELL ǟò'ǟb`Q`' KNAT7 ǟòKi~ī#99'ºĂĶŠ"'ǟòKi~ī<ſ

ò K~_8 KNOX2KNOX ǟò( KȁKNOX1 ǟò( K1ȁKNOX2 ǟò( K2ȁBELL

V|fGǍ(wkyYǟò( B #!ſ99'Ŭŗū"ȁŜīºƺ<ǎźÌȁM~âǓ¸Þò#!

Êŷ <şƭȁƼŷ <ƞƭ"ſGvq_t`P"( KNOX

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V|fGǍ# BELL ŊV|fGǍ(Ǣ©ò'ĤÔĐȁl\zW@p~8(Li et al. 2011; Li et al.

<ČĚ!ĤÔòºÆ<¿Ē8ǭņţ'¹ũ&°ƈ! KNOX

2012)KNOX2 Ǔ¸Þòǟò(KNOX1 ǟòİ#KNOX2 ǟòİ&ºĂ#ĥŝ'V~JY^'Õõ%$'Đ'ǁŀĔƾ"8 ȁ'÷
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ǟ  ò #KNOX2 ǟò(áǠ'o]?kv|<¿Ē8µǘ'V~JY^

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8Nz@b`Q`' KNOX ǟò( BELL ǟò##3&ƨò 'ºƺƗƣ'ƜġǿKNOX1Ȁ#ƗƣºÆǿKNOX2Ȁ&ŋƩȁKNOX
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K. Sakakibara and C. Furumizu- 6
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植物科学最前線 7:72 (2016)&¿Ē8ÒƩĕ ſÜ9(ßȅ; Furumizu et al. 2015)
 °&Ƽư<ČĚ8ņţ& ! KNOX1 ǟò Ƽư'ƼǱ<Ǚ8#<ǂÏ ȁq
XVcZIe`& 8ƼưČĚ&( KNOX1 ǟò Ĕƾ"7ȁKNOX1 ǟò'ǛÁŲũ(Ƽư<ŉĚ8þư'Łī<èÄ8(Hay and Tsiantis 2006)įȁqXVcZIe`" KNOX2 ǟò<ǛÁŲũ8#Ƽư'ƼǱ Ŝÿ8# 6ȁNz@b`Q`'áǠ'ČęČĚ#ÕŊ&ȁ
qXVcZIe`'ƼưČĚ& !3 KNOX2 ǟò( KNOX1 ǟò#Ðû'ĎÂ<ł#ſÜ9(Furumizu et al. 2015)96'Żƿ 6ȁƹòņţ&ƫ8Ƒƚ" KNOX1 ǟòİ#KNOX2 ǟòİ'ŋƩ íºÆ#ĥŝ98KNOX1 ǟò# KNOX2 ǟò'ŲũǸ
ã&Ǥ%7 %#, /Ǔ¸Þò#!ŋƩ'ǜ ĥŝ98# 6, ǟò'Ųũ<Ǉƍ8NPǸã#ǟò'ŭţ"8V|fGǍ #3&ëÆ#"ŋƩºÆ&ƫ#ƥ 69
8 /ȁƹòņţ"' KNOX ǟòİ'Ǔ¸Þò'ŋƩ'ëÆ<ï÷&ſ¢#!ȁNz@b`Q`4^p^"Õõ9ȁnsC_s@|<Ō KNOX ŊV|fGǍ KNATM 4 PTS/TKD1

Ģ

69896' DNA ƙÔƩ<ġ% KNOX ŊV|fGǍ(ǘć' KNOX Ǔ¸Þò'ŋƩ<ƊÔų&ǫøȁKNOX Ǔ¸Þò&58V~JY^ǟò'Ųũ¿Ē'ĝ¿Þò#!­#ſÜ9!8(Kimura et al. 2008; Magnani and Hake 2008)

