中国东北地区水稻抽穗期性状研究进展
|
摘要
水稻抽穗期是指水稻从播种到抽穗(开花)所经历的时间,它是决定水稻产量和区域适应性的重要农艺性状之一。水稻抽穗期的数量性状很复杂,受内源多基因及外源多因素调控。本文介绍了水稻中2条保守的抽穗期调控途径,论述了各抽穗期基因间相互调控关系,并重点阐述了调控东北地区水稻抽穗期的主效基因和微效基因以及东北地区水稻中不同抽穗期基因的单体类型,同时解析了不同基因型组合形式对东北水稻区域分布的决定性作用,并探讨了通过基因编辑手段,对抽穗期抑制基因进行定点突变,缩短品种熟期,扩大优良品种种植面积,旨为从南方引进新水稻种质资源提供理论和实践指导。此外,针对目前东北地区水稻育种的现状,对分子设计育种与常规育种相结合的技术方式进行了展望。
Rice heading date( HD) refers to the time from sowing to heading( flowering). HD is one of the most important factors determining rice distribution and grain yield. HD,as a complex quantitative trait,is controlled by a number of genes and diverse environmental factors. In this review,we introduce two conserved flowering pathway,and the complex interaction among various HD genes in rice. We mainly describe the major and minor effect HD genes which control the rice distribution in Northeast China,and diverse haplotypes of each HD gene. We highlight that combination of different haplotype of multiple HD genes determine the rice distribution in Northeast China. In addition,we describe a CRISPR/cas9-mediated breeding technique by which the HD of cultivar can be efficiently shortened by mutating the heading date repressors genes,and adapted growth region of elite cultivar can move northward and planting area will be increased,simultaneously. This technique will facilitate to introduce new rice germplasms from South China to Northeast China. Based on the current situation of rice breeding and production in Northeast China,we propose to strengthen improving rice quality and increasing growth area by combining molecular design breeding and conventional breeding in future.
Rice heading date( HD) refers to the time from sowing to heading( flowering). HD is one of the most important factors determining rice distribution and grain yield. HD,as a complex quantitative trait,is controlled by a number of genes and diverse environmental factors. In this review,we introduce two conserved flowering pathway,and the complex interaction among various HD genes in rice. We mainly describe the major and minor effect HD genes which control the rice distribution in Northeast China,and diverse haplotypes of each HD gene. We highlight that combination of different haplotype of multiple HD genes determine the rice distribution in Northeast China. In addition,we describe a CRISPR/cas9-mediated breeding technique by which the HD of cultivar can be efficiently shortened by mutating the heading date repressors genes,and adapted growth region of elite cultivar can move northward and planting area will be increased,simultaneously. This technique will facilitate to introduce new rice germplasms from South China to Northeast China. Based on the current situation of rice breeding and production in Northeast China,we propose to strengthen improving rice quality and increasing growth area by combining molecular design breeding and conventional breeding in future.
引文
[1]BRAMBILLA V,GOMEZ-ARIZA J,CERISE M,et al.The importance of being on time:Regulatory networks controlling photoperiodic flowering in cereals[J].Frontiers in Plant Science,2017,8:665.
[2]AHN J H.Have florigen,will travel[J].Nature Plants,2016,2(6):16081.
[3]TAOKA K,OHKI I,TSUJI H,et al.Structure and function of florigen and the receptor complex[J].Trends in Plant Science,2013,18(5):287-294.
[4]EVANS L T.Flower induction and the florigen concept[J].Annual Review of Plant Physiology,1971,22:365-394.
[5]DU J J,LIU S P,LIU W H,et al.The application of photoperiodic control of the plant flowering[R].13th China International Forum on Solid State Lighting(Sslchina 2016),2016:101-103.
[6]HE Y H.Enabling photoperiodic control of flowering by timely chromatin silencing of the florigen gene[J].Nucleus,2015,6(3):179-182.
[7]MATSUBARA K,HORI K,OGISO-TANAKA E,et al.Cloning of quantitative trait genes from rice reveals conservation and divergence of photoperiod flowering pathways in Arabidopsis and rice[J].Frontiers in Plant Science,2014,5:193.
[8]舒黄英,郝园园,蔡庆泽,等.模式植物拟南芥开花时间分子调控研究进展[J].植物科学学报,2017,35(4):603-608.SHU H Y,HAO Y Y,CAI Q Z,et al.Recent research progress on the molecular regulation of flowering time in Arabidopsis thaliana[J].Plant Science Journal,2017,35(4):603-608.
