水稻CSSL-Z492单、双片段代换系构建及粒型QTL的遗传解析

doi:10.3864/j.issn.0578-1752.2025.03.001

摘要/Abstract

摘要：

【目的】水稻粒型是由多基因控制的数量性状，将其遗传分解于单片段代换系（SSSL）中，并解析其遗传方式，为后续基因的遗传机制研究和设计育种提供依据。【方法】以日本晴为遗传背景的水稻染色体片段代换系Z492为研究材料，应用混合线性模型（MLM）进行粒型性状QTL的遗传分解和解析。【结果】以日本晴/Z492构建F₂群体，共鉴定出4个粒型QTL，包括粒长qGL6、qGL7和长宽比qRLW7、qRLW12，构建这些QTL的3个单片段代换系（S1—S3）和3个双片段代换系（D1—D3）。利用3个SSSL进一步鉴定出8个粒型性状QTL，包括qGL6、qGL7，以及6个新鉴定的QTL（qGW6、qRLW6、qGW7、qGWT7、qGL12和qGW12）。同时，分析不同QTL在3个DSSL中的遗传模式。结果表明，qGL6（a=0.26 mm）和qGL7（a=0.21 mm）互作产生-0.21 mm的粒长上位性效应，导致D1的粒长遗传效应（0.26 mm）与二者的加性效应相当。因而，D1的粒长（7.98 mm）与含单个QTL的S2和S1粒长（7.89和7.98 mm）无显著差异，而比日本晴的粒长（7.47 mm）显著增加。表明在设计育种中选择qGL6和qGL7聚合对增加粒长无效。qGW6（a=0.07 mm）和qGW12（a=0.06 mm）在D2中独立遗传，二者聚合产生的遗传效应（0.13 mm）使D2的粒宽（3.65 mm）比相应单片段代换系显著增加。因而在设计育种中可选这两个QTL增加粒宽。qGW7（a=0.11 mm）和qGW12（a=0.06 mm）互作则产生-0.10 mm的粒宽上位性效应，导致D3的粒宽遗传效应（0.07 mm）与qGW12的加性效应相当。因而D3的粒宽（3.59 mm）与含qGW12的S3无显著差异，而比日本晴显著变宽（3.44 mm），比携带qGW7的S2显著变窄（3.66 mm）。【结论】以单片段代换系和双片段代换系鉴定不同性状QTL对设计育种是非常必要的。不同QTL聚合会产生不同的遗传模式，有些是独立遗传，有些呈现不同的上位性效应。同时，S1和S3杂交可实现长宽大粒的育种目标，S1和S2杂交可产生比相应单片段代换系更重的籽粒，而S2和S3杂交无实际意义。

关键词: 水稻, 粒型, QTL, 单片段代换系, QTL互作

Abstract:

【Objective】Rice grain size is a quantitative trait controlled by multiple genes. They can be dissected into a single segment substitution line (SSSL), which is of great significance for their genetic mechanism study and breeding by design. 【Method】Z492, a chromosome segment substitution line in the genetic background of Nipponbare, was used as material to dissect QTL for rice grain size by mixed linear model (MLM) method. 【Result】The F₂ population was constructed from Nipponbare/Z492 to identify four QTL for grain size, including qGL6 and qGL7 for grain length and qRLW7 and qRLW12 for rate of grain length to width. Then three single-segment substitution lines (SSSL, S1-S3) and 3 dual-segment substitution lines (DSSL, D1-D3) carrying these QTL were further constructed. And the SSSL were then used to detect eight QTL for grain size, including qGL6, qGL7 and six newly identified QTL (qGW6, qRLW6, qGW7, qGWT7, qGL12, qGW12). Simultaneously, the genetic model of different QTL in 3 DSSL were analyzed. The results showed that interaction of qGL6 (a=0.26 mm) and qGL7 (a=0.21 mm) produced -0.21 mm of grain length epistatic effect, which resulted in the genetic effect (0.26 mm) of D1 equal to the additive effect of each QTL. Thus, the grain length (7.98 mm) of D1 displayed no difference from those (7.89 and 7.98 mm) of S2 with qGL7 and S1 containing qGL6, while significantly longer than that (7.47 mm) of Nipponbare. The result indicated that it is not necessary to pyramid qGL6 and qGL7 in breeding by design for increasing grain length. qGW6 (a=0.07 mm) and qGW12 (a=0.06 mm) belonged to independent inheritance in D2, thus, the genetic effect (0.13 mm) after pyramiding of qGW6 and qGW12 caused the grain width (3.65 mm) of D2 broader significantly than any of the SSSL with the single QTL. So, qGW6 and qGW12 can be selected to increase grain width in breeding by design. Interaction of qGW7 (a=0.11 mm) and qGW12 (a=0.06 mm) yielded -0.10 mm of epistatic effect, causing the grain width genetic effect (0.07 mm) of D3 parallel to the additive effect of qGW12. Thus, the grain width (3.59 mm) of D3 exhibited no difference with that (3.56 mm) of S3 carrying qGW12, while wider significantly than that (3.44 mm) of Nipponbare and narrower significantly than that (3.66 mm) of S2. 【Conclusion】It is very necessary for breeding by design to identify QTL for different important traits using SSSL and DSSL. Pyramiding different QTL produce various genetic models. Some display independent inheritance, and others exhibit various epistatic effects. In addition, to cross with S1 and S3 can realize the goal of longer, wider and heavier rice grain, and to cross with S1 and S2 can reach the target of heavier grain weight, while to cross with S2 and S3 have no any effects in grain size.

