水稻温敏感叶色突变体研究进展

doi:10．3969/j．issn．1001G7216．2015．04．014

摘要/Abstract

摘要：

温度是影响水稻生长发育的一个重要因素,由温度引起水稻叶色改变的一类突变体统称为水稻温敏感叶色突变体。探讨温度影响水稻叶色的机理,对促进水稻遗传改良与高产育种具有重要意义。对已发掘的水稻温敏感突变体在温度对突变体的表型影响、基因定位及克隆、分子作用机理和育种利用及其研究进展等方面进行了分类讨论。

关键词: 水稻, 温敏感突变体, 叶色, 基因

Abstract:

Temperature is one of the factors affecting the growth and development of rice. The rice mutant whose leaf-color varied with temperature is refered to thermo-sensitive leaf coloration mutant. Understanding the mechanism of thermo-sensitive leaf coloration mutation is significant for promoting genetic improvement and high-yielding breeding in rice. We reviewed the recent research progresses about phenotypes, genetic mapping, cloning, molecular mechanism, breeding and utilization of thermo-sensitive leaf coloration mutants in rice.

Key words: rice, thermo-sensitive mutant, leaf color, gene

中图分类号:

Q343.5
Q754

刘艳霞, 林冬枝, 董彦君. 水稻温敏感叶色突变体研究进展[J]. 中国水稻科学, 2015, 29(4): 439-446.

Yan-xia LIU, Dong-zhi LIN, Yan-jun DONG. Research Advances in Thermo-sensitive Leaf Coloration Mutants in Rice[J]. Chinese Journal OF Rice Science, 2015, 29(4): 439-446.

