氮肥对水稻节间和叶鞘非结构性碳水化合物积累转运特征的影响
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摘要
以两优培九和扬稻6号为材料进行盆栽试验,研究低氮(N1)和高氮(N2)处理下水稻主茎上部3个节间和叶鞘非结构性碳水化合物(NSC)的积累转运特征及对氮肥用量的响应。结果表明:1)与N2处理相比,N1处理总体上显著促进灌浆早期两优培九倒2、倒3节间和叶鞘NSC的转运,对扬稻6号的影响差异不明显;N1处理显著增加了扬稻6号上部3个节间及叶鞘和两优培九倒3节间及叶鞘NSC表观转运量,也增加了两品种上部3个节间及叶鞘的NSC(总)表观贡献率。2)两个氮肥处理下,供试品种的叶鞘(两优培九倒3叶鞘除外)NSC转运量和表观贡献率大于节间。3)两优培九倒2、倒3节间及叶鞘NSC表观转运量和贡献率、上部3节间及叶鞘的NSC总转运量与总表观贡献率在两个氮肥处理下均高于扬稻6号。4)N1处理下,两优培九和扬稻6号倒2、倒3节间及叶鞘的转运量分别占总转运量的92%和75%,贡献率分别占91%和76%;而N2处理下两优培九倒2、倒3节间及叶鞘的转运量占90%,扬稻6号倒2、倒3节间则无输出。本研究表明水稻上部3个节间和叶鞘的NSC(总)转运量品种间存在差异,并受施氮量和节/叶位的影响。在水稻高产和减氮栽培中,通过选用适宜品种和优化氮肥管理,增加花前茎鞘NSC积累和花后NSC再分配,对提高水稻产量潜力和氮效率,特别是对花后逆境和减氮条件下产量稳定具有重要意义。
Two indica rice varieties,Liangyoupeijiu(LYP9)and Yangdao 6(YD6),were grown in pots under low(N1)and high(N2)nitrogen application rates.The accumulation and translocation of non-structural carbohydrates(NSC)in the top-most three internodes and leaf sheaths of main stems and its contributions to grain yield were investigated.1)Compared with N2 treatment,N1treatment enhanced NSC translocation of the second and third internodes and sheaths in LYP9 during the early grain filling stage,and also increased the apparent transferred mass of NSC(ATMNSC)of the three internodes from the top and sheaths in YD6 and the third internode from the top and sheath in LYP9,respectively.However,N1 treatment had no obvious effect on ATMNSC of the top first and second internodes and sheaths in LYP9.Noticeably,N1 treatment increased the apparent contributions of NSC to grain yield(ACNSC)of the three internodes and sheaths in both two varieties.2)Except the top-most third sheath in LYP9,the ATMNSCand ACNSCof sheaths were higher than those of internodes in both two varieties under N1 and N2treatments.3)Under N1 and N2treatments,ATMNSCand ACNSCof the top-most second and third internodes and sheaths,and total ATMNSCand ACNSCof the three internodes and sheaths in LYP9 were higher than those in YD6,respectively.4)Under N1 treatment,the top-most second and third internodes and sheaths totally contributed 92% and 91% of total ATMNSC and ACNSCof the three internodes and sheaths in LYP9,and 75% and 76%in YD6.Under N2 treatment,the top-most second and third internodes and sheaths totally contributed 90% and 91%in LYP9,and no NSC translocation from the top-most second and third internodes and totally accounted for 77% of total ACNSC.Our results indicated that ATMNSC of the three internodes and sheaths showed genotypic variation and depended on internode/sheath,and was affected by N application rate.Optimizing N fertilizer application and selecting suitable rice variety might enhance pre-anthesis NSC accumulation and post-anthesis NSC translocation,which could further increase rice yield potential and nitrogen use efficiency,especially stability of grain yield under adverse environments and nitrogen fertilizer-reducing practice.
