植物科学最前線 4: 45( 2013)

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812-8581 ƞđƓƞđėőÄƫĒ 6-10-1
Molecular mechanism of chloroplast photorelocation movement in land plants
Key words: actin, Arabidopsis, chloroplast movement, organelle movement, phototropin
Noriyuki Suetsugu & Masamitsu Wada
Department of Biology, Faculty of Sciences, Kyushu University
Hakozaki 6-10-1, Higashi-ku, Fukuoka 812-8581, Japan

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N. Suetsugu & M. Wada - 1

BSJ- Revi ew 4: 45( 2013)

植物科学最前線 4: 46( 2013)2009ȨȧèȭȩȮȨ    b{X\:[i.8Z_fX
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BSJ- Revi ew 4: 46( 2013)

植物科学最前線 4: 47( 2013)36ǉƸǺÁ:1ŚŽ:Û/³Ƞƽ ȩ%Ǧ ç-71%&`CHzo:ƁĦ³Ƞƽ 6
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Oz?_]R]ȧArabidopsis thalianaȨ& phot1 ! phot2 %ȭ%gB[[zf|:ĸphot1 ! phot2 &!1$ȖÖÎī:¹ħȧSakai et al. 2001Ȩ
ȩ$ phot2

ǲȁÎī:¹ħ6ȧJarillo et al. 2001, Kagawa et al. 2001Ȩ
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ȧèȯȨdpZwE`LJȧPhyscomitrella

patensȨ$&ȭƢȠȧPpphotA ! PpphotBȨȱ%gB[[zf| üé6 ȧKasahara et al.
2004, Rensing et al., 2008ȨȩǚŒ7ȯȧPpphotA1, PpphotA2, PpphotB1, PpphotB2Ȩ&
71ȖÖÎī!ǲȁÎī:¹ħ

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ȧèȯȨ

N. Suetsugu & M. Wada - 3

BSJ- Revi ew 4: 47( 2013)

植物科学最前線 4: 48( 2013)        b{~pt2*+8Z_fXBCFM șǆƗÉ&șǆ©ÎīȩǦǆƗÉ&Ǧǆ©Îī:Ɲāƹ&ǝń75ȩƛƹ&ǝń7# Ôǀĭ ȤdNCw&ĵę#ĖõǉƸ:Ƴƿ­$ĸȩğ©
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2005ȨȩgB[[zf|&ŎǥƎ$ȖÖÎī!ǲȁÎī% Ŀ:¹ħ6Šǀ:ĸ63
 6āȕ$ȩOz?_]R]% phot2 òƊ 1țĘ$ġ© ǲȁÎī ǡĉ76
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BSJ- Revi ew 4: 48( 2013)

植物科学最前線 4: 49( 2013)

ȌŦȔ!Ǜ6gB[[zf|%=naȅȃµ$36³úƭƵǚŒ 4ȩgB[[zf|Ȁú%Ȇǔ&77%³Ƞƽ ƀƦ$ǧ 4'ƢúŚŽ% phot1 ! phot2 %Ȁ
úȆǔ&ǓúŚŽ!ǐúŚŽ%³Đ%» ǧ5ȩKJŚŽS|Ƞ &ƀǃ$ photA! photB $³ 7)K=GGRJSK;LPT268~Z^j3[i Ȃ%ĺÖǌȠȧdNCw! Mesotaenium caldariorumȨ
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2007aȨ
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ȧèȭȨ
`CHzo&gB[[zf|%Šǀ\p?|:+¢ĸ
% ȩƳƿǂ$ČéǉƸǺÁ:¹ħ6g>[Hzo%ā 6Ôǀĭ ġ Ɲá7āȕ$ȩk@v?OW%Ǧǆ©$36©čĭ!ǉƸǺÁ Ŀ:šĽȧșǆ©Îī&ŤĘ 6ȨòƊ rapȧKadota & Wada 1999Ȩ&ȩǣ+ȏ5+%v?| `CHzoȀú$šĽ ȧKawai et al. 2003Ȩ
4$`CHzoȀú: rap òƊ$ĉª
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2003, Schneider et al. 2004Ȩ Ȣ + !$ȩĴ&ĺÖǌȠdNCw 41`CHzo:ƌǗȧSuetsugu et al.
2005bȨ
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N. Suetsugu & M. Wada - 5

