籼粳交超级稻甬优538花后氮素积累模型与特征分析

韦还和　孟天瑶　李　超　张洪程,*　史天宇　马荣荣　王晓燕杨筠文　戴其根,*　霍中洋　许　轲　魏海燕　郭保卫

1扬州大学农业部长江流域稻作技术创新中心/江苏省作物遗传生理重点实验室,江苏扬州 225009;2浙江省宁波市农业科学院作物研究所,浙江宁波 315101;3浙江省宁波市种子公司,浙江宁波 315101;4浙江省宁波市鄞州区农业技术服务站,浙江宁波 315100



籼粳交超级稻甬优538花后氮素积累模型与特征分析

韦还和1孟天瑶1李超1张洪程1,*史天宇1马荣荣2王晓燕3杨筠文4戴其根1,*霍中洋1许轲1魏海燕1郭保卫1

1扬州大学农业部长江流域稻作技术创新中心/江苏省作物遗传生理重点实验室,江苏扬州 225009;2浙江省宁波市农业科学院作物研究所,浙江宁波 315101;3浙江省宁波市种子公司,浙江宁波 315101;4浙江省宁波市鄞州区农业技术服务站,浙江宁波 315100

摘要:以甬优籼粳交超级稻甬优538为试材,常规粳稻镇稻18和杂交籼稻中浙优1号为对照,研究甬优籼粳杂交稻花后氮素积累特征以及比较不同类型品种花后氮素积累特征差异。结果表明:(1)两年中,甬优538平均产量为12.5 t hm(-2),显著高于对照。抽穗期、成熟期及抽穗-成熟期氮素积累量均呈甬优538>镇稻18>中浙优1号。百千克籽粒吸氮量以镇稻18最高,甬优538最低;氮肥偏生产力则以甬优538最高,镇稻18最低。甬优538花后茎鞘氮素转运量和转运率、穗部氮素积累量显著高于对照。(2)镇稻18和甬优538花后各时期氮素积累量均高于中浙优1号,且各品种花后氮素积累量与花后天数均以Richards方程拟合效果较好(R2均大于0.995)。分析特征参数,花后最大氮素积累速率、花后平均氮素积累速率及花后到达最大氮素积累速率的时间均呈镇稻18>中浙优1号>甬优538,花后氮素有效吸收时间则呈甬优538>镇稻18>中浙优1号。花后氮素积累渐增期、缓增期的持续天数、积累速率和积累量以镇稻18最高、甬优538最低,快增期的持续天数和积累量以甬优538显著高于对照。本研究结果表明,甬优538花后较强的氮素积累优势集中在快增期,此期氮素积累量占其花后氮素积累总量的86.1％ (两年平均值)。且甬优538在花后快增期较强的氮素积累能力主要在于其较长的快增期持续天数。

关键词:籼粳交超级稻;品种类型;花后氮素积累

本研究由农业部超级稻专项(02318802013231),国家公益性行业(农业)科研专项(201303102),宁波市重大科技项目(2013C11001),江苏省重点研发项目(BE2015340)和扬州大学研究生创新培养计划项目(KYLX15_1371),扬州大学科技创新培育基金(2015CXJ042)和基于模型与GIS的高邮市小麦精确管理和诊断调控技术的开发与示范推广(SXGC[2013]248)资助。

This study was supported by the Special Program of Super Rice of the Ministry of Agricultural (02318802013231),China Special Fund for Agro-scientific Research in the Public Interest (201303102),the Major Technology Project of Ningbo City (2013C11001),the Key Projects of Jiangsu Province (BE2015340),and Innovative Program for Graduate Students of Yangzhou University (KYLX15_1371),Science and Technology Innovation Fund of Yangzhou University (2015CXJ042),and Precise Diagnosis and Management of Control Technology Based on Modelling and GIS of Gaoyou City (SXGC[2013]248).