图/表 1

参考文献 70

[1]	Dong F G, Zhu X D, Xiong Z D, et al.Breeding of a photo-thermo-period sensitive genie male sterile indica rice with a pale-green-leaf marker.Chin J Rice Sci, 1995, 9(2): 65-70.
[2]	Larkin R M, Alonso J M, Ecker J R, et al.GUN4, a regulator of chlorophyll synthesis and intracellular signaling.Science, 2003, 299: 902-906.
[3]	李育红, 王宝和, 戴正元, 等. 水稻叶色突变体及其基因定位、克隆的研究进展. 江苏农业科学, 2011, 39(2): 34-39.
[4]	Gustafsson A.The plastid development in various types of chlorophyll mutations.Hereditas, 1942, 28(3/4): 483-492.
[5]	Awan M A, Konzak C F, Rutger J N, et al.Mutagenic effect of sodium azide in rice.Crop Sci, 1980, 20(5): 663-668.
[6]	张力科, 高用明. 水稻叶色突变体及其基因定位和克隆的研究进展. 作物杂志, 2009, 2: 12-16.
[7]	Falbel T G,Staehelin L A.Characterization of a family of chlorophyll-deficient wheat(Triticum) and barly(Hordeum vulgare) mutants with defects in the magnesium-insertion step of chlorophyll biosynthesis.Plant Physiol, 1994, 104: 639-648.
[8]	吴殿星, 舒庆尧, 夏英武. 60Coγ射线诱发的籼型温敏核不育水稻叶色突变系变异分析. 作物学报, 1999, 25(3): 64-69.
[9]	Kusumi K, Komori H, Satoh H, et al.Characterization of a zebra mutant of rice with increased susceptibility to light stress.Plant Cell Physiol, 2000, 41(2): 158-164.
[10]	夏九成. 水稻白化突变体和云南软米的超微结构观察及相关基因的遗传分析和分子标记定位[D]. 成都: 四川农业大学, 2006.
[11]	朱明库, 胡宗利, 周爽, 等. 植物叶色白化研究进展. 生命科学, 2012, 24(3): 255-261.
[12]	杨麟, 罗大刚. 水稻叶色突变体的研究进展. 安徽农业科学, 2013, 41(18): 3341-3343, 3378.
[13]	Jiang Q, Mei J, Gong X D, et al.Importance of the rice TCD9 encoding α subunit of chaperonin protein60 (Cpn60α) for the chloroplast development during the early leaf stage.Plant Sci, 2014(215/216): 172-179.
[14]	Alberte R S, Hesketh J D, Hofstra G, et al.Composition and activity of the photosynthetic apparatus in temperature-sensitive mutants of higher plants.Proc Natl Acad Sci USA, 1974, 71(6): 2414-2418.
[15]	Markwell J P, Danko S J, Bauwe H, et al.A temperature-sensitive chlorophyll b-deficient mutant of sweetclover (Melilotus alba).Plant Physiol, 1986, 81(2): 329-334.
[16]	Markwell J P, Osterman J C.Occurrence of temperature-sensitive phenotypic plasticity in chlorophyll-deficient mutants of Arabidopsis thaliana.Plant Physiol, 1992, 98(1): 392-394.
[17]	Galova E, Bohmova B, Sevcovicova A.Analysis of some barley chlorophyll mutants and their response to temperature stress.Photosynthetica, 2000, 38(1): 29-35.
[18]	Pasini L, Bruschini S, Bertoli A, et al.Photosynthetic performance of cold-sensitive mutants of maize at low temperature.Physiol Plant, 2005, 124(3): 362-370.
[19]	Kusumi K, Mizutani A, Nishimura M, et al.A virescent gene V1 determines the expression timing of plastid genes for transcription/translation apparatus during early leaf development in rice.Plant J, 1997, 12(6): 1241-1250.
[20]	Sugimoto H, Kusumi K, Tozawa Y, et al.The virescent-2 mutation inhibits translation of plastid transcripts for the plastid genetic system at an early stage of chloroplast differentiation.Plant Cell Physiol, 2004, 45(8): 985-996.
[21]	王军, 杨杰, 陈志德, 等. 水稻白化转绿突变体v13(t)的生理特性和基因定位. 中国农业科学, 2011, 44(10): 1973-1979.
[22]	崔海瑞, 夏英武, 高明尉, 等. 温度对水稻突变体W1叶色及叶绿素生物合成的影响. 核农学报, 2001, 15(5): 269-273.
[23]	舒庆尧, 刘贵付, 夏英武. 温敏水稻叶色突变体的研究. 核农学报, 1996, 10(1): 6-10.
[24]	董彦君, 董文其, 张小明, 等. 突变体Fan5苗色低温敏感性状的遗传分析. 中国水稻科学, 1995, 9(4): 249-250.
[25]	陈佳颖, 赵剑, 刘晓, 等. 一个新水稻温敏感叶色突变休的遗传分析及其基因分子定位. 植物学报, 2010, 45(4): 419-425.