BSJ- Revi ew 4: 49( 2013)

植物科学最前線 4: 50( 2013)>[Hzo1Ó

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1üé6ȧSparkes 2010, Deeks & Hussey 2005Ȩ ȩ%!8 Arp2/3 ǔÖ%ŚŽ%CxE`vǺÁ*%ȍ&ń4&#ċ# !1ƢúŚŽ$ȩƔŗƆŽ$¬Ǵ#CxE`vȧn[K|\w=ȩLxPȩjxCFOU~oȩĊƿȨ%ǺÁ&nCO|$ ü6! ňǱń4 $#ȧSparkes 2010Ȩ # 4ǉƸǺÁ&ȩArp2/3 ǔÖ%Šǀ šĽòƊ 1ŤĘ 5ȧKadota et al. 2009Ȩ
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N. Suetsugu & M. Wada - 6

BSJ- Revi ew 4: 50( 2013)

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N. Suetsugu & M. Wada - 7

BSJ- Revi ew 4: 51( 2013)

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6! ³ 6ȆǕ#!$ȩ=HX|ƻƷ ü%CxE`vǺÁ%ȎĂ¼!Ƙ476
2,3-hV|PC| 2-qaFOo ±ƄƳƿ &ȩǉƸǺÁ&þ«$ȎĂ76 ȩ7
1#ġ©ŹĈŅ$Ǘ476 cp-=HX|ƻƷ%ǻƎŲ÷!ż¦*%¥éÃ&ǙĆ7ȧYamada et al. 2011Ȩ
%ƴŔ&ȩcp-=HX|ƻƷ%ż¦*%¥é&ǉƸǺÁ%ƴŔ &# ȩǺÁ%ǧå!#5Ħ6!:Ɲ6 Ƴƿ­ Țţ6ǉƸ&Ƴƿǥ$ű6% &# ȩ=HX|ƻƷ$ üƳƿǂ$ĺƕ6 ȧTakagi et al. 2009Ȩȩ7 1%ï 9 $Á6Ƴƿ%ě
Ƭç$ğ©:ŹĈ6!ȩǉƸ%«Þç