[9]BLMEL M,DALLY N,JUNG C.Flowering time regulation in crops-what did we learn from Arabidopsis?[J].Current Opinion in Biotechnology,2015,32:121-129.
[10]SUN C H,CHEN D,FANG J,et al.Understanding the genetic and epigenetic architecture in complex network of rice flowering pathways[J].Protein and Cell,2014,5(12):889-898.
[11]MURAKAMI M,TAGO Y,YAMASHINO T,et al.Comparative overviews of clock-associated genes of Arabidopsis thaliana and Oryza sativa[J].Plant and Cell Physiology,2007,48(1):110-121.
[12]IZAWA T.Adaptation of flowering-time by natural and artificial selection in Arabidopsis and rice[J].Journal of Experimental Botany,2007,58(12):3091-3097.
[13]TAOKA K,OHKI I,TSUJI H,et al.14-3-3 proteins act as intracellular receptors for rice Hd3a florigen[J].Nature,2011,476:332-335.
[14]SONG S Y,CHEN Y,LIU L,et al.Os FTIP1-mediated regulation of florigen transport in rice is negatively regulated by the ubiquitin-like domain kinase Os Ub DKγ4[J].Plant Cell,2017,29(3):491-507.
[15]KOMIYA R,IKEGAMI A,TAMAKI S,et al.Hd3a and RFT1 are essential for flowering in rice[J].Development,2008,135(4):767-774.
[16]KOJIMA S,TAKAHASHI Y,KOBAYASHI Y,et al.Hd3a,a rice ortholog of the Arabidopsis FT gene,promotes transition to flowering downstream of Hd1 under short-day conditions[J].Plant and Cell Physiology,2002,43(10):1096-1105.
[17]SUN C H,FANG J,ZHAO T L,et al.The histone methyltransferase SDG724 mediates H3K36me2/3 deposition at MADS50 and RFT1 and promotes flowering in rice[J].The Plant Cell,2012,24(8):3235-3247.
[18]YANO M,KATAYOSE Y,ASHIKARI M,et al.Hd1,a major photoperiod sensitivity quantitative trait locus in rice,is closely related to the arabidopsis flowering time gene CONSTANS[J].Plant Cell,2000,12(12):2473-2483.
[19]G?MEZ-ARIZA J,GALBIATI F,GORETTI D,et al.Loss of floral repressor function adapts rice to higher latitudes in Europe[J].Journal of Experimental Botany,2015,66(7):2027-2039.
[20]GORETTI D,MARTIGNAGO D,LANDINI M,et al.Transcriptional and post-transcriptional mechanisms limit heading date 1(Hd1)function to adapt rice to high latitudes[J].PLo S Genetics,2017,13(1):e1006530.
[21]TAKAHASHI Y,SHIMAMOTO K.Heading date 1(Hd1),an ortholog of Arabidopsis CONSTANS,is a possible target of human selection during domestication to diversify flowering times of cultivated rice[J].Genes Genetic System,2011,86(3):175-182.
[22]YANG Y,FU D B,ZHU C M,et al.The RING-finger ubiquitin ligase HAF1 mediates heading date 1 degradation during photoperiodic flowering in rice[J].Plant Cell,2015,27(9):2455-2468.
[23]SUN X H,ZHANG Z G,WU J X,et al.The Oryza sativa regulator HDR1 associates with the kinase Os K4 to control photoperiodic flowering[J].PLo S Genetics,2016,12(3):e1005927.
[24]DOI K,IZAWA T,FUSE T,et al.Ehd1,a B-type response regulator in rice,confers short-day promotion of flowering and controls FT-like gene expression independently of Hd1[J].Genes and Development,2004,18(8):926-936.
[25]ZHAO J,CHEN H Y,REN D,et al.Genetic interactions between diverged alleles of Early heading date 1(Ehd1)and Heading date 3a(Hd3a)/RICE FLOWERING LOCUS T1(RFT1)control differential heading and contribute to regional adaptation in rice(Oryza sativa)[J].New Phytologist,2015,208(3):936-948.
[26]ENDO-HIGASHI N,IZAWA T.Flowering time Genes Heading date 1 and Early heading date 1 together control panicle development in rice[J].Plant and Cell Physiology,2011,52(6):1083-1094.