[  W/9 W/9XRP9ACM907+02RZ5TD Nz@b`Q`Jdr&( 13 §' BELL ǟò ×/98
ŋƩŌĨëű 'ǟôų%ǁŀ&5
7, STM ǟò%$' KNOX1 ǟò( PENNYWISE 4 POUND-FOOLISH %$' BELL ǟò##3&ŋƩ8# ſ9!8(Byrne et al. 2003; Rutjens et al. 2009; Smith and Hake 2003)į, ŋƩŌĨëű 'Ʒũâ'Ǻ4ǛÁŲũņţ 'ǁŀ 6 KNOX2 ǟò( BELL1ȁSAWTOOTH1 (SAW1)ȁ
SAW2 %$' BELL ǟò##3&ŋƩ8ÒƩĕ ſÜ9! 7ȁNz@b`Q`' BELL ǟò&(ŋƩ'ºÆ 8# ĥŝ98(Furumizu et al. 2015)96'ŽƆ 6ȁBELL ǟòİ¶"'ǟòǤƼ#'Đ'ŋƩºÆ3 KNOX ǟò#'ŷ¡Ů'Ɨ0Ô;'ìŊÆ&ù#ƥ 698KNOX ǟò# BELL ǟò'Ɨ0Ô;'ìŊÆ 57ǾĊ%Ǔ¸ǇƍcY^{~G'ŉƏ<ÒƩ&ȁ'ƙłȁƹòņţ'áǠ'ƼǱ%o]?kv|'ǙÆ&3ǌŧ'"(% #ƥ 698["
 2014 Ĉ&´Ǩ9NuOGtƳǺGyjTxq]?Ar'Jdrėæ<×2Ƒƚǁŀ&57ȁ
KNOX ǟòİ(ǭņţ'ìŊÆ&°ƈ!ȁKNOX1 # KNOX2 'ǟòİ&º 9!
# ĵ6 #%ǿßȃ; Hori et al. 2014; Sakakibara 2016)NuOGtƳǺ"( (æØ9! 6ȁ 6ȁǭņţ'Ƒƚ"' KNOX1 # KNOX2 'ǟòİ"ĮǀŋƩŨđ

#&57ȁƨò 'ƼǱ%ìƔƨ ¿'žƈ&ŋƩ85 &%#ƥ 698NuOGtƳ
Ǻ& !96' KNOX1 ǟòȁKNOX2 ǟò 99$'5 %ŋƩ<ğ!8 (ƬK. Sakakibara and C. Furumizu- 7
BSJ-Review 7:72 (2016)

植物科学最前線 7:73 (2016)Ù ś  #  :   ǖ Ĉ ȁ N u O G t Ƴ Ǻ n N q _ z Ŷ ' g s q D [ F t Closterium
peracerosum-strigosum-littorale complex #V\jB Penium margaritaceum & !ČǍǓħƑ'žƈ
ŷō"æØ9!