cp-=HX|ƻƷ ñ¾7$ǉƸ%Á 1ğ-ǳ$ġ©ŹĈ & cp-=HX|ƻƷ Ų÷ȩǉƸ&õÁ 3$#

ȧKadota et al. 2009Ȩ
%!&ȩ©ġĝ$ ü cp-=HX|ƻƷ%Ȉ%òÃ Ƴƿ
ǂ*%ĺƕ%ơĝ:ŬĀ6!:Ɲá6
 ©$ ü cp-=HX|ƻƷ%ÁıòÃ!ǉƸ©ĀǺÁ!%ȍǷ&ȩgB[[zf|šĽòƊ:ǚŒ$354$ń4 !#phot1 òƊ & phot2 $ üȖ
ÖÎī!ǲȁÎī !1$ǧ5ȩcp-=HX|ƻƷ%Áı$1ȇƆë!Ũǭõ #ǽ
Ǘ47# ȧKadota et al. 2009Ȩ ġ©ŹĈ$3ǧ6 cp-=HX|ƻƷ%Ų
÷!%ĥ%¥éÃ ȇƆë!Ũǭ9 $łV?n|I ǧ5ȩ%ƴŔ!ǉƸ ǲȁù06V?n|I1ł ȧIchikawa et al. 2011Ȩ
phot2 òƊ&ǲȁÎī:š
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植物科学最前線 4: 53( 2013)Ľ6% ȩġ© 1ȖÖÎī ǡĉ76ȧJarillo et al. 2001, Kagawa et al. 2001Ȩ%òƊ%ǉƸ$ġ©:ŹĈ6!ȩġ© ŹĈ7Ȃ³ % cp-=HX|ƻƷ%Ų÷&ǧ4ȩǳ$ cp-=HX|ƻƷ%ȆÖ ¡Ǹ7ȩȖÖÎī ǡĉ7ȧIchikawa et al. 2011,
Kadota et al. 2009Ȩ
phot1phot2 òƊ & cp-=HX|ƻƷ%© üƎ#òÃ&« ǧ4ȩǉƸ%©ĀǺÁ1Ƴƿǂ*%ĺƕ%©¹ħ1Ǘ47# ȧIchikawa et al. 2011, Kadota
et al. 2009Ȩ
74%ƴŔ 4ȩgB[[zf|& cp-=HX|ƻƷ%Áı:¹ħ6!
ń4 !#  <AES|z3[i298ac
 Oz?_]R]$6³úȀýƎǚŒ$35ȩ©ÑăgB[[zf| # ǉƸǺÁ%£ÕǼƭ2ǺÁƭ$ȍ96Ş#åú ×Ā7ȧèȲȨĴ&ȩ74%å
ú%òƊ$ȩǉƸǺÁ$ cp-=HX|ƻƷ%Áı:ǟƳ$ǙĆ6!$35ȩ
74%åú

cp-=HX|ƻƷ%¹ħ$"%3$ȍ6 :ĻĆèȲȩOz?_]R]$6ǉƸǺÁ%£ÕǼƭ2ǺÁƭ$ȍ96åú) " #$%%#% #$ Ĵ ÇȗòƊ%

chup1 òƊ&ň1ģǥ ġ ȩǉƸ©ĀǺÁ:šĽ6 # ȩǉƸ Ƴƿǂ 4ó7-°ȖȧKasahara et al. 2002, Oikawa et
al. 2003Ȩ
ǉŕƳƿ:ǙĆ6!ȩȇƆë &ºŶ© ŹĈ7#ȏ5ǉƸ&%Ƽ:,!;"ò# ȩchup1 òƊ%ǉƸ&ÊģǥŰÁ$Ŷ Ƴƿ­:ƠÁȧKadota et al. 2009, Suetsugu et al. 2010cȨ
%òƊ &Ƴƿǥ%=HX|ƻƷ%Áı&ȇƆë!ò94# ȩcp-=HX|ƻƷ « .47# ȧOikawa et al. 2003, Kadota et al.
2009Ȩ
%!& cp-=HX|ƻƷ ȩǉƸ%©ĀǺÁ%.#4Ƴƿǂ*%ĺƕ$1Īȝ 6!:4$Ǒ6CHUP1 &ǔľ%Šǀ\p?|:ĸǉƸ%óÂǂ
N. Suetsugu & M. Wada - 9

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植物科学最前線 4: 54( 2013)$Čé6ȒŚŽ$žƊƎ#V|cHǥ 6ȧOikawa et al. 2003Ȩ
ȧèȲȨ
=naōƧ¦%ƋŪĭȞì&ǉƸóÂǂČé$Īȝ 6ȧOikawa et al. 2003, 2008, Schmidt von Braun &Schleiff 2008Ȩ%ĥ8¦%=naōƧ¦Å³%K?x\K?x\p?|&ȩǉƸ Ƴƿǂ$ĺƕ60$ĪǕ 5ȧOikawa et al. 2008Ȩ
ȩ-Ǟȡƪ­%āȡƴŔ &ȈÃ$1
ĪǕ 6ȧLehmann et al. 2011Ȩ
K?x\K?x%Ţ$&=HX|ƴÖȂ üéȩǞȡƪ­ %=HX|ƻƷƴÖǀ ƜǠ7ȧOikawa et al. 2003Ȩ
DxlFOōƧ¦&izw|
ťí$ą/Ȟì!ȒŚŽ% CHUP1 %Ȍ țĘ$¢üĭ%ȤȞì:Û/ȧOikawa et al.
2003Ȩ
izw|ťí$ą/Ȟì&ÁŽƩ%=HX|ȆÖ$ȍ96åú$1Ǘ47ȩ=HX|³ú!ƴÖȩ%ȆÖ:¡Ǹ6izg>w|!ƴÖ6Ȟì 6! Ƙ476ȧHolt & Koffer 2001Ȩ
āȕ$ CHUP1 &izg>w|ƴÖǀ:1ȧSchmidt von Braun &Schleiff 2008Ȩ
%! 4 CHUP1 &ǉƸ$Čéȩcp-=HX|ƻƷ%ȆÖ$ȍ6
åú 6Ôǀĭ Ȥ ȩ%!8%ǝķ&Ÿ) # "$"$#" #$&$ ȖÖÎī%òƊ%QHw~^|I%ǻơ
;"šĽ6#

