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식물의 엽록소 생합성 및 조절 기작에 대한 주요 연구동향

김다혜1, 양주희1, 김현정2, 이주희1, 이종렬1,*, 임선형2,*

Recent Advances in Genetic Regulation of Chlorophyll Metabolism in Plants

Da-Hye Kim1, Ju-Hee Yang1, Hyoun-Joung Kim2, JuHee Rhee1, Jong-Yeol Lee1,*, Sun-Hyung Lim2,*
Korean Journal of Breeding Science 2020;52(4):281-296.
Published online: December 1, 2020

1농촌진흥청 국립농업과학원

2국립한경대학교 원예생명공학과

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*Corresponding Author: Sun-Hyung Lim (E-mail: limsh2@hknu.ac.kr, Tel: +82-31-670-5105, Fax: +82-31-670-5109)
*Co-corresponding Author: Jong-Yeol Lee (E-mail: jy0820@korea.kr, Tel: +82-63-238-4616, Fax: +82-63-238-4602)
• Received: August 14, 2020   • Revised: September 24, 2020   • Accepted: October 4, 2020

Copyright © 2020 by the Korean Society of Breeding Science

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Recent Advances in Genetic Regulation of Chlorophyll Metabolism in Plants
Korean. J. Breed. Sci.. 2020;52(4):281-296.   Published online December 1, 2020
DOI: https://doi.org/10.9787/KJBS.2020.52.4.281
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Recent Advances in Genetic Regulation of Chlorophyll Metabolism in Plants
Korean. J. Breed. Sci.. 2020;52(4):281-296.   Published online December 1, 2020
DOI: https://doi.org/10.9787/KJBS.2020.52.4.281
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Recent Advances in Genetic Regulation of Chlorophyll Metabolism in Plants
Image Image Image Image
Fig. 1 Schematic representation of chlorophyll metabolic pathways in Arabidopsis. The dotted grey lines represent the individual four distinct pathways. The abbreviations of enzyme names and metabolites are as follows: 1) Common pathway; GluRS, glutamyl-tRNA synthetase; GluTR, glutamate-tRNA reductase; GSA, glutamate 1-semialdehyde aminotransferase; ALAD, 5-aminolevulinic acid dehydratase; PBGD, porphobilinogen deaminase; UROS, uroporphyrinogen III synthase; UROD, uroporphyrinogen III decarboxylase; CPO, coprophyrinogen III oxidase; Glu, glutamate; GSA, Glu 1-semialdehyde; ALA, 5-aminolevulinic acid; PBG, monopyrrole porphobilinogen; HMB, hydrocymethylbilane; Urogen III, uroporphyrinogen III; Coprogen III, coprophyrinogen III; FC, ferrochelatase. 2) Chlorophyll specific biosynthetic pathway; PPO, protoporphyrinogen IX oxidase; CHLD/CHLH/CHLI, Mg-chelatase subunit D/H/I; GUN4, genomes uncoupled 4; CHLM, Mg-protoporphyrin IX methyltransferase; CHL27/LCAA, low chlorophyll accumulation A, Mg-protoporphyrin IX monomethylester cyclase subunit; Mg-protoporphyrin IX monomethyl ester cyclase; POR, NADPH:protochlorophyllide oxidoreductase; DVR, 3,8-divinyl-chlorophyllide reductase; CHLP, geranylgeranyl reductase; CHLG, chlorophyll synthase; Protogen IX, protoporphyrinogen IX; Proto IX, protoporphyrin IX; Mg-Proto IX, Mg-protoporphyrin IX; Mg-ProtoME, Mg-protoporphyrin IX monomethylester; DV-Pchlide, protochlorophyllide; DV-Chlide, 3,8-divinyl-chlorophyllide; MV Chlide a, 3-vinyl chlide a; GGPP, geranylgeranyl pyrophosphate; Phytyl-PP, phytyl pyrophosphate. 3) Chlorophyll cycle pathway; Chl(ide) a, chlorophyll(ide) a; HMChl (ide) a, 7-hydrocymethyl chlorophyll(ide) a; Chl(ide) b, chlorophyll (ide) b; HMChl a, 7-hydrocymethyl chlorophyll a. 4) Chlorophyll degradation pathway; RCC, red chlorophyll catabolite; pFCC, primary fluorescent chlorophyll catabolites; mFCC, modified primary fluorescent chlorophyll catabolites; NCC, nonfluorescent chlorophyll catabolite.
Fig. 2 Schematic diagram of chlorophyll cycle. R indicates either protons or phytols. CAO, chlorophyllide a oxygenase; CBR, chlorophyll b reductase; HCAR, 7-hydroxymethyl chlorophyll a reductase.
Fig. 3 Proposed route of chlorophyll degradation. Chlorophyll a is hydrolyzed by chlase to from chlorophyllide and phytol. Also, chlorophyll is released of Mg by Mg-dechelatase. Chlase, chlorophyllase; SGR, stay-green; PPH, pheophytinase; PAO, pheophorbide a oxygenase; RCCR, red chlorophyll catabolite reductase; pFCC, primary fluorescent chlorophyll catabolites; NCC, nonfluorescent chlorophyll catabolite.
Fig. 4 Chlorophyll biosynthetic pathway controlled by transcription factors in day (A) and night (B). Bold letters and lines represent activated status. Solid arrows and solid bars indicate the positive and negative regulation, respectively. Dotted lines indicate indirect effects (Behringer & Schwechheimer 2015, Chattopadhyay et al. 1998, Goslings et al. 2004, Leivar & Quail 2011, Ma et al. 2014, Matsumoto et al. 2004, Tamai et al. 2002, Tang et al. 2012, Waters et al. 2009, Xu et al. 2015, Zhu et al. 2015).

