甬优系列籼粳杂交稻的冠层结构与光合特性

引用本文

姜元华, 许轲, 赵可, 孙建军, 韦还和, 许俊伟, 魏海燕, 郭保卫, 霍中洋, 戴其根, 张洪程. 甬优系列籼粳杂交稻的冠层结构与光合特性. 41(2): 286-296
JIANG Yuan-Hua, XU Ke, ZHAO Ke, SUN Jian-Jun, WEI Huan-He, XU Jun-Wei, WEI Hai-Yan, GUO Bao-Wei, HUO Zhong-Yang, DAI Qi-Gen, ZHANG Hong-Cheng. Canopy Structure and Photosynthetic Characteristics of Yongyou Series of Indica-Japonica Hybrid Rice under High-yielding Cultivation Condition . Acta Agronomica Sinica, 41(2): 286-296 复制到剪切板

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作物学报编辑部

甬优系列籼粳杂交稻的冠层结构与光合特性

姜元华, 许轲, 赵可, 孙建军, 韦还和, 许俊伟, 魏海燕, 郭保卫, 霍中洋, 戴其根, 张洪程^*

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

^* 通讯作者(Corresponding author): 张洪程, E-mail: hczhang@yzu.edu.cn, Tel: 0514-87979220

第一作者联系方式: E-mail: 447685778@qq.com, Tel: 15995114072

收稿日期:2014-09-30

基金:本研究由国家粮食丰产科技工程项目(2011BAD16B03), 国家公益性行业(农业)科研专项(201303102), 农业部超级稻专项(02318802013231)和江苏省科技支撑计划项目(BE2012301)资助;

摘要

旨在阐明甬优系列籼粳杂交稻的冠层结构与光合特性。以甬优籼粳杂交稻(A)为研究对象, 以三系杂交粳稻(B)、超级常规粳稻(C)和超级杂交籼稻(D)为对照, 在机插高产栽培条件下, 对不同类型水稻的叶片受光姿态和群体冠层光分布、光合特性系统比较研究。结果表明: (1)上3张叶片的长度、宽度表现为A>B>D>C, 叶基角表现为D>A>B>C, 披垂度表现为D>B>C>A。冠层上部叶面积密度表现为D>A>B>C, 冠层下部叶面积密度表现为C>B>A>D, 最大叶面积密度表现为A>D>B>C, 最大叶面积密度出现的相对高度表现为D>A>B>C。冠层上部相对光照表现为A>B>C>D, 冠层下部相对光照表现为B>A>C>D, 冠层平均相对光照表现为B>A>C>D, 冠层消光系数表现为C>D>B>A。(2)抽穗期群体叶面积指数、高效叶面积率均呈D>A>B>C趋势, 有效叶面积率呈A>B>C>D趋势; 颖花/叶和实粒/叶均表现为A>B>C>D。经济产量、生物产量均表现为A>B>C>D, 经济系数呈D>C>B>A趋势; 蜡熟期和成熟期剑叶的叶绿素含量、类胡萝卜素含量、PSII的光化学效率及净光合速率呈A>B>C>D趋势; 抽穗至成熟期剑叶的MDA含量呈D>C>B>A趋势, SOD、POD、CAT活性呈A>B>C>D趋势。与其他3种类型水稻相比, 甬优系列籼粳杂交稻的冠层结构与光合特性具有显著优势, 这是甬优系列杂交稻产量潜力正常发挥的生态生理基础, 也是进一步提高亚种间杂交稻群体生产力的重要途径。

关键词: 籼粳杂交稻; 甬优系列; 冠层结构; 光合特性

Canopy Structure and Photosynthetic Characteristics of Yongyou Series of Indica-Japonica Hybrid Rice under High-yielding Cultivation Condition

JIANG Yuan-Hua, XU Ke, ZHAO Ke, SUN Jian-Jun, WEI Huan-He, XU Jun-Wei, WEI Hai-Yan, GUO Bao-Wei, HUO Zhong-Yang, DAI Qi-Gen, ZHANG Hong-Cheng^*

Innovation Center of Rice Cultivation Technology in the Yangtze Valley, Ministry of Agriculture / Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China