Key words: rice, grain size, QTL, single segment substitution lines, QTL interaction

李璐, 谢庄, 谢可盈, 张瀚, 赵卓文, 向奥妮, 李巧龙, 凌英华, 何光华, 赵芳明. 水稻CSSL-Z492单、双片段代换系构建及粒型QTL的遗传解析[J]. 中国农业科学, 2025, 58(3): 401-415.

LI Lu, XIE Zhuang, XIE KeYing, ZHANG Han, ZHAO ZhuoWen, XIANG AoNi, LI QiaoLong, LING YingHua, HE GuangHua, ZHAO FangMing. Construction of Single and Dual-Segment Substitution Lines from Rice CSSL-Z492 and Genetic Dissection of QTL for Grain Size[J]. Scientia Agricultura Sinica, 2025, 58(3): 401-415.

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参考文献 39

[1]	HORI K, SUN J. Rice grain size and quality. Rice, 2022, 15(1): 33. doi: 10.1186/s12284-022-00579-z pmid: 35776387
[2]	PONCE K, ZHANG Y, GUO L B, LENG Y J, YE G Y. Genome-wide association study of grain size traits in Indica rice multiparent advanced generation intercross (MAGIC) population. Frontiers in Plant Science, 2020, 11: 395.
[3]	XING Y Z, ZHANG Q F. Genetic and molecular bases of rice yield. Annual Review of Plant Biology, 2010, 61: 421-442. doi: 10.1146/annurev-arplant-042809-112209 pmid: 20192739
[4]	PARK J R, SEO J, PARK S, JIN M N, JEONG O Y, PARK H S. Identification of potential QTLs related to grain size in rice. Plants, 2023, 12(9): 1766.
[5]	LENG Y J, WANG S L, WANG R A, TAO T, JIA S W, SONG T, XU L N, CAI X L, JIN S K, GAO J P. Multi-environmental genetic analysis of grain size traits based on chromosome segment substitution line in rice (Oryza sativa L.). Phyton-International Journal of Experimental Botany, 2022, 91(5): 943-958.
[6]	徐建军, 梁国华. 水稻染色体片段代换系群体的构建及应用研究进展. 安徽农业科学, 2011, 39(4): 1935-1938.
	XU J J, LIANG G H. Research progress of construction and application of rice (Oryza sativa L.) chromosome segment substitution lines. Journal of Anhui Agricultural Sciences, 2011, 39(4): 1935-1938. (in Chinese)
[7]	BALAKRISHNAN D, SURAPANENI M, MESAPOGU S, NEELAMRAJU S. Development and use of chromosome segment substitution lines as a genetic resource for crop improvement. Theoretical and Applied Genetics, 2019, 132(1): 1-25. doi: 10.1007/s00122-018-3219-y pmid: 30483819
[8]	DING G M, HU B L, ZHOU Y, YANG W L, ZHAO M M, XIE J K, ZHANG F T. Development and characterization of chromosome segment substitution lines derived from Oryza rufipogon in the background of the Oryza sativa indica restorer line R974. Genes, 2022, 13(5): 735.
[9]	向思茜, 李儒香, 徐光益, 邓岢莉, 余金琎, 李苗苗, 杨正林, 凌英华, 桑贤春, 何光华, 赵芳明. 基于水稻长大粒染色体片段代换系Z66的粒型QTL的鉴定及其聚合分析. 作物学报, 2023, 49(3): 731-743. doi: 10.3724/SP.J.1006.2023.12081