第一作者联系方式:E-mail:920964110@qq.com

水稻花后氮素积累水平是影响花后光合物质积累与分配、调节籽粒蛋白质及组分含量、影响加工和食味品质的重要因素。因此,定量分析水稻花后氮素积累的动态变化是揭示作物产量形成机理和稻米品质研究中的重要内容[1-3]。当前就禾谷类作物氮素积累动态已有相关报道,但大都侧重于全生育期,如水稻方面,孙成明等[4]报道了武香粳14在FACE处理下的氮素动态变化,纪洪亭等[5]研究了超级杂交稻养分积累动态模型及相关特征。小麦方面,丁锦峰等[6]研究了长江中下游稻茬小麦超高产群体氮素积累、分配与利用特征,党红凯等[7]分析了超高产冬小麦对氮素的吸收、积累等方面特征。此外,王宜伦等[8]、景立权等[9]也就超高产玉米群体氮素积累与分配进行了相关研究。与大面积种植的常规粳稻和杂交籼稻相比,甬优系列籼粳杂交稻已在生产上表现出较明显的产量优势[10-11]。当前就甬优系列籼粳杂交稻的研究多集中在产量优势形成机制方面,如分析产量构成因素、光合物质生产、茎秆特性、根系形态生理等[12-16]。就甬优系列籼粳杂交稻氮磷钾养分积累、吸收等的报道较少,且与常规粳稻和杂交籼稻相比,甬优系列籼粳杂交稻在花后氮素积累动态及特征上的差异分析迄今尚缺乏报道。本研究采用作物生长模型对不同类型水稻品种氮素积累与花后天数的关系进行曲线估计,利用推导出的特征参数定量分析不同类型品种花后氮素积累特征及其差异,以期为甬优系列籼粳杂交稻花后氮素积累动态和管理决策提供量化工具和实践依据。

1　材料与方法

1.1试验材料与栽培管理概况

以甬优籼粳交品种甬优538为试材、常规粳稻镇稻18和杂交籼稻中浙优1号为对照。2012—2013年参试品种的主要生育期见表1。

表1　主要生育期以及生育阶段天数Table 1　Duration of growing stages of the tested varieties

试验于2012—2013年在浙江省宁波市鄞州区洞桥镇百梁桥村进行。土壤类型为黄化青紫泥田,pH 5.51、含有机质38.37 g kg-1、全氮0.16％、碱解氮82.45 mg kg-1、速效磷20.14 mg kg-1、速效钾78.45 mg kg-1、水溶性盐总量0.13 g kg-1。两年中参试品种水稻生长期间的月平均温度、日照时数、降雨量见表2。

表2　2012-2013年水稻生长期间的月平均温度、日照时数和降雨量Table 2　Mean temperature,sunshine hours,and rainfall during rice growing seasons in 2012-2013

试验采取完全随机区组设计,小区面积20 m2,3次重复。小区间作埂隔离,并用塑料薄膜覆盖埂体,保证单独排灌。毯苗育秧,秧龄20 d,栽插株行距为30.0cm×13.2cm。对籼粳杂交稻甬优538、杂交籼稻中浙优1号每穴2苗栽插,常规粳稻镇稻18每穴4苗栽插。对甬优538和镇稻18施纯氮270 kg hm-2,中浙优1号施纯氮 225 kg hm-2,氮肥按基蘖肥∶穗粒肥=6∶4施用。各小区磷、钾肥施用量一致,即施过磷酸钙(含12％ P2O5) 1150 kg hm-2,全部基施。氯化钾(含60％ K2O) 450 kg hm-2,按基蘖肥∶穗粒肥= 4∶6施用。移栽后采用湿润灌溉为主,建立浅水层;群体达到目标穗数的80％时搁田,控制无效分蘖发生;抽穗扬花期田间保持3 cm水层,灌浆结实期间歇灌溉,干湿交替,收割前7 d断水搁田。按常规高产栽培要求防治病虫害。

1.2测定项目与方法

1.2.1花后氮素积累动态甬优538于花后0 d(抽穗期)田取样,此后每隔5 d取一次样,直至花后第65天。镇稻18于花后0 d (抽穗期)田取样,此后每隔5 d取一次样,直至花后50 d。中浙优1号于花后0 d (抽穗期)田取样,此后每隔5 d取一次样,直至花后45 d。每次取样均按各小区的平均茎蘖数取5穴植株,样品分成叶、茎鞘、穗3个部分,105℃杀青30 min,75℃烘干至恒重后,测定干物质量。称重后样品用万能粉碎机粉碎,过80目筛后采用半微量凯氏定氮法测定氮素含量。