Two indica rice varieties,Liangyoupeijiu(LYP9)and Yangdao 6(YD6),were grown in pots under low(N1)and high(N2)nitrogen application rates.The accumulation and translocation of non-structural carbohydrates(NSC)in the top-most three internodes and leaf sheaths of main stems and its contributions to grain yield were investigated.1)Compared with N2 treatment,N1treatment enhanced NSC translocation of the second and third internodes and sheaths in LYP9 during the early grain filling stage,and also increased the apparent transferred mass of NSC(ATMNSC)of the three internodes from the top and sheaths in YD6 and the third internode from the top and sheath in LYP9,respectively.However,N1 treatment had no obvious effect on ATMNSC of the top first and second internodes and sheaths in LYP9.Noticeably,N1 treatment increased the apparent contributions of NSC to grain yield(ACNSC)of the three internodes and sheaths in both two varieties.2)Except the top-most third sheath in LYP9,the ATMNSCand ACNSCof sheaths were higher than those of internodes in both two varieties under N1 and N2treatments.3)Under N1 and N2treatments,ATMNSCand ACNSCof the top-most second and third internodes and sheaths,and total ATMNSCand ACNSCof the three internodes and sheaths in LYP9 were higher than those in YD6,respectively.4)Under N1 treatment,the top-most second and third internodes and sheaths totally contributed 92% and 91% of total ATMNSC and ACNSCof the three internodes and sheaths in LYP9,and 75% and 76%in YD6.Under N2 treatment,the top-most second and third internodes and sheaths totally contributed 90% and 91%in LYP9,and no NSC translocation from the top-most second and third internodes and totally accounted for 77% of total ACNSC.Our results indicated that ATMNSC of the three internodes and sheaths showed genotypic variation and depended on internode/sheath,and was affected by N application rate.Optimizing N fertilizer application and selecting suitable rice variety might enhance pre-anthesis NSC accumulation and post-anthesis NSC translocation,which could further increase rice yield potential and nitrogen use efficiency,especially stability of grain yield under adverse environments and nitrogen fertilizer-reducing practice.
引文
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[2]Pan J F,Cui K H,Wei D,et al.Relationships of non-structural carbohydrates accumulation and translocation with yield formation in rice recombinant inbred lines under two nitrogen levels.Physiol Plant,2011,141(4):321-331.
[3]潘俊峰,崔克辉,向镜,等.不同库容量类型基因型水稻茎鞘非结构性碳水化合物积累转运特征.华中农业大学学报,2015,34(1):9-15.Pan J F,Cui K H,Xiang J,et al.Characteristics of non-structural carbohydrate accumulation and translocation in rice genotypes with various sink-capacity.J Huazhong Agric Univ,2015,34(1):9-15.(in Chinese with English abstract)
[4]王志琴,杨建昌,朱庆森,等.水稻抽穗期茎鞘中储存的可用性糖与籽粒充实的关系.江苏农学院学报,1997,18(4):13-17.Wang Z Q,Yang J C,Zhu Q S,et al.Relation of the usable carbohydrate reserved in stems and sheaths at heading stage with grain filling in rice plants.J Jiangsu Agric Coll,1997,18(4):13-17.(in Chinese with English abstract)
[5]Fu J,Huang Z H,Wang Z Q,et al.Pre-anthesis non-structural carbohydrate reserve in the stem enhances the sink strength of inferior spikelets during grain filling of rice.Field Crops Res,2011,123(2):170-182
[6]Kashiwagi T,Ishimaru K.Identification and functional analysis of a locus for improvement of lodging resistance in rice.Plant Physiol,2004,134(2):676-683.
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[10]Isopp H,Frehner M,Almeida J P F,et al.Nitrogen plays a major role in leaves when source-sink relations change:C and N metabolism in Loliumperenne growing under free air CO2enrichment.Aust J Plant Physiol,2000,27(9):851-858.
[11]Takahashi T,Chevalier P M,Rupp R A.Storage and remobilization of soluble carbohydrates after heading in different plant parts of a winter wheat cultivar.PlantProdSci,2001,4(3):160-165.
[12]Sugimura Y,Michiyama H,Hirano T.Involvement ofα-amylase genes in starch degradation in rice leaf sheaths at the post-heading stage.Plant Prod Sci,2015,18(3):277-283.
[13]李国辉,钟旭华,田卡,等.施氮对水稻茎秆抗倒伏能力的影响及其形态和力学机理.中国农业科学,2013,46(7):1323-1334.Li G H,Zhong X H,Tian K,et al.Effect of nitrogen application on stem lodging resistance of rice and its morphological and mechanical mechanisms.Sci Agric Sin,2013,46(7):1323-1334.(in Chinese with English abstract)
[14]曾建敏,崔克辉,黄见良,等.水稻生理生化特性对氮肥的反应及与氮利用效率的关系.作物学报,2007,33(7):1168-1176.Zeng J M,Cui K H,Huang J L,et al.Responses of physio-biochemical properties to n-fertilizer application and its relationship with nitrogen use efficiency in rice(Oryza sativa L.).Acta Agron Sin,2007,33(7):1168-1176.(in Chinese with English abstract)
[15]Scofield G N,Ruuska S A,Aoki N,et al.Starch storage in the stems of wheat plants:Localization and temporal changes.Ann Bot(Lond),2009,103(6):859-868.