	XIANG S Q, LI R X, XU G Y, DENG K L, YU J J, LI M M, YANG Z L, LING Y H, SANG X C, HE G H, ZHAO F M. Identification and pyramid analysis of QTLs for grain size based on rice long-large-grain chromosome segment substitution line Z66. Acta Agronomica Sinica, 2023, 49(3): 731-743. (in Chinese) doi: 10.3724/SP.J.1006.2023.12081
[10]	LI J, YANG H X, XU G Y, DENG K L, YU J J, XIANG S Q, ZHOU K, ZHANG Q L, LI R X, LI M M, LING, Y H, YANG Z L, HE G H, ZHAO F M. QTL analysis of Z414, a chromosome segment substitution line with short, wide grains, and substitution mapping of qGL11 in rice. Rice, 2022, 15(1): 25. doi: 10.1186/s12284-022-00571-7 pmid: 35532865
[11]	RIAZ A, WANG H M, ZHANG Z H, ZEGEYE W A, LI Y H, WANG H, XUE P, PENG Z Q, SHEN X H, CHENG S H, ZHANG Y X. Development of chromosome segment substitution lines and genetic dissection of grain size related locus in rice. Rice Science, 2021, 28(4): 322-324. doi: 10.1016/j.rsci.2021.05.003
[12]	LIANG P X, WANG H, ZHANG Q L, ZHOU K, LI M M, LI R X, XIANG S Q, ZHANG T, LING Y H, YANG Z L, HE G H, ZHAO F M. Identification and pyramiding of QTLs for rice grain size based on short-wide grain CSSL-Z563 and fine-mapping of qGL3-2. Rice, 2021, 14(1): 35. doi: 10.1186/s12284-021-00477-w pmid: 33847838
[13]	黄涛, 王燕宁, 钟奇, 程琴, 杨朦朦, 王鹏, 吴光亮, 黄诗颖, 李才敬, 余剑峰, 贺浩华, 边建民. 利用染色体片段置换系群体定位和分析水稻粒重和粒型QTL. 中国水稻科学, 2022, 36(2): 159-170. doi: 10.16819/j.1001-7216.2021.210204
	HUANG T, WANG Y N, ZHONG Q, CHENG Q, YANG M M, WANG P, WU G L, HUANG S Y, LI C J, YU J F, HE H H, BIAN J M. Mapping and analysis of QTLs for rice grain weight and grain shape using chromosome segment substitution line population. Chinese Journal of Rice Science, 2022, 36(2): 159-170. (in Chinese) doi: 10.16819/j.1001-7216.2021.210204
[14]	郑镇武, 赵宏源, 梁晓娅, 王一珺, 王驰航, 巩高洋, 黄金燕, 张桂权, 王少奎, 刘祖培. 水稻qGL3.4调控籽粒大小与株型. 遗传, 2023, 45(9): 835-844.
	ZHENG Z W, ZHAO H Y, LIANG X Y, WANG Y J, WANG C H, GONG G Y, HUANG J Y, ZHANG G Q, WANG S K, LIU Z P. qGL3.4 controls kernel size and plant architecture in rice. Hereditas, 2023, 45(9): 835-844. (in Chinese)
[15]	王军, 朱金燕, 周勇, 杨杰, 范方军, 李文奇, 梁国华, 仲维功. 基于染色体单片段代换系的水稻粒形QTL定位. 作物学报, 2013, 39(4): 617-625.
	WANG J, ZHU J Y ZHOU Y, YANG J, FAN F J, LI W Q, LIANG G H, ZHONG W G. Mapping of QTLs for grain shape using chromosome single segment substitution lines in rice (Oryza sativa L.). Acta Agronomica Sinica, 2013, 39(4): 617-625. (in Chinese)
[16]	展世杰. 水稻粒型QTL GL5的表型分析及精细定位[D]. 雅安: 四川农业大学, 2021.
	ZHAN S J. Phenotypic analysis and fine mapping of rice grain shape QTL GL5[D]. Yaan: Sichuan Agricultural University, 2021. (in Chinese)
[17]	张波, 裴瑞琴, 杨维丰, 朱海涛, 刘桂富, 张桂权, 王少奎. 利用单片段代换系鉴定巴西陆稻IAPAR9中的粒型基因. 作物学报, 2021, 47(8): 1472-1480. doi: 10.3724/SP.J.1006.2021.02056