1.2.2计算方法与数据处理

籽粒生产率(kg grain kg-1)=籽粒产量/成熟期植株氮素吸收量氮肥偏生产力(kg kg-1)=施氮区产量/小区施氮量氮素转运量(kg hm-2)=抽穗期茎鞘、叶氮素积累量-成熟期茎鞘、叶氮素积累量

氮素转运率(％)=单位面积植株成熟期叶、茎鞘氮素的表观转运量/抽穗期叶、茎鞘氮素积累量×100

氮素转运贡献率(％)=氮素转运量/抽穗至成熟期穗部氮素积累量×100

本文以Curve Expert 1.3软件对各品种花后天数(抽穗当天为0)和群体氮素积累量之间的关系进行拟合,发现用Richards方程拟合的效果较好(R2均大于0.995)。Richards方程式中,W为花后各期群体氮素积累量(kg hm-2),A为终极群体氮素积累量(kg hm-2),t为开花后天数(d),B、N、K为方程参数。参照朱庆森等[17]的方法,求出以下参数。

花后氮素积累阶段的渐增期(0-t1)、快增期(t1-t2)和缓增期(t2-t3)计算公式如下:

渐增期平均氮素积累速率

缓增期平均氮素积累速率(kg hm-2d-1)

1.3数据处理

运用Microsoft Excel软件录入数据、计算,用DPS软件作统计分析。

2　结果与分析

2.1产量及花后氮素吸收量

两年中,甬优538平均产量为12.5 t hm-2,显著高于镇稻18和中浙优1号。抽穗期氮素积累量呈甬优538>镇稻18>中浙优1号,成熟期和抽穗-成熟期氮素积累量也呈类似趋势。分析氮素利用指标,两年中甬优538籽粒生产率为63.7 kg grain kg-1,显著高于镇稻18 (55.3 kg grain kg-1)和中浙优1号(59.8 kg grain kg-1);氮肥偏生产力以甬优538最高,镇稻18最低(表3)。

表3　产量及花后氮素吸收量Table 3　Grain yield and nitrogen uptake after heading in the varieties

抽穗至成熟期阶段,叶片氮素转运量和转运率以镇稻18最高、中浙优1号最低;茎鞘氮素转运量和转运率以甬优538最高、镇稻18最低;两年中甬优538穗部氮素养分增加量为79.8 kg hm-2,显著高于镇稻18 (75.4 kg hm-2)和中浙优1号(64.4 kg hm-2);花后氮素转运贡献率以甬优538和镇稻18显著高于中浙优1号(表4)。

2.2花后氮素积累动态

甬优538、镇稻18和中浙优1号花后氮素积累均大致呈慢-快-慢的趋势。镇稻18和甬优538花后各时期氮素积累量均高于中浙优1号(图1)。各品种花后氮素积累速率均呈先缓慢上升后下降趋势。在到达花后最大氮素积累速率前,甬优538氮素积累速率低于镇稻18和中浙优1号。与镇稻18和中浙优1号相比,甬优538最大氮素积累速率值在到达花后呈较平缓下降趋势(图2)。

表4　抽穗至成熟期叶片、茎鞘氮素的转运Table 4　Nitrogen transferred from leaf,stem and sheath from heading to maturity

图1　各品种花后氮素积累量动态Fig.1　Dynamics in nitrogen uptake after heading in the tested varieties

图2　各品种花后氮素积累速率动态Fig.2　Dynamics in nitrogen uptake rate after heading in the tested varieties