7(Abe et al. 2011; Sørensen et al. 2014)ȁĐȁNuOGtƳǺ"'KNOX1/KNOX2 ǟòİ4 BELL ǟò'ŋƩǁŀ ď98
 /ȁƓƳ 63 KNOX ǟò# BELL ǟò 990 ! 7(Matsuzaki et al. 2004;
Mukherjee et al. 2009)ȁKNOX-BELL Ƙǐ'ǙÆ<ƥ 8"96'ŋƩǁŀ3+=ƬÙś
KNOX ǟòİ# BELL ǟòİ'ǏŞ< '.!08#ȁ96'ǟòİ(ƓƳ#ƞƭņţ'µǘƀ°"'ǟòǤƼ&5!Ŭ#ĥŝ9ȁǏŞ'ÑǟòǤƼ#'Đ'ǙÆ ņţ'o]?kv|4ǭņţ'ŬŗÓ'ǙÆ&í%čǴ< #ƥ 698 Ļƅ"(ŲŬǟôų%ŽƆ ¯÷!8ƞƳǺ#KIņţȁ!ƹòņţ'ŽƆ<ē&ƕ! ȁ'ºǺƤ& 8ǟòŋƩ'ǁŀ&Ö÷ǽƑ'Ĭ«3ǖĈȁǻǑų&Ǚ="8NWņţ& !(ũà´Ǩ9!8þưǺ'@bDVgeSelaginella moellendorffiiJdr
"%ȁƲ' NW"8wXu~_qQ{vh%$'Jdrǁŀ3Ǚ269! 7(Sessa et al.
2014)ȁwXu~_qQ{vh%$"ǟòý² ÒƩ%# æØ9 (Muthukumar et al. 2013;
Plackett et al. 2014)ĐȁNWņţ"'99'ǟòİ¶"'ǟòī ĵ6 #%7ȁ96'ǟò'ŋƩǁŀ Ǚ2698#"ȁƜƌļņţ"'KNOXǟòİ#BELLǟòİ'ǟòǤƼ#ŋƩºÆ<ŐǮų&Ǉ,8# ÒƩ#%8: /ȁũó'KIņţ't]xȁV@Ǻ'SaLIȁR|Ǻ'gsZwEcLI&Ä ȁƋ'KIņţ"8ZdLI't]xÆ3ñ/!
8(Szövényi et al. 2015)Đȁ96'ºǺƤ& 8BELL/KNOXǟò'ŋƩ4ǭņţ"Įǀ&Ũđ#ƥ 698ŋƩ_s@|'ǁŀ Ǚ1#ȁKNOXǟòİÏ*BELLǟòİ'ǟòǤƼ#9&ƛĮǀŋƩŨđ $'5 &ǭņţ'ČęǙÆ4ŬŗÓ'ǙÆ<36' '³
ǋ ĵ6 #%8: SU C~P^vw> Monash íô' John L. Bowman Ëê&ĲǶ 6'ĸŵ% discussion <Ęǈ/
:F?DAbe, J., Hori, S., Tsuchikane, Y., Kitao, N., Kato, M., & Sekimoto, H. 2011. Stable nuclear transformation of the
Closterium peracerosum-strigosum-littorale complex. Plant Cell Physiol. 52: 1676-1685.
Barton, M.K., & Poethig, R.S. 1993. Formation of the shoot apical meristem in Arabidopsis thaliana: an analysis
of development in the wildtype and in the shoot meristemless mutant. Development 119: 823-831.
Bharathan, G., Janssen, B.-J., Kellogg, E.A., & Sinha, N. 1999. Phylogenetic relationships and evolution of the
KNOTTED class of plant homeodomain proteins. Mol. Biol. Evol. 16: 553-563.
Blein, T., Pulido, A., Vialette-Guiraud, A., Nikovics, K., Morin, H., Hay, A., Johansen, I.E., Tsiantis, M., & Laufs,
P. 2008. A conserved molecular framework for compound leaf development. Science 322: 1835-1839.
Byrne, M.E., Barley, R., Curtis, M., Arroyo, J.M., Dunham, M., Hudson, A., & Martienssen, R.A. 2000.
Asymmetric leaves1 mediates leaf patterning and stem cell function in Arabidopsis. Nature 408: 967-971.

K. Sakakibara and C. Furumizu- 8
BSJ-Review 7:73 (2016)

植物科学最前線 7:74 (2016)