[27]XUE W Y,XING Y Z,WENG X Y,et al.Natural variation in Ghd7 is an important regulator of heading date and yield potential in rice[J].Nature genetics,2008,40(6):761-767.
[28]LI X F,LIU H Z,WANG M Q,et al.Combinations of Hd2 and Hd4 genes determine rice adaptability to Heilongjiang province,northern limit of China[J].Journal of Integrative Plant Biology,2015,57(8):698-707.
[29]WEI X J,XU J F,GUO H N,et al.DTH8 suppresses flowering in rice,influencing plant height and yield potential simultaneously[J].Plant Physiology,2010,153(4):1747-1758.
[30]YAN W H,WANG P,CHEN H X,et al.A major QTL,Ghd8,plays pleiotropic roles in regulating grain productivity,plant height,and heading date in rice[J].Molecular Plant,2011,4(2):319-330.
[31]GAO H,JIN M N,ZHENG X M,et al.Days to heading 7,a major quantitative locus determining photoperiod sensitivity and regional adaptation in rice[J].Proceedings of the National Academy of Sciences of the United States of America,2014,111(46):16337-16342.
[32]YAN W H,LIU H Y,ZHOU X C,et al.Natural variation in Ghd7.1 plays an important role in grain yield and adaptation in rice[J].Cell Research,2013,23(7):969-971.
[33]KOO B H,YOO S C,PARK J W,et al.Natural variation in Os PRR37 regulates heading date and contributes to rice cultivation at a wide range of latitudes[J].Molecular Plant,2013,6(6):1877-1888.
[34]HORI K,OGISO-TANAKA E,MATSUBARA K,et al.Hd16,a gene for casein kinase I,is involved in the control of rice flowering time by modulating the day-length response[J].The Plant Journal,2013,76(1):36-46.
[35]GAO H,ZHENG X M,FEI G L,et al.Ehd4 encodes a novel and Oryza-genus-specific regulator of photoperiodic flowering in rice[J].Plos Genetics,2013,9(2):e1003281.
[36]MATSUBARA K,YAMANOUCHI U,NONOUE Y,et al.Ehd3,encoding a plant homeodomain finger-containing protein,is a critical promoter of rice flowering[J].The Plant Journal,2011,66(4):603-612.
[37]WU C Y,YOU C L,LI C S,et al.RID1,encoding a Cys2/His2-type zinc finger transcription factor,acts as a master switch from vegetative to floral development in rice[J].Proceedings of the National Academy of Sciences of the United States of America,2008,105(35):12915-12920.
[38]SHIBAYA T,HORI K,OGISO-TANAKA E,et al.Hd18,encoding histone acetylase related to Arabidopsis FLOWERING LOCUS D,is involved in the control of flowering time in rice[J].Plant and Cell Physiology,2016,57(9):1828-1838.
[39]RYU C H,LEE S Y,CHO L H,et al.Os MADS50 and Os MADS56 function antagonistically in regulating long day(LD)-dependent flowering in rice[J].Plant,Cell and Environment,2009,32(10):1412-1427.
[40]ZHAO J M,HUANG X,OUYANG X H,et al.Os ELF3-1,an ortholog of Arabidopsis early flowering 3,regulates rice circadian rhythm and photoperiodic flowering[J].Plo S one,2012,7(8):e43705.
[41]MATSUBARA K,OGISO-TANAKA E,HORI K,et al.Natural variation in Hd17,a homolog of Arabidopsis ELF3 that is involved in rice photoperiodic flowering[J].Plant and Cell Physiology,2012,53(14):709-716.
[42]NEMOTO Y,NONOUE Y,YANO M,et al.Hd1,a CONSTANS ortholog in rice,functions as an Ehd1 repressor through interaction with monocot-specific CCT-domain protein Ghd7[J].The Plant Jounal,2016,86(3):221-233.
[43]WU W X,ZHENG X M,LU G W,et al.Association of functional nucleotide polymorphisms at DTH2 with the northward expansion of rice cultivation in Asia[J].Proceedings of the National Academy of Sciences of the United States of America,2013,110(8):2775-2780.
[44]KIM S K,YUN C H,LEE J H,et al.Os CO3,a CONSTANS-LIKE gene,controls flowering by negatively regulating the expression of FT-like genes under SD conditions in rice[J].Planta,2008,228(2):355-365.