2.3花后氮素积累拟合方程及氮素吸收参数

利用Curve Expert 1.3 软件对各品种花后天数(抽穗当天为0)和群体氮素积累量之间的关系进行拟合,发现用Richards方程拟合的效果较好(R2均大于0.995),具体方程见表5。分析方程参数,两年中镇稻18花后最大氮素积累速率和花后平均氮素积累速率分别为0.7473、0.3841 kg hm-2d-1,显著高于中浙优1号(0.6834、0.3534 kg hm-2d-1)和甬优538 (0.6052、0.3186 kg hm-2d-1);花后达到最大氮素吸收速率的时间亦以镇稻18最高、甬优538最低;花后氮素有效吸收时间呈甬优538 (48.3 d)>镇稻18 (39.3 d)>中浙优1号(39.9 d)(表6)。

表5　各品种花后氮素积累的拟合方程Table 5　Simulation equations of nitrogen uptake after heading in the tested varieties

表6　各品种花后氮素积累参数Table 6　Parameters of nitrogen uptake after heading among the tested varieties

分析花后氮素积累的渐增、快增、缓增期特征(表7)。花后氮素积累渐增期和缓增期的平均积累速率和阶段天数呈镇稻18>中浙优1号>甬优538;花后渐增期和缓增期的氮素积累量亦以镇稻18>中浙优1号>甬优538。甬优538花后快增期氮素积累量为22.59 kg hm-2,显著高于镇稻18 (10.72 kg hm-2)和中浙优1号(12.20 kg hm-2);甬优538花后快增期天数为40.4 d,显著高于对照。

表7　各品种花后氮素积累渐增、快增、缓增期3个阶段的特征Table 7　Characteristics of nitrogen uptake in the early,middle and late stage after heading among the tested varieties

3　讨论

3.1不同类型水稻品种氮素利用效率

当前,在提高水稻产量的同时,如何协同提高氮肥利用效率是水稻研究和生产中面临的一大难题,高产氮高效品种的选育也一直是育种和栽培领域关注的热点[18-19]。水稻氮素利用参数包括氮肥吸收利用率、氮肥生理利用率、籽粒生产率、百千克籽粒吸氮量、氮肥偏生产力、农学利用率等[20]。目前,就不同基因型水稻品种氮素利用效率差异已有较多研究[20-23]。龚金龙等[21]研究表明,粳稻氮素生理利用率、氮素吸收利用率和农学利用率高于籼稻,干物质生产效率和氮肥偏生产力则显著低于籼稻。孟天瑶等[22]研究表明,甬优籼粳杂交稻氮肥偏生产力显著高于常规粳稻、杂交籼稻和杂交粳稻。Koutroubas 等[23]研究地中海地区种植的粳稻和籼稻的产量和氮素利用率表明,籼稻品种籽粒生产率高于粳稻品种,这主要是由于其较高的氮收获指数。本试验因未设置氮素空白对照处理,因此选择籽粒生产率和氮肥偏生产力分析不同基因型品种氮素利用效率差异,结果表明,两年中甬优538平均产量为12.5 t hm-2,显著高于镇稻18 (10.6 t hm-2)和中浙优1号(10.3 t hm-2)。籽粒生产率和氮肥偏生产力以甬优538最高,镇稻18最低。该结果说明,甬优538较好地协同提高了籽粒产量和氮肥利用效率。此外,常规粳稻镇稻18籽粒生产率和氮肥偏生产力低于杂交籼稻中浙优1号。

3.2不同类型水稻品种花后氮素转运特征

不同基因型水稻花后氮素转移特征存在明显差异。赵敏等[24]研究表明,中籼中熟杂交稻较中籼迟熟杂交稻和粳稻具有较高的氮素积累、转运和利用效率。霍中洋等[25]研究表明,与产量较低群体相比,产量较高群体具有花后茎鞘氮素转运量和转运率适中、以及较高的叶片氮素转运量、转运率和穗部氮素积累量。龚金龙等[21]研究表明,粳稻花后叶片氮素转运量低于籼稻,茎鞘氮素转运量则显著高于籼稻。董桂春等[26]研究表明,氮素籽粒生产效率较高的品种具有灌浆结实期叶、茎鞘氮素转运量大、运转率高等特点。李敏等[27]研究发现,较中等生产力类型,高生产力品种花后茎叶氮素转运量和转运率均较高。本试验条件下,产量较高的甬优538花后穗部氮素积累量、氮素转运贡献率均显著高于对照,茎鞘氮素转运量和转运率亦高于对照,叶片氮素转运量和转运率则介于镇稻18和中浙优1号之间。该结果表明,与对照相比,甬优538具有花后叶片、茎鞘氮素转运量大、转运率较高的特点。