Byrne, M.E., Groover, A.T., Fontana, J.R., & Martienssen, R.A. 2003. Phyllotactic pattern and stem cell fate aredetermined by the Arabidopsis homeobox gene BELLRINGER. Development 130: 3941–3950.
Chuck, G., Lincoln, C., & Hake, S. 1996. KNAT1 induces lobed leaves with ectopic meristems when
overexpressed in Arabidopsis. Plant Cell 8: 1277-1289.
Derelle, R., Lopez, P., Le Guyader, H., & Manuel, M. 2007. Homeodomain proteins belong to the ancestral
molecular toolkit of eukaryotes. Evol. Dev. 9: 212-219.
Furumizu, C., Alvarez, J.P., Sakakibara, K., & Bowman, J.L. 2015. Antagonistic roles for KNOX1 and KNOX2
genes in patterning the land plant body plan following an ancient gene duplication. PloS Genet. 11:
e1004980.
Graham, L.E., Cook, M.E., & Busse, J.S. 2000. The origin of plants: body plan changes contibuting to a major
evolutionary radiation. Proc. Natl. Acad. Sci. USA 97: 4535-4540.
Hake, S., Smith, H.M.S., Holtan, H., Magnani, E., Mele, G., & Ramirez, J. 2004. The role of KNOX genes inplant development. Annu. Rev. Cell Dev. Biol. 20: 125-151.
Hareven, D., Gutfinger, T., Parnis, A., Eshed, Y., & Lifschitz, E. 1996. The making of a compound leaf: Genetic
manipulation of leaf architecture in tomato. Cell 84: 735-744.
Hay, A., & Tsiantis, M. 2006. The genetic basis for differences in leaf form between Arabidopsis thaliana and its
wild relative Cardamine hirsuta. Nat. Genet. 38: 942-947.
Hay, A., & Tsiantis, M. 2010. KNOX genes: versatile regulators of plant development and diversity. Development
137: 3153-3165.
Hori, K., Maruyama, F., Fujisawa, T., Togashi, T., Yamamoto, N., Seo, M., Sato, S., Yamada, T., Mori, H.,
Tajima, N., Moriyama, T., Ikeuchi, M., Watanabe, M., Wada, H., Kobayashi, K., Saito, M., Masuda, T.,
Sasaki-Sekimoto, Y., Mashiguchi, K., Awai, K., Shimojima, M., Masuda, S., Iwai, M., Nobusawa, T., Narise,T., Kondo, S., Saito, H., Sato, R., Murakawa, M., Ihara, Y., Oshima-Yamada, Y., Ohtaka, K., Satoh, M.,
Sonobe, K., Ishii, M., Ohtani, R., Kanamori, M., Honoki, R., Miyazaki, D., Mochizuki, H., Umetsu, J.,
Higashi, K., Shibata, D., Kamiya, Y., Sato, N., Nakamura, Y., Tabata, S., Ida, S., Kurokawa, K., & Ohta, H.
2014. Klebsormidium flaccidum genome reveals primary factors for plant terrestrial adaptation. Nat.
Commun. 5: 3978.
Horst, N.A., Katz, A., Pereman, I., Decker, E.L., Ohad, N., & Reski, R. 2016. A single homeobox gene triggers
phase transition, embryogenesis and asexual reproduction. Nat. Plants 2: 15209.
Kerstetter, R., Vollbrecht, E., Lowe, B., Veit, B., Yamaguchi, J., & Hake, S. 1994. Sequence analysis andexpression patterns divide the maize knotted1-like homeobox genes into two classes. Plant Cell 6:
1877-1887.
Kimura, S., Koenig, D., Kang, J., Yoong, F.Y., & Sinha, N. 2008. Natural variation in leaf morphology resultsfrom mutation of a novel KNOX gene. Curr. Biol. 18: 672-677.
Kumaran, M.K., Bowman, J.L., & Sundaresan, V. 2002. YABBY polarity genes mediate the repression of
KNOX homeobox genes in Arabidopsis. Plant Cell 14: 2761-2770.
Lee, J.-H., Lin, H., Joo, S., & Goodenough, U. 2008. Early sexual origins of homeoprotein heterodimerization andevolution of the plant KNOX/BELL family. Cell 133: 829-840.