kac1 òƊ Çȗ7 ȩȖÖÎī:,!

ȩǲȁÎī%ǵĝ1ȇƆë!Ũ+ǹ

2010cȨ
Oz?_]R]$& KAC1 $3 Ȁú KAC2ȧSuetsugu et al.

üé6kac2 òƊ%ǉƸǺÁ&ŤĘ  ȩkac1kac2 ȆòƊ &ǉƸ©ĀǺÁ « Ǘ47ȩǉƸƳƿǂ

4ó76! chup1 òƊ!3ģǥ:Ɲāȕ$ chup1 òƊ×Ş

kac1kac2 ȆòƊ & cp-=HX|ƻƷ « .47ȩkac1 òƊ% cp-=HX|ƻƷ%Ȉ&ȇƆë!Ũ+ċ# ȧSuetsugu et al. 2010cȨ
Ȣ + !$ȩKAC &ĩĊƪq~V~!Ƙ476F`O|Ş%V|cHǥ ȧèȲȨ
!8 %q~V~\p?|&ĩĊƪ$ƴÖ6ǀ½2q~V~ůĭ$Īȝ#_HyCX\¾Ū³ǚǀ:šĽ
KAC %DxlFOōƧ&%V|cHǥ!%Ƒ×ĭ&Ÿ ȩȒŚŽ% KAC %Ȍ țĘ$Ƒ×ĭȤ%Ȃ³&Ǟȡƪ­ğ#

41=HX|ƻƷ!ƴÖ6ǀ½

6ȧSuetsugu et al. 2010cȨ
KAC &Ƴƿǂ!Ƴƿǥ$Čé6% ȩcp-=HX|ƻƷ Ƴƿǂ$ƴÖ60$ĪǕ#åú 6!ƾ476)   "$"!%""" #$%%$"# #
 jac1 òƊ&ȖÖÎī:þ«$šĽ5ȧSuetsugu et al. 2005aȨ
ȩǲȁÎī&ȇƆë351ğ©ġĝ ǡĉ76 ȩ%ǵĝ&ǹȧIchikawa et al. 2011, Kodama et al. 2010ȨȖÖÎī:šĽ6% ȩğ© ©%ĿÙ$Ù  cp-=HX|ƻƷ%¥éÃ&Ǘ47# ȩȂ%ğ© 4ǲ6ǉƸ &ȩ©!Îć¦$ cp-=HX|ƻƷ ¥é6!
ǙĆ7ȧIchikawa et al. 2011ȨȜƍ!$ȩjac1 òƊ &ġ©ŹĈ$36 cp-=HX|ƻƷ%ǻƎŲ÷ ,!;"ǙĆ7# ž$Ƴƿ«$ġ©ŹĈ! $& cp=HX|ƻƷ%Ų÷1¥éÃ1ǙĆ7ȩƴŔ!ǉƸǲȁÎī&ǧ4ȩƳƿǂ$ġ ĺƕȧIchikawa et al. 2011Ȩ
JAC1 &DxlFOōƧ¦$³úOrjz|V|cH
ǥ$.476 J-\p?|!ß'76źOtYHV|cHǥ:¹ħ6\p?|:ĸ
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植物科学最前線 4: 55( 2013)ȧSuetsugu et al. 2005aȨ
ȧèȲȨž$C~FOw|!ß'76HvQw|Ċƿ:ǐǖ6V|
cHǥ$Ƒ×ĭ ȤȧEisenberg & Greene 2007Ȩāȕ$ƴŇŝǶǚŒ$35ȩJAC1 % J-\
p?|&C~FOw|% J-\p?|!×ƦŝǶ:ĸ!