[26]	Gong X D, Su Q Q, Lin D Z, et al.The rice OsV4 encoding a novel pentatricopeptide repeat protein is required for chloroplast development during the early leaf stage under cold stress.J Intege Plant Biol, 2014, 56(4): 400-410.
[27]	Song J, Wei X J, Shao G N, et al.The rice nuclear gene WLP1 encoding a chloroplast ribosome L13 protein is needed for chloroplast development in rice grown under low temperature conditions.Plant Mol Biol, 2014, 84: 301-314.
[28]	魏祥进, 宋健, 胡培松, 等. 水稻苗期低温白叶突变体cde2的鉴定和基因定位. 中国水稻科学, 2014, 28(2): 111-118.
[29]	郭涛, 黄宜, 黄永相, 等. 一个控制水稻叶色白化转绿及多分蘖矮杆性状基因hw-1(t)的鉴定. 作物学报, 2012, 38(1): 23-35.
[30]	Liu W, Fu Y, Hu G, et al.Identification and fine mapping of a thermo-sensitive chlorophyll deficient mutant in rice(Oryza sativa L. ).Planta, 2007, 226(3): 785-795.
[31]	He Y H, Zou G X, Rao T C, et al.Genetic analysis and gene mapping of a rice white stripe leaf mutant(st10).Plant Gene Trait, 2011, 2(4): 23-29.
[32]	吴军, 陈佳颖, 赵剑, 等. 2个水稻温敏感叶色突变体的光合特性研究. 中国农学通报, 2012, 28(21): 16-21.
[33]	郑加兴, 覃保祥, 邱永福, 等. 水稻低温白化转绿突变系ds93的形态生理特性及基因定位. 西南农业学报, 2013, 26(3): 843-849.
[34]	Yoo S C, Cho S H, Sugimoto H, et al.Rice virescent3 andStripe1encoding the large and small subunits of ribonucleotide reductase are required for chloroplast biogenesis during early leaf development.Plant Physiol, 2009, 150(1): 388-401.
[35]	赵剑. 一个受温度调节水稻叶绿素合成PPR基因TCD1的图位克隆及初步功能分析. 上海: 上海师范大学, 2011.
[36]	Dong Y J, Dong W Q, Shi S Y, et al.Identification and genetic analysis of a thermo-sensitive seedling-colour mutant in rice(Oryza sativa L. ).Breeding Sci, 2001, 51: 1-4.
[37]	Xia J, Wang Y, Ma B, et al.Ultrastructure and gene mapping of the albino mutant al12 in rice (Oryza sativa L. ).Acta Genet Sin, 2006, 33(12): 1112-1119.
[38]	Kusumi K, Sakata C, Nakamura T, et al.A plastid protein NUS1 is essential for build-up of the genetic system for early chloroplast development under cold stress conditions.Plant J, 2011, 68(6): 1039-1050.
[39]	郭鹏, 邵健丰, 刘洪家, 等. 水稻温敏黄转绿突变体v5的鉴定和基因定位. 浙江农业学报, 2011, 23(5): 857-861.
[40]	兰涛, 汪斌, 凌秋平, 等. 水稻苗期低温失绿基因cisc(t)的精细定位及其候选基因的确定. 科学通报, 2010, 55(22): 2183-2187.
[41]	祁鲁, 刘晓, 陈佳颖, 等. 一个新的水稻温敏感叶色突变体基因定位分析. 中国水稻科学, 2010, 24(3): 226-227.
[42]	董彦君, 林冬枝, 梅杰, 等. 一个水稻温度敏感失绿突变体的遗传分析及分子定位. 分子植物育种, 2013, 11(1): 1-7.
[43]	江少华, 周华, 林冬枝, 等. 水稻幼苗叶色温敏感突变体的鉴定及其基因定位. 中国水稻科学, 2013, 27(4): 359-364.
[44]	李超, 林冬枝, 董彦君, 等. 一个水稻苗期温敏感白色条斑叶突变体的遗传分析及基因定位. 中国水稻科学, 2010, 24(3): 223-227.
[45]	Peng Y, Zhang Y, Lv J, et al.Characterization and fine mapping of a novel rice albino mutant low temperature albino1.J Genet Genom, 2012, 39: 385-396.
[46]	何瑞锋, 丁毅, 余金洪, 等. 水稻温敏叶绿素突变体叶片超微结构的研究. 武汉植物学研究, 2001, 19(1): 1-5.
[47]	钱前, 程式华. 水稻遗传学和功能基因组学. 北京: 科学出版社, 2006: 35-43.
[48]	Funke L, Dakoji S, Bredt D S.Membrane-associated guanylate kinases regulate adhesion and plasticity at cell junctions.Annu Rev Biochem, 2005, 74: 219-245.
[49]	Kuhlendahl S, Spangenberg O, Konrad M, et al.Functional analysis of the guanylate kinase-like domain in the synapse-associated protein SAP97.Eur J Biochem, 1998, 252: 305-313.
[50]	Olsen O, Bredt D S.Functional analysis of the nucleotide binding domain of membrane-associated guanylate kinases.J Biol Chem, 2003, 278: 6873-6878.
[51]	Elpeleg O, Mandel H, Saada A.Depletion of the other genome-mitochondrial DNA depletion syndromes in humans.J Mol Med, 2002, 80: 389-396.