3.3不同类型水稻品种花后氮素积累模型及其特征

养分积累模拟模型是作物生长模拟系统的重要组成部分。纪洪亭等[5]采用Gompertz模型拟合了超级杂交稻干物质和养分积累动态。赵敏等[24]研究表明育插秧机械化条件下水稻植株氮素积累符合Logistic曲线增长规律。且近几年对玉米和棉花等作物氮素吸收动态都直接采用Logistic模型进行拟合[28-29]。杨京平等[30]借助水稻生长模型Oryza-0较好地模拟了水稻氮吸收和积累动态。本试验条件下,籼粳杂交稻甬优538、常规粳稻镇稻18、杂交籼稻中浙优1号花后氮素积累均以Richards模型拟合效果最好(R2均大于0.995),且Richards模型在水稻籽粒灌浆拟合[17,31]、干物质积累动态[32]等方面都有较好的应用。

此前,基于作物生长模型对作物养分吸收积累特征的描述多侧重于全生育期,而对花后养分吸收积累特征的研究较少。丁锦峰等[33]利用Richards模型拟合了小麦全生育期积累动态。郭文琦等[34]采用Logistic模型拟合了棉花花后氮、磷、钾积累动态。在水稻方面,纪洪亭等[5]研究表明,超级杂交稻氮积累的最大速率出现在孕穗前10 d,氮积累的快增期出现在拔节前12 d至抽穗期,此期氮素积累量占氮积累总量的65.6％。章明清等[35]报道了杂交籼稻根系吸收氮、磷、钾的动态模拟模型,结果表明,水稻根系吸收氮、磷、钾的积累吸收量在整个生育期呈“S”型特征,在生育前期随生育进程迅速提高并达最大值,随后按指数规律下降,且根系对氮、磷、钾的吸收速率基本同步。陈洁等[36]以两优培九、武育粳3号和武香粳14为试材,研究水稻植株氮素吸收特征表明,花前叶片和茎中的相对氮含量随播后生长度日线性增加;花后叶片和茎中的相对氮含量随花后生长度日线性递减;花后吸收的氮素随籽粒重的增加对数递增。本研究表明,甬优538抽穗至成熟期氮素积累量为28.1 kg hm-2,显著高于镇稻18 (26.7 kg hm-2)和中浙优1号(24.3 kg hm-2)。花后最大氮素积累速率和花后平均氮素积累速率均以镇稻18最高、甬优538最低。甬优538、镇稻18和中浙优1号分别在花后27.6、30.8和28.8 d达到最大氮素吸收速率。花后氮素有效吸收时间呈甬优538 (48.3 d) >镇稻18 (39.3 d) >中浙优1号(39.9 d)。该结果表明,甬优538花后较强的氮素积累量主要是由于其较长的花后氮素有效吸收天数,而非花后氮素吸收速率。

纪洪亭等[5]研究表明,超级杂交籼稻氮积累的快增期出现在拔节前12 d至抽穗期,此期氮素积累量占氮积累总量的65.6％,提出超级杂交籼稻养分积累的优势在于快增期持续天数长、中后期养分积累速率较快。章明清等[35]研究表明,早稻和晚稻在栽后50～60 d根系氮、磷、钾养分吸收能力达到峰值,且早稻和晚稻根系最大养分吸收量的时间均出现在生育后期。本试验条件下,花后渐增期和缓增期的氮素积累量以镇稻18最高、甬优538最低;甬优538花后快增期氮素积累量为22.59 kg hm-2,显著高于镇稻18 (10.72 kg hm-2)和中浙优1号(12.20 kg hm-2),此期氮素积累量占其花后氮素积累总量的86.1％ (两年平均值);此外,甬优538花后快增期天数为40.4 d,显著高于对照;而花后快增期阶段的平均氮素积累速率则显著低于对照。该结果说明,甬优538花后氮素积累优势集中在快增期,且甬优538在花后快增期较强的氮素积累能力主要在于其较长的快增期持续天数。