[45]LI D J,YANG C H,LI X B,et al.Sense and antisense Os Dof12 transcripts in rice[J].BMC Molecular Biology,2008,9:80.
[46]LIU T M,LIU H Y,ZHANG H,et al.Validation and characterization of Ghd7.1,a major quantitative trait locus with pleiotropic effects on spikelets per panicle,plant height,and heading date in rice(Oryza sativa L.)[J].Journal of Integrative Plant Biology,2013,55(10):917-927.
[47]薛为亚.水稻产量相关基因Ghd7的分离与鉴定[D].武汉:华中农业大学,2008.XUE W Y.Isolation and characterization of a pleiotropic gene,Ghd7,in rice[D].Wuhan:Huazhong Agricultural University,2008.
[48]LU L,YAN W H,XUE W Y,et al.Evolution and association analysis of Ghd7 in rice[J].Plo S ONE,2012,7(5):e34021.
[49]FUJINO K,YAMANOUCHI U,YANO M.Roles of the Hd5 gene controlling heading date for adaptation to the northern limits of rice cultivation[J].Theoretical and Applied Genetics,2013,126(3):611-618.
[50]FUJINO K,SEKIGUCHI H.Identification of QTLs conferring genetic variation for heading date among rice varieties at the northern-limit of rice cultivation[J].Breeding Science,2005,55:141-146.
[51]LI X F,SUN Y Q,TIAN X J,et al.Comprehensive identification of major flowering time genes and their combinations,which determined rice distribution in Northeast China[J].Plant Growth Regulation,2018,DOI:10.1007/s10725-017-0364-2.
[52]于秋竹.黑龙江省不同积温带水稻产量和品质及株型研究[D].沈阳:沈阳农业大学,2014.YU Q Z.Research of rice yield quality and plant type in different accumulated temperature zone in Heilongjiang province[D].Shenyang:Shenyang Agricultural University,2014.
[53]姜树坤,张凤鸣,白良明,等.黑龙江省第一积温带水稻育种趋势分析[J].作物杂志,2015(4):61-64.JIANG S K,ZHANG F M,BAI L M,et al.The Japonica rice breeding trend in 1staccumulated temperature zone of Heilongjiang province[J].Crops,2015(4):61-64.
[54]李秀峰.黑龙江省水稻抽穗期基因的效应研究及育种应用[D].北京:中国科学院大学,2016.LI X F.The study on effect of heading date genes in Heilongjiang rice cultivars and their application in molecular assistant breeding[D].Beijing:University of Chinese Academy of Sciences,2016.
[55]周文甲,田晓杰,任月坤,等.利用CRISPR/Cas9创造早熟香味水稻[J].土壤与作物,2017,6(2):146-52.ZHOU W J,TIAN X J,REN Y K,et al.Breeding of early-maturating and fragrant rice via CRISPR/Cas9 mediated genome editing[J].Soils and Crops,2017,6(2):146-52.
[56]LI X F,ZHOU W J,REN Y K,et al.High-efficiency breeding of early-maturing rice cultivars via CRISPR/Cas9-mediated genome editing[J].Journal of Genetics and Genomics,2017,44(2):175-178.
[57]CHEN S H,YANG Y,SHI W W,et al.Badh2,encoding betaine aldehyde dehydrogenase,inhibits the biosynthesis of 2-acetyl-1-pyrroline,a major component in rice fragrance[J].The Plant cell,2008,20(7):1850-1861.
[58]TIAN Z X,QIAN Q,LIU Q Q,et al.Allelic diversities in rice starch biosynthesis lead to a diverse array of rice eating and cooking qualities[J].Proceedings of the National Academy of Sciences,2009,106(51):21760-21765.
[2]AHN J H.Have florigen,will travel[J].Nature Plants,2016,2(6):16081.
[3]TAOKA K,OHKI I,TSUJI H,et al.Structure and function of florigen and the receptor complex[J].Trends in Plant Science,2013,18(5):287-294.
[4]EVANS L T.Flower induction and the florigen concept[J].Annual Review of Plant Physiology,1971,22:365-394.
[5]DU J J,LIU S P,LIU W H,et al.The application of photoperiodic control of the plant flowering[R].13th China International Forum on Solid State Lighting(Sslchina 2016),2016:101-103.