	ZHANG B, PEI R Q, YANG W F, ZHU H T, LIU G F, ZHANG G Q, WANG S K. Mapping and identification QTLs controlling grain size in rice (Oryza sativa L.) by using single segment substitution lines derived from IAPAR9. Acta Agronomica Sinica, 2021, 47(8): 1472-1480. (in Chinese) doi: 10.3724/SP.J.1006.2021.02056
[18]	MA F Y, DU J, WANG D C, WANG H, ZHAO B B, HE G H, YANG Z L, ZHANG T, WU R H, ZHAO F M. Identification of long-grain chromosome segment substitution line Z744 and QTL analysis for agronomic traits in rice. Journal of Integrative Agriculture, 2020, 19(5): 1163-1169.
[19]	王大川, 汪会, 马福盈, 杜婕, 张佳宇, 徐光益, 何光华, 李云峰, 凌英华, 赵芳明. 增加穗粒数的水稻染色体代换系Z747鉴定及相关性状QTL定位. 作物学报, 2020, 46(1): 140-146.
	WANG D C, WANG H, MA F Y, DU J, ZHANG J Y, XU G Y, HE G H, LI Y F, LING Y H, ZHAO F M. Identification of rice chromosome segment substitution Line Z747 with increased grain number and QTL mapping for related traits. Acta Agronomica Sinica, 2020, 46(1): 140-146. (in Chinese)
[20]	LI J, ZHOU J W, ZHANG Y, YANG Y, PU Q H, TAO D Y. New insights into the nature of interspecific hybrid sterility in rice. Frontiers in Plant Science, 2020, 11: 555572.
[21]	CUI Y R, LI R D, LI G W, ZHANG F, ZHU T T, ZHANG Q F, ALI J, LI Z K, XU S Z. Hybrid breeding of rice via genomic selection. Plant Biotechnology Journal, 2020, 18(1): 57-67. doi: 10.1111/pbi.13170 pmid: 31124256
[22]	NADIR S, KHAN S, ZHU Q, HENRY D, WEI L, LEE D S, CHEN L J. An overview on reproductive isolation in Oryza sativa complex. AoB PLANTS, 2018, 10(6): ply060.
[23]	ZHANG G Q. Prospects of utilization of inter-subspecific heterosis between indica and Japonica rice. Journal of Integrative Agriculture, 2020, 19(1): 1-10.
[24]	FURUTA T, UEHARA K, ANGELES-SHIM R B, SHIM J, ASHIKARI M, TAKASHI T. Development and evaluation of chromosome segment substitution lines (CSSLs) carrying chromosome segments derived from Oryza rufipogon in the genetic background of Oryza sativa L.. Breeding Science, 2014, 63(5): 468-475.
[25]	李儒香, 周恺, 王大川, 李巧龙, 向奥妮, 李璐, 李苗苗, 向思茜, 凌英华, 何光华, 赵芳明. 水稻CSSL-Z481代换片段携带的穗部性状QTL分析及次级代换系培育. 中国农业科学, 2023, 56(7): 1228-1247. doi: 10.3864/j.issn.0578-1752.2023.07.003.
	LI R X, ZHOU K, WANG D C, LI Q L, XIANG A N, LI L, LI M M, XIANG S Q/X, LING Y H, HE G H, ZHAO F M. Analysis of QTLs and breeding of secondary substitution lines for panicle traits based on rice chromosome segment substitution line CSSL-Z481. Scientia Agricultura Sinica, 2023, 56(7): 1228-1247. doi: 10.3864/j.issn.0578-1752.2023.07.003. (in Chinese)
[26]	ZHANG T, WANG S M, SUN S F, ZHANG Y, LI J, YOU J, SU T, CHEN W B, LING Y H, HE G H, ZHAO F M. Analysis of QTL for grain size in a rice chromosome segment substitution line Z1392 with long grains and fine mapping of qGL-6. Rice, 2020, 13(1): 40.
