The Divergent Roles of STAYGREEN (SGR) Homologs in Chlorophyll Degradation

다국어 초록 (Multilingual Abstract)

Degradation of chlorophyll (Chl) by Chl catabolic enzymes (CCEs) causes the loss of green color that typically occurs during senescence of leaves. In addition to CCEs, STAYGREEN1 (SGR1) functions as a key regulator of Chl degradation. Although sgr1 mutants in many plant species exhibit a staygreen phenotype, the biochemical function of the SGR1 protein remains elusive. Many recent studies have examined the physiological and molecular roles of SGR1 and its homologs (SGR2 and SGR-LIKE) in Chl metabolism, finding that these proteins have different roles in different species.
In this review, we summarize the recent studies on SGR and discuss the most likely functions of SGR homologs.

참고문헌 (Reference)

1 Jakoby, M., "bZIP transcription factors in Arabidopsis" 7 : 106-111, 2002

2 Sattler, S. E., "Vitamin E is essential for seed longevity and for preventing lipid peroxidation during germination" 16 : 1419-1432, 2004

3 Sato, Y., "Two short-chain dehydrogenase/reductases, NON-YELLOW COLORING 1 and NYC1-LIKE, are required for chlorophyll b and light-harvesting complex II degradation during senescence in rice" 57 : 120-131, 2009

4 Fujita, Y., "Three SnRK2 protein kinases are the main positive regulators of abscisic acid signaling in response to water stress in Arabidopsis" 50 : 2123-2132, 2009

5 Park, S. Y., "The senescence-induced staygreen protein regulates chlorophyll degradation" 19 : 1649-1664, 2007

6 Chao, W. S., "The impact of chlorophyll-retention mutations, d1d2 and cyt-G1, during embryogeny in soybean" 107 : 253-262, 1995

7 Fraser, P. D., "The biosynthesis and nutritional uses of carotenoids" 43 : 228-265, 2004

8 Rong, H., "The Stay-Green Rice like(SGRL)gene regulates chlorophyll degradation in rice" 170 : 1367-1373, 2013

9 Olinares, P. D., "The Clp protease system; a central component of the chloroplast protease network" 1807 : 999-1011, 2011

10 Mach, J. M., "The Arabidopsis-accelerated cell death gene ACD2 encodes red chlorophyll catabolite reductase and suppresses the spread of disease symptoms" 98 : 771-776, 2001

11 Tanaka, R., "The Arabidopsis-accelerated cell death gene ACD1 is involved in oxygenation of pheophorbide a : inhibition of the pheophorbide a oxygenase activity does not lead to the"stay-green"phenotype in Arabidopsis" 44 : 1266-1274, 2003

12 Hörtensteiner, S., "Stay-green regulates chlorophyll and chlorophyll-binding protein degradation during senescence" 14 : 155-162, 2009

13 Sakuraba, Y., "STAY-GREEN and chlorophyll catabolic enzymes interact at light-harvesting complex II for chlorophyll detoxification during leaf senescence in Arabidopsis" 24 : 507-518, 2012

14 Kusaba, M., "Rice NON-YELLOW COLORING1 is involved in lightharvesting complex II and grana degradation during leaf senescence" 19 : 1362-1375, 2007

15 Kato, Y., "Protein quality control in chloroplast : a current model of D1 protein degradation in the photosystem II repair cycle" 146 : 463-469, 2009

16 Forsberg, J., "Protease activities in the chloroplast capable of cleaving an LHCII N-terminal peptide" 123 : 21-29, 2005

17 Sakuraba, Y., "Phytochrome-interacting transcription factors PIF4 and PIF5 induced leaf senescence in Arabidopsis" 5 : 4636-, 2014

18 Schelbert, S., "Pheophytin pheophorbide hydrolase(pheophytinase)is involved in chlorophyll breakdown during leaf senescence in Arabidopsis" 21 : 767-785, 2009

19 Horie, Y., "Participation of chlorophyll b reductase in the initial step of the degradation of light-harvesting chlorophyll a/b-protein complexes in Arabidopsis" 284 : 17449-17456, 2009

20 Jiang, H., "Overexpression of SGR results in oxidative stress and lesion-mimic cell death in rice seedlings" 253 : 375-387, 2011

21 Liang, C., "OsNAP connects abscisic acid and leaf senescence by fine-tuning abscisic acid biosynthesis and directly targeting senescence-associated genes in rice" 111 : 10013-10018, 2014

22 Yamatani, H., "NYC4, the rice ortholog of Arabidopsis THF1, is involved in the degradation of chlorophyll-protein complexes during leaf senescence" 74 : 652-662, 2013

23 Jiang, H., "Molecular cloning and function analysis of the stay green gene in rice" 52 : 197-209, 2007

24 Hirashima, M., "Light-independent cell death induced by accumulation of pheophorbide a in Arabidopsis thaliana" 50 : 719-729, 2009

25 Gray, J., "Light-dependent death of maize lls1 cells is mediated by mature chloroplasts" 130 : 1894-1907, 2002

26 Lim, P. O., "Leaf senescence" 58 : 115-136, 2007

27 Tang, Y., "Knockdown of OsPAO and OsRCCR1 cause different plant death phenotypes in rice" 168 : 1952-1959, 2011

28 Cha, K. W., "Isolation, characterization, and mapping of the stay green mutant in rice" 104 : 526-532, 2002

29 Giraudat, J., "Isolation of the Arabidopsis ABI3 gene by positional cloning" 4 : 1251-1261, 1992

30 Wei, Q., "Isolation and characterization of a chlorophyll degradation regulatory gene from tall fescue" 30 : 1201-1207, 2011