[52]	Hedtke B, Wagner I, Börner T, et al.Inter-organellar crosstalk in higher plants: Impaired chloroplast development affects mitochondrial gene and transcript levels.Plant J, 1999, 19: 635-643.
[53]	Sugimoto H, Kusumi K, Noguchi K, et al.The rice nuclear gene, VIRESCENT2, is essential for chloroplast development and encodes a novel type of guanylate kinase targeted to plastids and mitochondria.Plant J, 2007, 52(3): 512-527.
[54]	Suzuki K, Nakanishi H, Bower J, et al.Plastid chaperonin proteins Cpn60α and Cpn60β are required for plastid division inArabidopsis thaliana. BMC Plant Biol, 2009, 9: 38.
[55]	Vitha S, McAndrew R S, Osteryoung K W. FtsZ ring formation at the chloroplast division site in plants.J Cell Biol, 2001, 153: 111-120.
[56]	McAndrew R S, Froehlich J E, Vitha S, et al. Colocalization of plastid division proteins in the chloroplast stromal compartment establishes a new functional relationship between FtsZ1 and FtsZ2 in higher plants.Plant Physiol, 2001, 127: 1656-1666.
[57]	Koussevitzky S, Stanne T M, Peto C A, et al.An Arabidopsis thaliana virescent mutant reveals a role for ClpR1 in plastid development.Plant Mol Biol, 2007, 63: 85-96.
[58]	Mandel M A, Feldmann K A, Herrera-Estrella L, et al.CLA1, a novel gene required for chloroplast development, is highly conserved in evolution.Plant J, 1996, 9: 649-658.
[59]	Hajdukiewicz P T, Allison L A, Maliga P.The two RNA polymerases encoded by the nuclear and the plastid compartments transcribe distinct groups of genes in tobacco plastids.EMBO J, 1997, 16: 4041-4048.
[60]	Maliga P.Two plastid RNA polymerases of higher plants: An evolving story.Trends Plant Sci, 1998, 3: 4-6.
[61]	Chandra Sanyal S, Liljas A.The end of the beginning: Struc-tural studies of ribosomal proteins.Curr Opin Struct Biol, 2000, 10: 633-636.
[62]	Maguire B A, Zimmermann R A.The ribosome in focus.Cell, 2001, 104: 813-816.
[63]	Ramakrishnan V, Moore P B.Atomic structures at last: The ribosome in 2000.Curr Opin Struct Biol, 2001, 11: 144-154.
[64]	Sharma M R, Wilson D N, Datta P P, et al.Cryo-EM study of the spinach chloroplast ribosome reveals the structural and functional roles of plastid-specific ribosomal proteins.Proc Natl Acad Sci USA, 2007, 104: 19315-19320.
[65]	Gerdes S Y, Scholle M D, Campbell J W, et al.Experimental determination and system level analysis of essential genes in Escherichia coli MG1655.J Bacteriol, 2003, 185: 5673-5684.
[66]	Eckhardt U, Grimm B, Hörtensteiner S.Recent advances in chlorophyll biosynthesis and breakdown in higher plants.Plant Mol Biol, 2004, 56: 1-14.
[67]	Lee S W, Cho B H, Park S G, et al.Aminoacyl-tRNA synthetase complexes: Beyond translation.J Cell Sci, 2004, 117: 3725-3734.
[68]	Park S G, Ewalt K L, Kim S.Functional expansion of aminoacyl-tRNA synthetases and their interacting factors: New perspectives on housekeepers.Trends Biochem Sci, 2005, 30: 569-574.
[69]	Qiu W, Zhou B, Darwish D, et al.Characterization of enzymatic properties of human ribonucleotide reductase holoenzyme reconstituted in vitro from hRRM1, hRRM2, and p53R2 subunits.Biochem Biophys Res Commun, 2006, 340: 428-434.
[70]	Chabouté M E, Combettes B, Clément B, et al.Molecular characterization of tobacco ribonucleotide reductase RNR1 and RNR2 cDNAs and cell cycle-regulated expression in synchronized plant cells.Plant Mol Biol, 1998, 38: 797-806.