[27]	WANG H, ZHANG J Y, FARKHANDA N, LI J, SUN S F, HE G H, ZHANG T, LING Y H, ZHAO F M. Identification of rice QTLs for important agronomic traits with long-kernel CSSL-Z741 and three SSSLs. Rice Science, 2020, 27(5): 414-422. doi: 10.1016/j.rsci.2020.04.008
[28]	沈文强, 赵冰冰, 于国玲, 李凤菲, 朱小燕, 马福盈, 李云峰, 何光华, 赵芳明. 优良水稻染色体片段代换系Z746的鉴定及重要农艺性状QTL定位及其验证. 作物学报, 2021, 47(3): 451-461. doi: 10.3724/SP.J.1006.2021.92002
	SHEN W Q, ZHAO B B, YU G L, LI F F, ZHU X Y, MA F Y, LI Y F, HE G H, ZHAO F M. Identification of an excellent rice chromosome segment substitution line Z746 and QTL mapping and verification of important agronomic traits. Acta Agronomica Sinica, 2021, 47(3): 451-461. (in Chinese)
[29]	李苗苗, 李儒香, 秦鱼河, 邓岢莉, 余金琎, 徐光益, 向思茜, 杨正林, 桑贤春, 凌英华, 何光华, 赵芳明. 基于水稻矮秆长粒CSSL-Z688的QTL鉴定及SSSLs培育. 西南大学学报(自然科学版), 2023, 45(1): 33-44.
	LI M M, LI R X, QIN Y H, DENG K L, YU J J, XU G Y, XIANG S Q/X, YANG Z L, SANG X C, LING Y H, HE G H, ZHAO F M. Identification of QTL based on a dwarf and long-large grain rice CSSL-Z688 and development of SSSLs. Journal of Southwest University (Natural Science Edition), 2023, 45(1): 33-44. (in Chinese)
[30]	赵志强, 傅亚萍, 杨鹍, 张玉满, 颜永胜, 方荣祥, 孙宗修, 陈晓英. 水稻小G蛋白OsPra2基因的克隆及功能分析. 生物工程学报, 2008, 24(12): 2027-2033.
	ZHAO Z Q, FU Y P, YANG K, ZHANG Y M, YAN Y S, FANG R X, SUN Z X, CHEN X Y. Cloning and function analysis of the rice small GTP-binding protein gene OsPra2. Chinese Journal of Biotechnology, 2008, 24(12): 2027-2033. (in Chinese)
[31]	ZHANG G, SONG X G, GUO H Y, WU Y, CHEN X Y, FANG R X. A small G protein as a novel component of the rice brassinosteroid signal transduction. Molecular Plant, 2016, 9(9): 1260-1271. doi: S1674-2052(16)30102-2 pmid: 27375203
[32]	ZHOU W, WANG X, ZHOU D, OUYANG Y D, YAO J L. Overexpression of the 16-kDa α-amylase/trypsin inhibitor RAG2 improves grain yield and quality of rice. Plant Biotechnology Journal, 2017, 15(5): 568-580. doi: 10.1111/pbi.12654 pmid: 27775871
[33]	YANG Y N, MA X S, XIA H, WANG L, CHEN S J, XU K, YANG F W, ZOU Y Q, WANG Y L, ZHU J M, LI T F, LUO Z, HU S P, LIAO Z G, LUO L J, YU S W. Natural variation of Alfin-like family affects seed size and drought tolerance in rice. The Plant Journal, 2022, 112(5): 1176-1193. doi: 10.1111/tpj.16003 pmid: 36219491
[34]	MA X S, FENG F J, ZHANG Y, ELESAWI I E, XU K, LI T F, MEI H W, LIU H Y, GAO N N, CHEN C L, LUO L J, YU S W. A novel rice grain size gene OsSNB was identified by genome-wide association study in natural population. PLoS Genetics, 2019, 15(5): e1008191.
[35]	XU X Y, ZHIGUO E, ZHANG D P, YUN Q B, ZHOU Y, NIU B X, CHEN C. OsYUC11-mediated auxin biosynthesis is essential for endosperm development of rice. Plant Physiology, 2021, 185(3): 934-950.