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Fig. 4

Recent Advances in Genetic Regulation of Chlorophyll Metabolism in Plants

Enzymes involved in chlorophyll biosynthetic pathway.

Metabolic pathway Enzyme name Arabidopsis Rice Reference
Common pathway GluTR At1g58290 Os10g0502400 McCormac et al. 2001, Zheng et al. 2012
At1g09940
At2g31250
GSA1 At5g63570 Os08g0532200 Dalal & Tripathy 2012, Ilag et al. 1994
GSA2 At3g48730
ALAD At1g69740 Os06g0704600 Dalal & Tripathy 2012, Tang et al. 2012
At1g44318
PBGD At5g08280 Os02g0168800 Lim et al. 1994, Parker et al. 2006
UROS At2g26540 Os03g0186100 Li et al. 2017, Tan et al. 2008
UROD1 At3g14930 Os01g0622300 Tanaka et al. 2011, Zheng et al. 2013
UROD2 At2g40490 Os03g0337600
CPO1 At1g03475 Os04g0610800 Ishikawa et al. 2001, Matsumoto et al. 2004, Sun et al. 2011
CPO2 At4g03205
Chlorophyll specific
biosynthetic pathway		 PPO1 At4g01690 Os01g0286600 Park et al. 2012, Phung et al. 2011, Zhang et al. 2014
PPO2 At5g14220 Os04g0490100
CHLD At1g08520 Os03g0811100 Kobayashi et al. 2008, Zhang et al. 2016
CHLH At5g13630 Os03g0323200 Adhikari et al. 2011, Jung et al. 2003
CHLI1 At4g18480 Os03g0563300 Kobayashi et al. 2008, Zhang et al. 2016
CHLI2 At5g45930
GUN4 At3g59400 Os11g0267000 Li et al. 2014, Richter et al. 2016
CHLM At4g25080 Os06g0132400 Richter et al. 2016, Wang et al. 2017
CHL27 At3g56940 Os01g0279100 Rzeznicka et al. 2005, Wang et al. 2017
LCAA At5g58250 Os03g0331600 Albus et al. 2012, Kong et al. 2016
PORA At5g54190 Os04g0678700 Eckhardt et al. 2004, Sakuraba et al. 2013
PORB At4g27440 Os10g0496900
PORC At1g03630
DVR At5g18660 Os03g0351200 Eckhardt et al. 2004, Wang et al. 2010
CHLP At1g74470 Os01g0265000 Kimura et al. 2018, Takahashi et al. 2014
CHLG At3g51820 Os05g0349700 Lin et al. 2014, Zhou et al. 2017
Chlorophyll cycle CAO At1g44446 Os10g0567100 Espineda et al. 1999, Lee et al. 2005
CBR At5g17770 Os05g0488900 Horie et al. 2009, Sato et al. 2009
HCAR At1g04620 Os04g0320100 Meguro et al. 2011, Piao et al. 2017
Chlorophyll
degradation
pathway		 Chlase1 At1g19670 Os10g0419600 Benedetti & Arruda 2002, Sato et al. 2007
Chlase2 At5g43860
SGR1 At4g22920 Os09g0532000 Jiang et al. 2007, Rong et al. 2013, Shimoda et al. 2016
SGR2 At4g11910
SGRL At1g44000 Os04g0692600
PPH At5g13800 Os06g0354700 Morita et al. 2009, Schelbert et al. 2009
PAO At3g44880 Os03g0146400 Pružinská et al. 2003, Tang et al. 2011
RCCR At4g37000 Os10g0389300 Pružinská et al. 2007, Tang et al. 2011

Transcription factors related on chlorophyll biosynthetic pathway.

Regulatory pattern Transcription factor Arabidopsis Rice Reference
Positive HY5 At5g11260 Os06g0601500 Burman et al. 2018, Toledo-Ortiz et al. 2014
Positive GLK1 At2g20570 Os06g0348800 Nakamura et al. 2009, Wang et al. 2013, Waters et al. 2009
GLK2 At5g44190 Os01g0239000
Positive GNC At5g56860 Os06g0571800 Hudson et al. 2011, Kusano et al. 2020
Positive CGA1/GNL At4g26150
Positive FHY3 At3g22170 Os03g0181600 Reyes et al. 2004, Tang et al. 2012
Positive FAR1 At4g15090 Os07g0615900
Positive RVE At5g17300 Os02g0685200 Gray et al. 2017, Xu et al. 2015
Negative PIF1 At2g20180 OS04g6618600 Li et al. 2016, Zhu et al. 2015
PIF3 At1g09530 Os01g0286100 Leivar & Quail 2011, Zhou et al. 2013
PIF4 At2g43010 Os03g0782500 Leivar et al. 2009, Piao et al. 2015
PIF5 At3g59060 Os12g0610200 Leivar et al. 2009, Nakamura et al. 2007
Negative FLU At3g14110 Os01g0510600 Kong et al. 2016, Meskauskiene et al. 2001
Negative SCL27 At2g45160 - Ma et al. 2014
Negative TOC1 At5g61380 Os02g0618200 Legnaioli et al. 2009, Murakami et al. 2007
Table 1 Enzymes involved in chlorophyll biosynthetic pathway.
Table 2 Transcription factors related on chlorophyll biosynthetic pathway.

- : It is unidentified in rice genome.

Table 1

Table 2

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