突变体名称 Mutant name	叶色表型(温度) Leaf phenotypes (temperature/℃)	临界温度 Critical temperature /℃	染色体位置 Chromosomal location	克隆基因 Cloned gene	参考文献 Reference
低温敏感型 Low-temperature sensitive type
cde2	白色(23), 绿色(30) White(23),Green(30)	27	1	-	[28]
mr21	黄白色(20), 绿色(30) Yellow-white (20),Green(30)	27.5	1	-	[25]
Fan5	白色(20), 绿色(28) White(20), Green(28)	26.1	-	-	[24]
7436S	白色(23.1), 绿色(30.1) White(23.1), Green(30.1)	28	11	-	[36]
al12	白色(24), 绿色(26) White(24), Green(26)	-	8	-	[37]
ds93	白色(20), 绿色(23) White(20), Green(23)	20	9	-	[33]
v1	黄白色(20), 绿色(30) Yellow-white(20), Green(30)	-	3	V1	[19,38]
v2	黄白色(20), 绿色(30) White(20), Green(30)	-	3	V2	[20]
v5^*	黄色(24), 绿色(26) Yellow(24), Green(26)	26	9	-	[39]
v13	白色 (28), 绿色(30) White(28), Green(30)	28	5	-	[21]
cisc(t)	白色(25), 绿色(30) White(25), Green(30)	-	9	-	[40]
W1	白色(20), 绿色(30) White(20), Green(30)	23	-	-	[22]
W17	白色 (25), 浅绿色(30) White(25), Green(30)	-	-	-	[23]
W25	白色(26), 浅绿色(30 ) White(26), Green(30)	-	-	-	[23]
mr20	黄白色 (20), 绿色(32) Yellow white (20), Green(32)	22.5	3	-	[41]
tcd9	白色(24), 绿色(28) White(24), Green(28)	28	9	TCD9	[13]
tcm12	黄白色转绿(20), 绿色(24) Green-revertible albino (20), Green(24)	24	12	-	[42]
tsl1	黄色(20), 绿色(32) Yellow(20), Green(32)	-	11	-	[43]
tws	白色条斑(20), 绿色(32) White stripe(20), Green(32)	28	4	-	[44]
osv4	白化转绿(20), 绿色(32) Green-revertible albino (20), Green(32)	-	4	OsV4	[26]
lta1	白化转绿 (20), 绿色(24) Green-revertible albino (20), Green(24)	-	11	-	[45]
1103S	间断失绿(23.1), 绿色(26) Green-yellow rand(23.1), Green(26)	23.1	-	-	[46]
wlp1	白色(23), 绿色(30) White(23), Green(30)	-	1	WLP1	[27]
v4	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	11	-	[47]
v5	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	3	-	[47]
v6	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	1	-	[47]
v7	黄白色 (20), 绿色(30) Yellow white (20),Green(30)	-	3	-	[47]
v8	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	8	-	[47]
v9	黄白色 (20), 绿色(30) Yellow white (20), Green(30)	-	11	-	[47]
v10	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	5	-	[47]
v11	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	7	-	[47]
v12	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	-	-	[47]
chs1	白色(17), 绿色(30) White(17), Green(30)	-	9	-	[47]
chs2	白色(17), 绿色(30) White(17), Green(30)	-	-	-	[47]
chs3	白色(17), 绿色(30) White(17), Green(30)	-	-	-	[47]
chs4	白色(17), 绿色(30) White(17), Green(30)	-	-	-	[47]
高温敏感型 High temperature sensitive type
cde1(t)	绿色(23), 黄绿色(26) Green(23), Yellow green(26)	28	2	Cde1	[30]
mr06	绿色(20), 白色(30) Green(20), White(30)	-	2	TCD1	[32,35]
W4	黄色(25), 白色(30) Yellow(25), White(30)	-	-	-	[23]
W11	黄色(25), 白色(30), Yellow(25), White(30)	-	-	-	[23]
st10	绿色(28), 白色条纹(32) Green(28), White stripe(32)	-	3	-	[31]
hfa-1	黄白色 (15~20), 白色(25~30) Yellow white(15-20), White(25-30)	-	4	-	[29]
特殊温敏感型 Special temperature sensitive type
v3	白化 (20或30), 绿色(30/20) Bleached (20 or 30), Green(30/20)	-	6	V3	[34]
st1	白化 (20或30), 绿色(30/20) Bleached (20 or 30), Green(30/20)	-	6	ST1	[34]