[16]Fukushima A,Akita S.Varietal differences of the course and differentiation time of large vascular bundles in the rachis of rice.Jpn J Crop Sci,1997,66(1):24-28.
[17]Liu G L,Mei H W,Yu X Q,et al.QTL analysis of panicle neck diameter,a trait highly correlated with panicle size,under well-watered and drought conditions in rice(Oryza sativa L.).Plant Sci,2008,174(1):71-77.
[18]Ruuska S A,Lewis D C,Kennedy G,et al.Large-scale transcriptome analysis of the effects of nitrogen nutrition on accumulation of stem carbohydrate in reproductive-stage wheat.Plant Mol Biol,2008,66(1-2):15-32.
[19]郭兆武,萧浪涛,罗孝和,等.超级杂交稻“两优培九”剑叶叶鞘的光合功能.作物学报,2007,33(9):1508-1515.Guo Z W,Xiao L T,Luo X H,et al.Photosynthetic function of the flag leaf sheath for super hybrid rice Liangyoupeijiu.Acta Agron Sin,2007,33(9):1508-1515.(in Chinese with English abstract)
[20]Nagata K,Yoshinaga S,Takanashi J,et al.Effects of dry matter production,translocation of nonstructural carbohydrates and nitrogen application on grain filling in rice cultivar Takanari,a cultivar bearing a large number of spikelets.Plant Prod Sci,2001,4(3):173-183.
[21]Hirano T,Saito Y,Ushimaru H,et al.The effect of the amount of nitrogen fertilizer on starch metabolism in leaf sheath of japonica and indicarice varieties during the heading period.Plant Prod Sci,2005,8(2):122-130.
[22]潘俊峰,李国辉,崔克辉.水稻茎鞘非结构性碳水化合物再分配及其在稳产和抗逆中的作用.中国水稻科学,2014,28(4):335-342.Pan J F,Li G H,Cui K H.Re-partitioning of non-structural carbohydrates in rice stems and their roles in increasing yield stability and stress tolerance.Chin J Rice Sci,2014,28(4):335-342.(in Chinese with English abstract)
[23]Gebbing T,Schnyder H.Pre-anthesis reserve utilization for protein and carbohydrate synthesis in grains of wheat.Plant Physiol,1999,121(3):871-878.
[24]薛艳凤,郎有忠,吕川根,等.两优培九及其父本扬稻6号抽穗后叶片与根系衰老特点的研究.扬州大学学报:农业与生命科学版,2009,29(3):7-11.Xue Y F,Lang Y Z,Lv C G,et al.Study on leaf and root senescence of Liangyoupeijiu and it′s male parent Yangdao 6after heading.J Yangzhou Univ,2009,29(3):7-11.(in Chinese with English abstract)
[2]Pan J F,Cui K H,Wei D,et al.Relationships of non-structural carbohydrates accumulation and translocation with yield formation in rice recombinant inbred lines under two nitrogen levels.Physiol Plant,2011,141(4):321-331.
[3]潘俊峰,崔克辉,向镜,等.不同库容量类型基因型水稻茎鞘非结构性碳水化合物积累转运特征.华中农业大学学报,2015,34(1):9-15.Pan J F,Cui K H,Xiang J,et al.Characteristics of non-structural carbohydrate accumulation and translocation in rice genotypes with various sink-capacity.J Huazhong Agric Univ,2015,34(1):9-15.(in Chinese with English abstract)
[4]王志琴,杨建昌,朱庆森,等.水稻抽穗期茎鞘中储存的可用性糖与籽粒充实的关系.江苏农学院学报,1997,18(4):13-17.Wang Z Q,Yang J C,Zhu Q S,et al.Relation of the usable carbohydrate reserved in stems and sheaths at heading stage with grain filling in rice plants.J Jiangsu Agric Coll,1997,18(4):13-17.(in Chinese with English abstract)
[5]Fu J,Huang Z H,Wang Z Q,et al.Pre-anthesis non-structural carbohydrate reserve in the stem enhances the sink strength of inferior spikelets during grain filling of rice.Field Crops Res,2011,123(2):170-182
[6]Kashiwagi T,Ishimaru K.Identification and functional analysis of a locus for improvement of lodging resistance in rice.Plant Physiol,2004,134(2):676-683.
[7]Perez C M,Palmiano E,Baun L C,et al.Starch metabolism in the leaf sheaths and culm of rice.Plant Physiol,1971,47(3):404-408.
[8]He H Y,Koike M,Ishimaru T,et al.Temporal and spatial variations of carbohydrate content in rice leaf sheath and their varietal differences.Plant Prod Sci,2005,8(5):546-552.