[6]HE Y H.Enabling photoperiodic control of flowering by timely chromatin silencing of the florigen gene[J].Nucleus,2015,6(3):179-182.
[7]MATSUBARA K,HORI K,OGISO-TANAKA E,et al.Cloning of quantitative trait genes from rice reveals conservation and divergence of photoperiod flowering pathways in Arabidopsis and rice[J].Frontiers in Plant Science,2014,5:193.
[8]舒黄英,郝园园,蔡庆泽,等.模式植物拟南芥开花时间分子调控研究进展[J].植物科学学报,2017,35(4):603-608.SHU H Y,HAO Y Y,CAI Q Z,et al.Recent research progress on the molecular regulation of flowering time in Arabidopsis thaliana[J].Plant Science Journal,2017,35(4):603-608.
[9]BLMEL M,DALLY N,JUNG C.Flowering time regulation in crops-what did we learn from Arabidopsis?[J].Current Opinion in Biotechnology,2015,32:121-129.
[10]SUN C H,CHEN D,FANG J,et al.Understanding the genetic and epigenetic architecture in complex network of rice flowering pathways[J].Protein and Cell,2014,5(12):889-898.
[11]MURAKAMI M,TAGO Y,YAMASHINO T,et al.Comparative overviews of clock-associated genes of Arabidopsis thaliana and Oryza sativa[J].Plant and Cell Physiology,2007,48(1):110-121.
[12]IZAWA T.Adaptation of flowering-time by natural and artificial selection in Arabidopsis and rice[J].Journal of Experimental Botany,2007,58(12):3091-3097.
[13]TAOKA K,OHKI I,TSUJI H,et al.14-3-3 proteins act as intracellular receptors for rice Hd3a florigen[J].Nature,2011,476:332-335.
[14]SONG S Y,CHEN Y,LIU L,et al.Os FTIP1-mediated regulation of florigen transport in rice is negatively regulated by the ubiquitin-like domain kinase Os Ub DKγ4[J].Plant Cell,2017,29(3):491-507.
[15]KOMIYA R,IKEGAMI A,TAMAKI S,et al.Hd3a and RFT1 are essential for flowering in rice[J].Development,2008,135(4):767-774.
[16]KOJIMA S,TAKAHASHI Y,KOBAYASHI Y,et al.Hd3a,a rice ortholog of the Arabidopsis FT gene,promotes transition to flowering downstream of Hd1 under short-day conditions[J].Plant and Cell Physiology,2002,43(10):1096-1105.
[17]SUN C H,FANG J,ZHAO T L,et al.The histone methyltransferase SDG724 mediates H3K36me2/3 deposition at MADS50 and RFT1 and promotes flowering in rice[J].The Plant Cell,2012,24(8):3235-3247.
[18]YANO M,KATAYOSE Y,ASHIKARI M,et al.Hd1,a major photoperiod sensitivity quantitative trait locus in rice,is closely related to the arabidopsis flowering time gene CONSTANS[J].Plant Cell,2000,12(12):2473-2483.
[19]G?MEZ-ARIZA J,GALBIATI F,GORETTI D,et al.Loss of floral repressor function adapts rice to higher latitudes in Europe[J].Journal of Experimental Botany,2015,66(7):2027-2039.
[20]GORETTI D,MARTIGNAGO D,LANDINI M,et al.Transcriptional and post-transcriptional mechanisms limit heading date 1(Hd1)function to adapt rice to high latitudes[J].PLo S Genetics,2017,13(1):e1006530.
[21]TAKAHASHI Y,SHIMAMOTO K.Heading date 1(Hd1),an ortholog of Arabidopsis CONSTANS,is a possible target of human selection during domestication to diversify flowering times of cultivated rice[J].Genes Genetic System,2011,86(3):175-182.
[22]YANG Y,FU D B,ZHU C M,et al.The RING-finger ubiquitin ligase HAF1 mediates heading date 1 degradation during photoperiodic flowering in rice[J].Plant Cell,2015,27(9):2455-2468.
[23]SUN X H,ZHANG Z G,WU J X,et al.The Oryza sativa regulator HDR1 associates with the kinase Os K4 to control photoperiodic flowering[J].PLo S Genetics,2016,12(3):e1005927.
[24]DOI K,IZAWA T,FUSE T,et al.Ehd1,a B-type response regulator in rice,confers short-day promotion of flowering and controls FT-like gene expression independently of Hd1[J].Genes and Development,2004,18(8):926-936.