K. Sakakibara and C. Furumizu- 9
BSJ-Review 7:74 (2016)

植物科学最前線 7:75 (2016)

Lewis, L.A., & McCourt, R.M. 2004. Green algae and the origin of land plants. Am. J. Bot. 91: 1535-1556.
Li, E., Wang, S., Liu, Y., Chen, J.-G., & Douglas, C.J. 2011. OVATE FAMILY PROTEIN4 (OFP4) interaction
with KNAT7 regulates secondary cell wall formation in Arabidopsis thaliana. Plant J. 67: 328-341.
Li, E., Bhargava, A., Qiang, W., Friedmann, M.C., Forneris, N., Savidge, R.A., Johnson, L.A., Mansfield, S.D.,
Ellis, B.E., & Douglas, C.J. 2012. The Class II KNOX gene KNAT7 negatively regulates secondary wall
formation in Arabidopsis and is functionally conserved in Populus. New Phytol. 194: 102-115.
Long, J.A., Moan, E.I., Medford, J.I., & Barton, M.K. 1996. A member of the KNOTTED class of homeodomain
proteins encoded by the STM gene of Arabidopsis. Nature 163: 66-69.
Magnani, E., & Hake, S. 2008. KNOX lost the OX: The Arabidopsis KNATM gene defines a novel class of KNOX
transcriptional regulators missing the homeodomain. Plant Cell 20: 875-887.
Markel, H., Chandler, J., & Werr, W. 2002. Translational fusions with the engrailed repressor domain efficiently
convert plant transcription factors into dominant-negative functions. Nucleic Acids Res. 30: 4709-4719.
Matsuzaki, M., Misumi, O., Shin-i, T., Maruyama, S., Takahara, M., Miyagishima, S., Mori, T., Nishida, K.,
Yagisawa, F., Nishida, K., Yoshida, Y., Nishimura, Y., Nakao, S., Kobayashi, T., Momoyama, Y.,
Higashiyama, T., Minoda, A., Sano, M., Nomoto, H., Oishi, K., Hayashi, H., Ohta, F., Nishizaka, S., Haga,
S., Miura, S., Morishita, T., Kabeya, Y., Terasawa, K., Suzuki, Y., Ishii, Y., Asakawa, S., Takano, H., Ohta,
N., Kuroiwa, H., Tanaka, K., Shimizu, N., Sugano, S., Sato, N., Nozaki, H., Ogasawara, N., Kohara, Y., &Kuroiwa, T. 2004. Genome sequence of the ultrasmall unicellular red alga Cyanidioschyzon merolae 10D.
Nature 428: 653-657.
Mele, G., Ori, N., Sato, Y., & Hake, S. 2003. The knotted1-like homeobox gene BREVIPEDICELLUS regulates
cell differentiation by modulating metabolic pathways. Genes Dev. 17: 2088-2093.
Mukherjee, K., Brocchieri, L., & Bürglin, T.R. 2009. A comprehensive classification and evolutionary analysis ofplant homeobox genes. Mol. Biol. Evol. 26: 2775-2794.
Muthukumar, B., Joyce, B.L., Elless, M.P., & Stewart Jr., C.N. 2013. Stable Transformation of Ferns Using
Spores as Targets: Pteris vittata and Ceratopteris thalictroides. Plant Physiol. 163: 648-658.
Ohno, S. 1970. Evolution by gene duplication. Springer-Verlag. New York.
Plackett, A.R.G., Huang, L., Sanders, H.L., & Langdale, J.A. 2014. High-efficiency stable transformation of themodel fern species Ceratopteris richardii via microparticle bombardment. Plant Physiol. 165: 3-14.
Reiser, L., Sanchez-Baracaldo, P., & Hake, S. 2000. Knots in the family tree: evolutionary relationships andfunctions of knox homeobox genes. Plant Mol. Biol. 42: 151-166.
Rensing, S.A., Lang, D., Zimmer, A.D., Terry, A., Salamov, A., Shapiro, H., Nishiyama, T., Perroud, P.F.,
Lindquist, E.A., Kamisugi, Y., Tanahashi, T., Sakakibara, K., Fujita, T., Oishi, K., Shin-I, T., Kuroki, Y.,
Toyoda, A., Suzuki, Y., Hashimoto, S., Yamaguchi, K., Sugano, S., Kohara, Y., Fujiyama, A., Anterola, A.,
Aoki, S., Ashton, N.W., Barbazuk, W.B., Barker, E., Bennetzen, J.L., Blankenship, R., Cho, S.H., Dutcher,
S.K., Estelle, M., Fawcett, J.A., Gundlach, H., Hanada, K., Heyl, A., Hicks, K.A., Hughes, J., Lohr, M.,
Mayer, K., Melkozernov, A., Murata, T., Nelson, D.R., Pils, B., Prigge, M., Reiss, B., Renner, T., Rombauts,
S., Rushton, P.J., Sanderfoot, A., Schween, G., Shiu, S.-H., Stueber, K., Theodoulou, F.L., Tu, H., Van dePeer, Y., Verrier, P.J., Waters, E., Wood, A., Yang, L., Cove, D.J., Cuming, A.C., Hasebe, M., Lucas, S.,