[36]	WANG D C, ZHOU K, XIANG S Q, ZHANG Q L, LI R X, LI M M, LIANG P X, FARKHANDA N, HE G H, LING Y H, ZHAO F M. Identification, pyramid and candidate genes of QTLs for associated traits based on a dense erect panicle rice CSSL-Z749 and five SSSLs, three DSSLs and one TSSL. Rice, 2021, 14(1): 55. doi: 10.1186/s12284-021-00496-7 pmid: 34132908
[37]	张龙廷, 吴静, 熊喜娟, 董景芳, 张少红, 赵均良, 刘自强, 杨梯丰. 基于单片段代换系的水稻苗高QTL定位和上位性效应分析. 华南农业大学学报, 2023, 44(6): 881-888.
	ZHANG L T, WU J, XIONG X J, DONG J F, ZHANG S H, ZHAO J L, LIU Z Q, YANG T F. QTL mapping and epistatic effect analysis of seedling height based on single segment substitution lines in rice. Journal of South China Agricultural University, 2023, 44(6): 881-888. (in Chinese)
[38]	ABIOLA O, ANGEL J M, AVNER P, BACHMANOV A A, BELKNAP J K, BENNETT B, BLANKENHORN E P, BLIZARD D A, BOLIVAR V, BROCKMANN G A, BUCK K J, BUREAU J F, CASLEY W L, CHESLER E J, CHEVERUD J M, CHURCHILL G A, COOK M, CRABBE J C, CRUSIO W E, DARVASI A, DE HAAN G, DERMANT P, DOERGE R W, ELLIOT R W, FARBER C R, FLAHERTY L, FLINT J, GERSHENFELD H, GIBSON J P, GU J, GU W K, HIMMELBAUER H, HITZEMANN R, HSU H C, HUNTER K, IRAQI F F, JANSEN R C, JOHNSON T E, JONES B C, KEMPERMANN G, LAMMERT F, LU L, MANLY K F, MATTHEWS D B, MEDRANO J F, MEHRABIAN M, MITTLEMANN G, MOCK B A, MOGIL J S, MONTAGUTELLI X, MORAHAN G, MOUNTZ J D, NAGASE H, NOWAKOWSKI R S, O’HARA B F, OSADCHUK A V, PAIGEN B, PALMER A A, PEIRCE J L, POMP D, ROSEMANN M, ROSEN G D, SCHALKWYK L C, SELTZER Z, SETTLE S, SHIMOMURA K, SHOU S M, SIKELA J M, SIRACUSA L D, SPEAROW J L, TEUSCHER C, THREADGILL D W, TOTH L A, TOYE A A, VADASZ C, VAN ZANT G, WAKELAND E, WILLIAMS R W, ZHANG H G, ZOU F, CONSORTIUM C T. The nature and identification of quantitative trait loci: A community' s view. Nature Reviews Genetics, 2003, 4(11): 911-916.
[39]	杨梯丰, 董景芳, 赵均良, 周炼, 杨武, 马雅美, 王健, 陈洛, 陈建松, 吴伟, 李雯慧, 刘斌, 张少红. 基于单片段代换系的水稻芽期耐冷QTL定位和聚合. 植物遗传资源学报, 2022, 23(6): 1726-1736. doi: 10.13430/j.cnki.jpgr.20220420002
	YANG T F, DONG J F, ZHAO J L, ZHOU L, YANG W, MA Y M, WANG J, CHEN L, CHEN J S, WU W, LI W H, LIU B, ZHANG S H. Identification and pyramiding of QTLs for cold tolerance at the bud bursting stage by use of single segment substitution lines in rice (Oryza sativa L.). Journal of Plant Genetic Resources, 2022, 23(6): 1726-1736. (in Chinese)

性状 Trait	QTL	染色体 Chr.	连锁标记 Linked marker	估计效应 Estimated effect	贡献率 Var. (%)	P值 P value
粒长Grain length (mm)	qGL6	6	RM494	0.07	4.97	0.0314
粒长Grain length (mm)	qGL7	7	RM5672	0.15	24.16	<0.0010
长宽比Rate of length to width	qRLW7	7	RM5672	0.03	6.24	0.0208
长宽比Rate of length to width	qRLW12	12	RM491	0.03	4.57	0.0400