[25]ZHAO J,CHEN H Y,REN D,et al.Genetic interactions between diverged alleles of Early heading date 1(Ehd1)and Heading date 3a(Hd3a)/RICE FLOWERING LOCUS T1(RFT1)control differential heading and contribute to regional adaptation in rice(Oryza sativa)[J].New Phytologist,2015,208(3):936-948.
[26]ENDO-HIGASHI N,IZAWA T.Flowering time Genes Heading date 1 and Early heading date 1 together control panicle development in rice[J].Plant and Cell Physiology,2011,52(6):1083-1094.
[27]XUE W Y,XING Y Z,WENG X Y,et al.Natural variation in Ghd7 is an important regulator of heading date and yield potential in rice[J].Nature genetics,2008,40(6):761-767.
[28]LI X F,LIU H Z,WANG M Q,et al.Combinations of Hd2 and Hd4 genes determine rice adaptability to Heilongjiang province,northern limit of China[J].Journal of Integrative Plant Biology,2015,57(8):698-707.
[29]WEI X J,XU J F,GUO H N,et al.DTH8 suppresses flowering in rice,influencing plant height and yield potential simultaneously[J].Plant Physiology,2010,153(4):1747-1758.
[30]YAN W H,WANG P,CHEN H X,et al.A major QTL,Ghd8,plays pleiotropic roles in regulating grain productivity,plant height,and heading date in rice[J].Molecular Plant,2011,4(2):319-330.
[31]GAO H,JIN M N,ZHENG X M,et al.Days to heading 7,a major quantitative locus determining photoperiod sensitivity and regional adaptation in rice[J].Proceedings of the National Academy of Sciences of the United States of America,2014,111(46):16337-16342.
[32]YAN W H,LIU H Y,ZHOU X C,et al.Natural variation in Ghd7.1 plays an important role in grain yield and adaptation in rice[J].Cell Research,2013,23(7):969-971.
[33]KOO B H,YOO S C,PARK J W,et al.Natural variation in Os PRR37 regulates heading date and contributes to rice cultivation at a wide range of latitudes[J].Molecular Plant,2013,6(6):1877-1888.
[34]HORI K,OGISO-TANAKA E,MATSUBARA K,et al.Hd16,a gene for casein kinase I,is involved in the control of rice flowering time by modulating the day-length response[J].The Plant Journal,2013,76(1):36-46.
[35]GAO H,ZHENG X M,FEI G L,et al.Ehd4 encodes a novel and Oryza-genus-specific regulator of photoperiodic flowering in rice[J].Plos Genetics,2013,9(2):e1003281.
[36]MATSUBARA K,YAMANOUCHI U,NONOUE Y,et al.Ehd3,encoding a plant homeodomain finger-containing protein,is a critical promoter of rice flowering[J].The Plant Journal,2011,66(4):603-612.
[37]WU C Y,YOU C L,LI C S,et al.RID1,encoding a Cys2/His2-type zinc finger transcription factor,acts as a master switch from vegetative to floral development in rice[J].Proceedings of the National Academy of Sciences of the United States of America,2008,105(35):12915-12920.
[38]SHIBAYA T,HORI K,OGISO-TANAKA E,et al.Hd18,encoding histone acetylase related to Arabidopsis FLOWERING LOCUS D,is involved in the control of flowering time in rice[J].Plant and Cell Physiology,2016,57(9):1828-1838.
[39]RYU C H,LEE S Y,CHO L H,et al.Os MADS50 and Os MADS56 function antagonistically in regulating long day(LD)-dependent flowering in rice[J].Plant,Cell and Environment,2009,32(10):1412-1427.
[40]ZHAO J M,HUANG X,OUYANG X H,et al.Os ELF3-1,an ortholog of Arabidopsis early flowering 3,regulates rice circadian rhythm and photoperiodic flowering[J].Plo S one,2012,7(8):e43705.
[41]MATSUBARA K,OGISO-TANAKA E,HORI K,et al.Natural variation in Hd17,a homolog of Arabidopsis ELF3 that is involved in rice photoperiodic flowering[J].Plant and Cell Physiology,2012,53(14):709-716.