[9]Ishimaru K,Kosone M,Sasaki H,et al.Leaf contents differ depending on the position in a rice leaf sheath during sinksource transition.Plant Physiol Biochem,2004,42(11):855-860.
[10]Isopp H,Frehner M,Almeida J P F,et al.Nitrogen plays a major role in leaves when source-sink relations change:C and N metabolism in Loliumperenne growing under free air CO2enrichment.Aust J Plant Physiol,2000,27(9):851-858.
[11]Takahashi T,Chevalier P M,Rupp R A.Storage and remobilization of soluble carbohydrates after heading in different plant parts of a winter wheat cultivar.PlantProdSci,2001,4(3):160-165.
[12]Sugimura Y,Michiyama H,Hirano T.Involvement ofα-amylase genes in starch degradation in rice leaf sheaths at the post-heading stage.Plant Prod Sci,2015,18(3):277-283.
[13]李国辉,钟旭华,田卡,等.施氮对水稻茎秆抗倒伏能力的影响及其形态和力学机理.中国农业科学,2013,46(7):1323-1334.Li G H,Zhong X H,Tian K,et al.Effect of nitrogen application on stem lodging resistance of rice and its morphological and mechanical mechanisms.Sci Agric Sin,2013,46(7):1323-1334.(in Chinese with English abstract)
[14]曾建敏,崔克辉,黄见良,等.水稻生理生化特性对氮肥的反应及与氮利用效率的关系.作物学报,2007,33(7):1168-1176.Zeng J M,Cui K H,Huang J L,et al.Responses of physio-biochemical properties to n-fertilizer application and its relationship with nitrogen use efficiency in rice(Oryza sativa L.).Acta Agron Sin,2007,33(7):1168-1176.(in Chinese with English abstract)
[15]Scofield G N,Ruuska S A,Aoki N,et al.Starch storage in the stems of wheat plants:Localization and temporal changes.Ann Bot(Lond),2009,103(6):859-868.
[16]Fukushima A,Akita S.Varietal differences of the course and differentiation time of large vascular bundles in the rachis of rice.Jpn J Crop Sci,1997,66(1):24-28.
[17]Liu G L,Mei H W,Yu X Q,et al.QTL analysis of panicle neck diameter,a trait highly correlated with panicle size,under well-watered and drought conditions in rice(Oryza sativa L.).Plant Sci,2008,174(1):71-77.
[18]Ruuska S A,Lewis D C,Kennedy G,et al.Large-scale transcriptome analysis of the effects of nitrogen nutrition on accumulation of stem carbohydrate in reproductive-stage wheat.Plant Mol Biol,2008,66(1-2):15-32.
[19]郭兆武,萧浪涛,罗孝和,等.超级杂交稻“两优培九”剑叶叶鞘的光合功能.作物学报,2007,33(9):1508-1515.Guo Z W,Xiao L T,Luo X H,et al.Photosynthetic function of the flag leaf sheath for super hybrid rice Liangyoupeijiu.Acta Agron Sin,2007,33(9):1508-1515.(in Chinese with English abstract)
[20]Nagata K,Yoshinaga S,Takanashi J,et al.Effects of dry matter production,translocation of nonstructural carbohydrates and nitrogen application on grain filling in rice cultivar Takanari,a cultivar bearing a large number of spikelets.Plant Prod Sci,2001,4(3):173-183.
[21]Hirano T,Saito Y,Ushimaru H,et al.The effect of the amount of nitrogen fertilizer on starch metabolism in leaf sheath of japonica and indicarice varieties during the heading period.Plant Prod Sci,2005,8(2):122-130.
[22]潘俊峰,李国辉,崔克辉.水稻茎鞘非结构性碳水化合物再分配及其在稳产和抗逆中的作用.中国水稻科学,2014,28(4):335-342.Pan J F,Li G H,Cui K H.Re-partitioning of non-structural carbohydrates in rice stems and their roles in increasing yield stability and stress tolerance.Chin J Rice Sci,2014,28(4):335-342.(in Chinese with English abstract)
[23]Gebbing T,Schnyder H.Pre-anthesis reserve utilization for protein and carbohydrate synthesis in grains of wheat.Plant Physiol,1999,121(3):871-878.
[24]薛艳凤,郎有忠,吕川根,等.两优培九及其父本扬稻6号抽穗后叶片与根系衰老特点的研究.扬州大学学报:农业与生命科学版,2009,29(3):7-11.Xue Y F,Lang Y Z,Lv C G,et al.Study on leaf and root senescence of Liangyoupeijiu and it′s male parent Yangdao 6after heading.J Yangzhou Univ,2009,29(3):7-11.(in Chinese with English abstract)