突变体名称 Mutant name	叶色表型(温度) Leaf phenotypes (temperature/℃)	临界温度 Critical temperature /℃	染色体位置 Chromosomal location	克隆基因 Cloned gene	参考文献 Reference
低温敏感型 Low-temperature sensitive type
cde2	白色(23), 绿色(30) White(23),Green(30)	27	1	-	[28]
mr21	黄白色(20), 绿色(30) Yellow-white (20),Green(30)	27.5	1	-	[25]
Fan5	白色(20), 绿色(28) White(20), Green(28)	26.1	-	-	[24]
7436S	白色(23.1), 绿色(30.1) White(23.1), Green(30.1)	28	11	-	[36]
al12	白色(24), 绿色(26) White(24), Green(26)	-	8	-	[37]
ds93	白色(20), 绿色(23) White(20), Green(23)	20	9	-	[33]
v1	黄白色(20), 绿色(30) Yellow-white(20), Green(30)	-	3	V1	[19,38]
v2	黄白色(20), 绿色(30) White(20), Green(30)	-	3	V2	[20]
v5^*	黄色(24), 绿色(26) Yellow(24), Green(26)	26	9	-	[39]
v13	白色 (28), 绿色(30) White(28), Green(30)	28	5	-	[21]
cisc(t)	白色(25), 绿色(30) White(25), Green(30)	-	9	-	[40]
W1	白色(20), 绿色(30) White(20), Green(30)	23	-	-	[22]
W17	白色 (25), 浅绿色(30) White(25), Green(30)	-	-	-	[23]
W25	白色(26), 浅绿色(30 ) White(26), Green(30)	-	-	-	[23]
mr20	黄白色 (20), 绿色(32) Yellow white (20), Green(32)	22.5	3	-	[41]
tcd9	白色(24), 绿色(28) White(24), Green(28)	28	9	TCD9	[13]
tcm12	黄白色转绿(20), 绿色(24) Green-revertible albino (20), Green(24)	24	12	-	[42]
tsl1	黄色(20), 绿色(32) Yellow(20), Green(32)	-	11	-	[43]
tws	白色条斑(20), 绿色(32) White stripe(20), Green(32)	28	4	-	[44]
osv4	白化转绿(20), 绿色(32) Green-revertible albino (20), Green(32)	-	4	OsV4	[26]
lta1	白化转绿 (20), 绿色(24) Green-revertible albino (20), Green(24)	-	11	-	[45]
1103S	间断失绿(23.1), 绿色(26) Green-yellow rand(23.1), Green(26)	23.1	-	-	[46]
wlp1	白色(23), 绿色(30) White(23), Green(30)	-	1	WLP1	[27]
v4	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	11	-	[47]
v5	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	3	-	[47]
v6	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	1	-	[47]
v7	黄白色 (20), 绿色(30) Yellow white (20),Green(30)	-	3	-	[47]
v8	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	8	-	[47]
v9	黄白色 (20), 绿色(30) Yellow white (20), Green(30)	-	11	-	[47]
v10	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	5	-	[47]
v11	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	7	-	[47]
v12	黄白色(20), 绿色(30) Yellow white (20), Green(30)	-	-	-	[47]
chs1	白色(17), 绿色(30) White(17), Green(30)	-	9	-	[47]
chs2	白色(17), 绿色(30) White(17), Green(30)	-	-	-	[47]
chs3	白色(17), 绿色(30) White(17), Green(30)	-	-	-	[47]
chs4	白色(17), 绿色(30) White(17), Green(30)	-	-	-	[47]
高温敏感型 High temperature sensitive type
cde1(t)	绿色(23), 黄绿色(26) Green(23), Yellow green(26)	28	2	Cde1	[30]
mr06	绿色(20), 白色(30) Green(20), White(30)	-	2	TCD1	[32,35]
W4	黄色(25), 白色(30) Yellow(25), White(30)	-	-	-	[23]
W11	黄色(25), 白色(30), Yellow(25), White(30)	-	-	-	[23]
st10	绿色(28), 白色条纹(32) Green(28), White stripe(32)	-	3	-	[31]
hfa-1	黄白色 (15~20), 白色(25~30) Yellow white(15-20), White(25-30)	-	4	-	[29]
特殊温敏感型 Special temperature sensitive type
v3	白化 (20或30), 绿色(30/20) Bleached (20 or 30), Green(30/20)	-	6	V3	[34]
st1	白化 (20或30), 绿色(30/20) Bleached (20 or 30), Green(30/20)	-	6	ST1	[34]