³

ȧSuetsugu et al.

2010b, Takano et al. 2010Ȩ
JAC1 % J-\p?| ǉƸǺÁ%¹ħ$Īȝ 6!&ń4 ȧTakano et al. 2010ȨȩJAC1 ǃ%Šǀ&³ #) '" #$&$%"%$ 1   #$&$ " Ĵ&ǲȁÎīšĽòƊ%QHw~^|I$35ȩǲȁÎī Ǌ ğȭ%òƊ
web1 ! pmi2 :ÇȗȧKodama et al. 2010, Luesse et al. 2006Ȩ
74%òƊ &ȖÖÎīȩ
ǲȁÎī!1$ǉƸǺÁ%ǵĝ ǹ ȩž$ǲȁÎī%ƊĘ Ǌ āȕ$ġ©ŹĈ$36 cp-=HX|ƻƷ%Ų÷ ,!;"ǧ4ȩ 9 $¥éÃ ǧ6%. ȧKodama et al. 2010Ȩ òƊ%ģǥ&,!;"× 5ȩȆòƊ177%Ç
%òƊ!ò94#% ȩ
WEB1 ! PMI2 &țĘ$Ąĺ$Šǀ6! ĻĆ7
WEB1 ! PMI2 & ȩ ! 1 $ K ? x \ K ? x \ p ? | : ĸ   V | c H ǥ g < n w ~ȧWEB1/PMI2-related V|cHǥg<nw~Ȩ$ď6ȧKodama et al. 2010, 2011ȨȧèȲȨȀýƎǚŒ!V|cHǥ%ŝǶ 4į73$ȩȄŧZ~b?hwY\ƭȩŚŽƳƿ­%³úǍ©ƑǒȧBiFCȨŮ#"$35 WEB1 ! PMI2 &ƴÖ6!ȩ4$ BiFC Ů$
3ȩ%ƴÖ Ƴƿǥ ǧ6! ǙĆ7ȧKodama et al. 2010Ȩ
ȀýƎ
ǚŒ$35 WEB1 ! PMI2 %Šǀ$& JAC1

ĪǕ 6! Ɲá7 ȩWEB1/PMI2! JAC1 %ƴÖ&%!8ś²7#ȧKodama et al. 2010Ȩġ© % cp-=HX|
ƻƷ%Áı%ƊĘ ȩjac1ȩweb1ȩpmi2 %%òƊ 6! 4ȧIchikawa et al.
2011, Kodama et al. 2010ȨȩWEB1/PMI2 & JAC1 !Ǳ!8 Šǀ6Ôǀĭ 6)/3V ǜ%$1 PMI1ȧplastid movement impaired 1Ȩ
ȧDeBlasio et al. 2005Ȩ2 THRUMIN1ȧWhippo
et al. 2011Ȩ#" ǉƸǺÁ$ȍ6! Ƙ476 ȩ74%åú

cp-=HX|

ƻƷ%¹ħ$ȍ6 Ú &îÝ7#ž$ THRUMIN1 &Ƴƿ­ =HX|ƻƷ!ƴÖȩ4$Ǟȡƪ­ =HX|ƻƷ%ŏÃ:¡Ǹ6! Ƙ476% ȧWhippo
et al. 2011Ȩȩthrumin1 òƊ$6 cp-=HX|ƻƷ%ÁıǙĆ&Īȝ 6 *972 ǉƸǺÁ&ǁŗƳƿ 1ǡĉ76% ȩȀúƌƃ:#țĘ$łÇƱ#ƭ¹ħ76! Ɲá76ȧWada 1988ȨƸǆŚŽ&Ƴƿǂ$Čé6©Ñă
gB[[zf|$3ǉƸǺÁ:¹ħ6pD^Ro:ƌǼ6!$35ȧèȮȨȩȀúƌƃ:#ȩƳƿǂ!ǉƸȌ!ňƙǨȗ ǰǵ$±Ƅ