K. Sakakibara and C. Furumizu- 10
BSJ-Review 7:75 (2016)

植物科学最前線 7:76 (2016)

Mishler, B.D., Reski, R., Grigoriev, I.V., Quatrano, R.S., & Boore, J.L. 2008. The Physcomitrella genome
reveals evolutionary insights into the conquest of land by plants. Science 319: 64-69.
Rensing, S.A. 2014. Gene duplication as a driver of plant morphogenetic evolution. Curr. Opin. Plant Biol. 17:
43-48.
Rutjens, B., Bao, D., Van Eck-Stouten, E., Brand, M., Smeekens, S., & Proveniers, M. 2009. Shoot apicalmeristem function in Arabidopsis requires the combined activities of three BEL1-like homeodomain proteins.
Plant J. 58: 641-654.
Sakakibara, K., Nishiyama, T., Deguchi, H., & Hasebe, M. 2008. Class 1 KNOX genes are not involved in shootdevelopment in the moss Physcomitrella patens but do function in sporophyte development. Evol. Dev. 10:555-566.
Sakakibara, K., Ando, S., Yip, H.K., Tamada, Y., Hiwatashi, Y., Murata, T., Deguchi, H., Hasebe, M., &Bowman, J.L. 2013. KNOX2 transcription factors regulate the haploid to diploid morphological transition inland plants. Science 339: 1067-1070.
ňÍĖò 2013. KIņţ'Bo]o 6ƿ !ƨò 'ƼǱÆ. =%Čȁ=%" ―Bo]oŽƆķÀƠȇņţƎȈ.  1: 39-44.
Sakakibara, K. 2016. Technological innovations give rise to a new era of plant evolutionary developmental
biology. In: Rensing, S.A. (ed.) Genomes and evolution of charophytes, bryophytes, club mosses and ferns.
Elsevier Limited, Oxford. pp. 3-35.
Sano, R., Juárez, C.M., Hass, B., Sakakibara, K., Ito, M., Banks, J.A., & Hasebe, M. 2005. KNOX homeobox
genes potentially have similar function in both diploid unicellular and multicellular meristems, but not in
haploid meristems. Evol. Dev. 7: 69-78.
Serikawa, K.A., Martinez-Laborda, A., Kim, H.S., & Zambryski, P.C. 1997a. Localization of expression of
KNAT3, a class 2 knotted1-like gene. Plant J. 11: 853-861.
Serikawa, K.A., & Zambryski, P.C. 1997b. Domain exchanges between KNAT3 and KNAT1 suggest specificityof the kn1-like homeodomains requires sequences outside of the third helix and N-terminal arm of thehomeodomain. Plant J. 11: 863-869.
Sessa, E.B., Banks, J.A., Barker, M.S., Der, J.P., Duffy, A.M., Graham, S.W., Hasebe, M., Langdale, J., Li, F.-W.,
Marchant, D.B., Pryer, K.M., Rothfels, C.J., Roux, S.J., Salmi, M.L., Sigel, E.M., Soltis, D.E., Soltis, P.S., &Stevenson, D.W., Wolf, P.G. 2014. Between two fern genomes. Gigascience 3: 15.
Shani, E., Burko, Y., Ben-Yaakov, L., Berger, Y., Amsellem, Z., Goldshmidt, A., Sharon, E., & Ori, N. 2009.
Stage-specificregulationofSolanum