4　结论

以Richards模型拟合花后氮素积累动态效果最好。甬优538花后氮素积累量显著高于对照,其花后较强的氮素积累优势主要集中在快增期,且快增期较强的氮素积累能力主要在于其较长的快增期持续天数。

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URL:http://www.cnki.net/kcms/detail/11.1809.S.20160125.1622.002.html

Dynamic Model and Its Characteristics Analysis for Nitrogen Accumulation after Heading in Yongyou 538

WEI Huan-He1,MENG Tian-Yao1,LI Chao1,ZHANG Hong-Cheng1,*,SHI Tian-Yu1,MA Rong-Rong2,WANG Xiao-Yan3,YANG Jun-Wen4,DAI Qi-Gen1,*,HUO Zhong-Yang1,XU Ke1,WEI Hai-Yan1,and GUO Bao-Wei11Innovation Center of Rice Cultivation Technology in Yangtze River Valley,Ministry of Agriculture/Key Laboratory of Crop Genetics and Physiology of Jiangsu Province,Yangzhou University,Yangzhou 225009,China;2Crop Research Institute,Ningbo Academy of Agricultural Sciences of Zhejiang Province,Ningbo 315101,China;3Ningbo Seed Company of Zhejiang Province,Ningbo 315101,China;4Agricultural Technology Extension and Service of Yinzhou District,Ningbo City,Zhejiang Province,Ningbo 315100,China

Abstract:A field experiment was conducted using indica/japonica hybrid super rice Yongyou 538 as the material,conventional japonica rice Zhendao 18 and hybrid indica rice Zhongzheyou 1 as the check to study the characteristics of nitrogen accumulation after heading and compare the differences in characteristics of nitrogen accumulation after heading in different types of rice variety.Resultsbook=541,ebook=80followed that,on an average across two years,grain yield of Yongyou 538 was 12.5 t ha(-1),significantly higher than that of check varieties.Nitrogen accumulation at heading,at maturity,and from heading to maturity showed a trend of Yongyou 538 >Zhendao 18 >Zhongzheyou 1.Nitrogen absorption of 100 kg seeds of Zhendao 18 was the highest,followed by Zhongzheyou 1,and Yongyou 538,while opposite trends was observed for N partial factor productivity.Compared with the check,nitrogen output from stem and nitrogen increase in panicle of Yongyou 538 were consistently higher.Nitrogen accumulation in Yongyou 538 and Zhendao 18 was both higher than that of Zhongzheyou 1 after heading,and Richards' equation was fit to simulate the relationship between nitrogen accumulation and days after heading for three types of rice variety (R2≥ 0.995).Maximum rate of nitrogen accumulation,mean rate of nitrogen accumulation,and days to maximum rate of nitrogen accumulation after heading in Zhendao 18 were the highest,while those in Yongyou 538 were the lowest.Effective nitrogen accumulation duration showed a trend of Yongyou 538 >Zhendao 18 >Zhongzheyou 1.Duration days,nitrogen accumulation rate,and nitrogen accumulation in early and late stage of Zhendao 18 were the highest,while those of Yongyou 538 were the lowest.Duration days and nitrogen accumulation in middle stage of Yongyou 538 was consistently higher compared with the check.Our results indicated that greater nitrogen uptake of Yongyou 538 was mainly occurred in the middle stage,which was 86.1％ of the nitrogen accumulation from heading to maturity.And higher nitrogen uptake in middle stage after heading of Yongyou 538 was mainly attributed to the longer duration days in this stage.

Keywords:Indica/japonica hybrid super rice of Yongyou series;Different rice type of cultivars;Nitrogen accumulation after heading

收稿日期Received():2015-10-25;Accepted(接受日期):2016-01-11;Published online(网络出版日期):2016-01-25.

*通讯作者(

Corresponding authors):张洪程,E-mail:hczhang@yzu.edu.cn;戴其根,E-mail:qgdai@yzu.edu.cn

DOI:10.3724/SP.J.1006.2016.00540