6Îīƭ:ƁĦƳƿǂǱ %£ÕǼ$ȍ&ȩDxO@o?C|%Ð5ǯ.Ʃ%ȍ Ɲá7
6 ȩŌƜǝ&#ȧSuetsugu et al. 2009Ȩ

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植物科学最前線 4: 56( 2013) ŚŽ Ȓ¶ŊȩȒŚŽ&ƐĺŁ©$476ƅð$ ȩŢƨ$ŋƆOWȩ
ǓúŚŽȩǐúŚŽƩ ƺǇ63$#6!ȩœĜ 35ô %©:Ħ6! ŚŽ%Ɔü
$Īȝ!#©ġĝ ȟƺ$òÁ6Ƌœ%œĜ ȩÀƂ3©Öĳ:Ǐȩ ©Ľ§
:ȁ4763ǿī60ȩŚŽ$!&ƶ©%ĿÙ2ġğ:ĆƘȩǉƸ ǿ´#Ƽ$Ʋł Á !

63#ǺÁpD^Ro ĪǕ ȧèȳȨnCO|$36=HX|ƻƷ$ŭǺÁ&ǵǵĝ:Ʒĸ
ŀü%=HX|ƻƷ$ŭĿÙ$ ƠÁ61%%ȧShimmen & Yokota, 2004ȨȩȩĬ#ĿÙǫļ&æȘ 6Ŀȩcp-=HX|ƻƷ$36ǺÁpD^Ro &ȩ=HX|ƻƷǃ ¤%ǉƸ$3ǃĤƎ$ȩ 1ƙŅȌ$®ŝĳ76% ȩ¤%ǉƸ&ƀƦ$Ƴƿ­:ǃƉ#ĿÙ$Á !
6ǉƸǺÁ$ cp-=HX|ƻƷ%© ü%Áı&Oz?_]R] # ȩKJȠ2OWȠ 1Ǘ476ȧTsuboi & Wada, 2012, Yamashita et al. 2011Ȩ
- cp-=HX|ƻƷ%ȆÖ6&¹ħ$ȆǕ#åú CHUP1 ! KAC &ȩƭƵýƎ$&ȒŚŽ%íȂ$Ƽ6
T^LJ$1üé6ȧSuetsugu et al. 2010aȨ
#9 cp-=HX|ƻƷ ü%ǉƸǺÁ%
pD^Ro&ȩŚŽ%Ȓ¶Ŋ$&ŀ$ƁĦ7ȩŶ òÁ6Ȓ%©ƅð ŚŽ ƺŖ60%õ #¿!#!ƾ476ȧèȮȩȳȨ bUpt2*+8>Q@IO\ƔŗŚŽ$¬Ǵ#CxE`v&ȩƳƿǥ$ŀü%=HX|ƻƷ$ŭnCO| ü%ǵǺÁ:Ǐ ȩ%ĿĞ &ĿÙǫļ ȘĿǉƸ&©%ġ2ĿÙ$ī cp-=H
X|ƻƷ%ȆÖ:ǃ³ǃǩ ¹ħ6!$35ȩƳƿ­:ǃƉ#ĿÙ$Á !

6©ÑăgB[[zf|! cp-=HX|ƻƷ%¹ħåú 6 CHUP1 ! KAC &ȒŚŽ$¬
Ǵ#åú 6ǉƸ "%3$ cp-=HX|ƻƷ:¸ƇǺÁ%ĻǸ½:Ɔ.²6 &ń4#

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