lycopersicum

leaf

maturation

by

class

1

KNOTTED1-LIKEHOMEOBOX proteins. Plant Cell 21: 3078-3092.
Smith, H.M.S., & Hake, S. 2003. The interaction of two homeobox genes, BREVIPEDICELLUS andPENNYWISE, regulates internode patterning in the Arabidopsis inflorescence. Plant Cell 15: 1717-1727.
Sørensen, I., Fei, Z., Andreas, A., Willats, W.G.T., Domozych, D.S., & Rose, J.K.C. 2014. Stable transformation
and reverse genetic analysis of Penium margaritaceum: a platform for studies of charophyte green algae, theimmediate ancestors of land plants. Plant J. 77: 339-351.
Szövényi, P., Frangedakis, E., Ricca, M., Quandt, D., Wicke, S., & Langdale, J.A. 2015. Establishment of

K. Sakakibara and C. Furumizu- 11
BSJ-Review 7:76 (2016)

植物科学最前線 7:77 (2016)

Anthoceros agrestis as a model species for studying the biology of hornworts. BMC Plant Biol. 15: 98.
Truernit, E., Siemering, K.R., Hodge, S., Grbic, V., & Haseloff, J. 2006. A map of KNAT gene expression in the
Arabidopsis root. Plant Mol. Biol. 60: 1-20.
Vollbrecht, E., Veit, B., Sinha, N., & Hake, S. 1991. The developmental gene Knotted-1 is a member of a maize
homeobox gene family. Nature 350: 241-243.
Wellman, C.H., Osterloff, P.L., & Mohiuddin, U. 2003. Fragments of the earliest land plants. Nature 425:282-285.
Wickett, N.J., Mirarab, S., Nguyen, N., Warnow, T., Carpenter, E., Matasci, N., Ayyampalayam, S., Barker, M.S.,
Burleigh, J.G., Gitzendanner, M.A., Ruhfel, B.R., Wafula, E., Der, J.P., Graham, S.W., Mathews, S.,
Melkonian, M., Soltis, D.E., Soltis, P.S., Miles, N.W., Rothfels, C.J., Pokorny, L., Shaw, A.J., DeGironimo,
L., Stevenson, D.W., Surek, B., Villarreal, J.C., Roure, B., Philippe, H., DePamphilis, C.W., Chen, T.,
Deyholos, M.K., Baucom, R.S., Kutchan, T.M., Augustin, M.M., Wang, J., Zhang, Y., Tian, Z., Yan, Z., Wu,
X., Sun, X., Wong, G.K.-S., & Leebens-Mack, J. 2014. Phylotranscriptomic analysis of the origin and early
diversification of land plants. Proc. Natl Acad. Sci. USA 111: E4859-E4868.
Zhao, H., Lu, M., Singh, R., Snell, & W.J. 2001. Ectopic expression of a Chlamydomonas mt+-specifichomeodomain protein in mt gametes initiates zygote development without gamete fusion. Genes Dev. 15:
2767-2777.
Zhong, R.Q., Lee, C.H., Zhou, J., L., McCarthy, R.L., & Ye, Z.H. 2008. A battery of transcription factors involvedin the regulation of secondary cell wall biosynthesis in Arabidopsis. Plant Cell 20: 2763-2782.

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