31 Pružinská, A., "In vivo participation of red chlorophyll catabolite reductase in chlorophyll breakdown" 19 : 369-387, 2007

32 Koornneef, M., "In vivo inhibition of seed development and reserve protein accumulation in recombinants of abscisic acid biosynthesis and responsiveness mutants in Arabidopsis thaliana" 90 : 463-469, 1989

33 Meguro, M., "Identification of the 7-hydroxymethyl chlorophyll a reductase of the chlorophyll cycle in Arabidopsis" 23 : 3442-3453, 2011

34 Ren, G., "Identification of a novel chloroplast protein AtNYE1 regulating chlorophyll degradation during leaf senescence in Arabidopsis" 144 : 1429-1441, 2007

35 Zhou, C., "From model to crop : functional analysis of a STAY-GREEN gene in the model legume Medicago truncatula and effective use of the gene for alfalfa improvement" 157 : 1483-1496, 2011

36 Thomas, H., "Five ways to stay green" 51 : 329-337, 2000

37 Johnson-Flanagan, A. M., "Ethylene production during development of mustard(Brassica juncea)and canola(Brassica napus)seed" 106 : 601-606, 1994

38 Wang, Q., "Deletion of the chloroplastlocalized Thylakoid formation1 gene product in Arabidopsis leads to deficient thylakoid formation and variegated leaves" 136 : 3594-3604, 2004

39 Sakuraba, Y., "Delayed degradation of chlorophylls and photosynthetic proteins in Arabidopsis autophagy mutants during stress-induced leaf yellowing" 65 : 3915-3925, 2014

40 Morita, R., "Defect in non-yellow coloring 3, an alpha/beta hydrolase-fold family protein, causes a stay-green phenotype during leaf senescence in rice" 59 : 940-952, 2009

41 Armstead, I., "Cross-species identification of Mendel's/locus" 315 : 73-, 2007

42 Fang, C., "Concerted evolution of D1 and D2 to regulate chlorophyll degradation in soybean" 77 : 700-712, 2014

43 Nakagawara, E., "Clp protease controls chlorophyll b synthesis by regulating the level of chlorophyllide a oxygenase" 49 : 800-809, 2007

44 Zhang, W., "Chlorophyll degradation : the tocopherol biosynthesis-related phytol hydrolase in Arabidopsis seeds is still missing" 166 : 70-79, 2014

45 Hörtensteiner, S., "Chlorophyll breakdown in higher plants" 1807 : 977-988, 2011

46 Pružinská, A., "Chlorophyll breakdown : pheophorbide a oxygenase is a Rieske-type iron-sulfur protein, encoded by the accelerated cell death 1 gene" 100 : 15259-15264, 2003

47 Nakajima, S., "Chlorophyll b reductase plays an essential role in maturation and storability of Arabidopsis seeds" 160 : 261-273, 2012

48 Clerkx, E. J., "Characterization of green seed, an enhancer of abi3-1 in Arabidopsis that affects seed longevity" 132 : 1077-1084, 2003

49 Guiamet, J. J., "Characterization of cytoplasmic and nuclear mutations affecting chlorophyll and chlorophyll-binding proteins during senescence in soybean" 96 : 227-231, 1991

50 Biswal, B., "Carotenoid catabolism during leaf senescence and its control by light" 30 : 3-13, 1995

51 Fraser, P. D., "Carotenoid biosynthesis during tomato fruit development(evidence for tissue-specific gene expression)" 105 : 405-413, 1994

52 Huang, W., "Arabidopsis thylakoid formation 1 is a critical regulator for dynamics of PSII-LHCII complexes in leaf senescence and excess light" 6 : 1673-1691, 2013

53 Sakuraba, Y., "Arabidopsis STAYGREEN-LIKE(SGRL)promotes abiotic stress-induced leaf yellowing during vegetative growth" 588 : 3830-3837, 2014

54 Sakuraba, Y., "Arabidopsis STAY-GREEN2is a negative regulator of chlorophyll degradation during leaf senescence" 7 : 1288-1302, 2014

55 Barry, C. S., "Amino acid substitutions in homologs of the STAY-GREEN protein are responsible for the green-flesh and chlorophyll retainer mutations of tomato and pepper" 147 : 179-187, 2008

56 Delmas, F., "ABI3 controls embryo degreening through Mendel's I locus" 110 : 3888-3894, 2013

57 Gray, J., "A novel suppressor of cell death in plants encoded by the Lls1 gene of maize" 89 : 25-31, 1997

58 Mecey, C., "A critical role of STAYGREEN/Mendel's I locus in controlling disease symptom development during Pseudomonas syringae pv tomato infection of Arabidopsis" 157 : 1965-1974, 2011

59 Luo, Z., "A STAY-GREEN protein SlSGR1 regulates lycopene and β-carotene accumulation by interacting directly with SlPSY1 during ripening processes in tomato" 198 : 442-452, 2013

60 Sakuraba, Y., "7-Hydroxymethyl chlorophyll a reductase functions in metabolic channeling of chlorophyll breakdown intermediates during leaf senescence" 430 : 32-37, 2013