[42]NEMOTO Y,NONOUE Y,YANO M,et al.Hd1,a CONSTANS ortholog in rice,functions as an Ehd1 repressor through interaction with monocot-specific CCT-domain protein Ghd7[J].The Plant Jounal,2016,86(3):221-233.
[43]WU W X,ZHENG X M,LU G W,et al.Association of functional nucleotide polymorphisms at DTH2 with the northward expansion of rice cultivation in Asia[J].Proceedings of the National Academy of Sciences of the United States of America,2013,110(8):2775-2780.
[44]KIM S K,YUN C H,LEE J H,et al.Os CO3,a CONSTANS-LIKE gene,controls flowering by negatively regulating the expression of FT-like genes under SD conditions in rice[J].Planta,2008,228(2):355-365.
[45]LI D J,YANG C H,LI X B,et al.Sense and antisense Os Dof12 transcripts in rice[J].BMC Molecular Biology,2008,9:80.
[46]LIU T M,LIU H Y,ZHANG H,et al.Validation and characterization of Ghd7.1,a major quantitative trait locus with pleiotropic effects on spikelets per panicle,plant height,and heading date in rice(Oryza sativa L.)[J].Journal of Integrative Plant Biology,2013,55(10):917-927.
[47]薛为亚.水稻产量相关基因Ghd7的分离与鉴定[D].武汉:华中农业大学,2008.XUE W Y.Isolation and characterization of a pleiotropic gene,Ghd7,in rice[D].Wuhan:Huazhong Agricultural University,2008.
[48]LU L,YAN W H,XUE W Y,et al.Evolution and association analysis of Ghd7 in rice[J].Plo S ONE,2012,7(5):e34021.
[49]FUJINO K,YAMANOUCHI U,YANO M.Roles of the Hd5 gene controlling heading date for adaptation to the northern limits of rice cultivation[J].Theoretical and Applied Genetics,2013,126(3):611-618.
[50]FUJINO K,SEKIGUCHI H.Identification of QTLs conferring genetic variation for heading date among rice varieties at the northern-limit of rice cultivation[J].Breeding Science,2005,55:141-146.
[51]LI X F,SUN Y Q,TIAN X J,et al.Comprehensive identification of major flowering time genes and their combinations,which determined rice distribution in Northeast China[J].Plant Growth Regulation,2018,DOI:10.1007/s10725-017-0364-2.
[52]于秋竹.黑龙江省不同积温带水稻产量和品质及株型研究[D].沈阳:沈阳农业大学,2014.YU Q Z.Research of rice yield quality and plant type in different accumulated temperature zone in Heilongjiang province[D].Shenyang:Shenyang Agricultural University,2014.
[53]姜树坤,张凤鸣,白良明,等.黑龙江省第一积温带水稻育种趋势分析[J].作物杂志,2015(4):61-64.JIANG S K,ZHANG F M,BAI L M,et al.The Japonica rice breeding trend in 1staccumulated temperature zone of Heilongjiang province[J].Crops,2015(4):61-64.
[54]李秀峰.黑龙江省水稻抽穗期基因的效应研究及育种应用[D].北京:中国科学院大学,2016.LI X F.The study on effect of heading date genes in Heilongjiang rice cultivars and their application in molecular assistant breeding[D].Beijing:University of Chinese Academy of Sciences,2016.
[55]周文甲,田晓杰,任月坤,等.利用CRISPR/Cas9创造早熟香味水稻[J].土壤与作物,2017,6(2):146-52.ZHOU W J,TIAN X J,REN Y K,et al.Breeding of early-maturating and fragrant rice via CRISPR/Cas9 mediated genome editing[J].Soils and Crops,2017,6(2):146-52.
[56]LI X F,ZHOU W J,REN Y K,et al.High-efficiency breeding of early-maturing rice cultivars via CRISPR/Cas9-mediated genome editing[J].Journal of Genetics and Genomics,2017,44(2):175-178.
[57]CHEN S H,YANG Y,SHI W W,et al.Badh2,encoding betaine aldehyde dehydrogenase,inhibits the biosynthesis of 2-acetyl-1-pyrroline,a major component in rice fragrance[J].The Plant cell,2008,20(7):1850-1861.
[58]TIAN Z X,QIAN Q,LIU Q Q,et al.Allelic diversities in rice starch biosynthesis lead to a diverse array of rice eating and cooking qualities[J].Proceedings of the National Academy of Sciences,2009,106(51):21760-21765.