Abstract

Compared with japonica hybrid rice cultivars(B), conventional japonica rice cultivars (C) and indica hybrid rice cultivars (D), Yongyou series of indica-japonica hybrid rice cultivars (A) were adopted to analyze the lightened posture of leave, light distribution of canopy and photosynthetic characteristics of plant systematically. Results were as follows: (1) leaf length, leaf width of top three leaves showed the trend of A>B>D>C, canopy extinction coefficient showed the trend of C>D>B>A, leaf basic angle of top three leaves and relative height of the largest leaf area density were D>A>B>C, lower canopy leaf area density was C>B>A>D, plant height and the largest leaf area density showed the trend of A>D>B>C, leaf drop angle of top three leaves showed the trend of D>B>C>A, upper canopy leaf area density observed D>A>B>C, lower canopy relative light showed the trend of A>B>C>D, lower canopy relative light and canopy average relative light showed the trend of B>A>C>D; (2) population LAI and high valid leaf area rate showed the trend of D>A>B>C at heading, valid leaf area rate, spikelets per cm² leaf area, filled grains per cm² leaf area, total biomass, economic yield, chlorophyll ( a+b) content, carotenoid content, PSII actual photochemical efficiency, net photosynthetic rate, enzyme activities of SOD, POD, and CAT after heading showed the trend of A>B>C>D, harvest index and MDA content showed the trend of D>C>B>A. Therefore, compared with other three types of rice, Yongyou Series of indica-japonica hybrid rice had significant advantages in the canopy structure and photosynthetic characteristics, which are both normal ecological and physiological bases of high yield, and maybe an important way to improve the population productivity of inter-subspecific hybrid rice.

Keyword: Indica-japonica hybrid rice; Yongyou series; Canopy structure; Photosynthetic characteristics

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持续提高单产是稻作研究的永恒主题。回顾我国水稻单产的两次重大飞跃(20世纪50年代末的矮化育种增产及70年代中期的三系杂交育种增产), 归结其根本的原因是水稻株型得到了改良^{[1, 2, 3]}。众多水稻学者一致认为, 优良的植株形态是实现水稻超高产第三次突破的骨架^{[4, 5, 6]}。因此, 水稻株型的研究越来越受到人们的重视, 群体冠层结构和光合特性是研究的重要内容之一。

前人围绕水稻高产、超高产的理想株型及其机能开展了大量研究, 并根据地域和品种特性等提出了诸多理想株型模式^{[7, 8, 9, 10, 11, 12]}, 也从水稻亚种属性^{[13, 14]}、品种演进^{[15, 16]}以及高产高效协同型品种^{[17, 18]}等角度对水稻的株型特征进行了深入研究, 如张洪程等^[13]研究认为与籼稻相比, 粳稻生育中期能形成适宜的叶面积指数, 较紧凑的株型以及较挺拔的叶片, 群体内部相对通风透光, 并且抽穗期群体干物质量积累适中, 较高的粒叶比保证最终库源关系较协调。杨建昌等^[15]研究发现, 品种改良明显减小了顶部3叶的着生角度, 增加了抽穗期的叶面积, 但颖花量的增加超过叶面积的增加, 导致粒叶比(颖花数与叶面积之比)提高。张庆等^[17]研究表明, 与低产低效型品种相比, 高产高效型品种水稻上三叶的叶长和叶披垂度较低, 上三叶的叶宽、群体LAI、有效LAI和比叶重等均较高。上述研究结果及其结论为水稻高产、优质、高效栽培及品种选用与改良提供了重要参考依据。

籼粳亚种间杂交稻蕴藏着巨大的杂种优势, 其应用一直被认为是进一步提高水稻产量的有效途径^{[19, 20]}。近年来, 浙江省宁波市农业科学院和宁波市种子有限公司从高光效株型及育种方法研究入手, 开展粳型核质互作雄性不育系和对粳型不育系强恢的恢复系及籼粳杂交稻强优组合的选育, 育成了甬优6号、甬优9号、甬优11、甬优12、甬优13、甬优15和甬优2640等多个典型籼粳杂交稻品种, 在生产试验和大面积应用中表现出较大超高产潜力, 产生了较显著的社会和经济效益, 多个品种已被农业部认定为主推超级稻品种。目前, 国内已有一些关于该系列组合的形态生理特征研究的报道^{[21, 22]}, 但供试材料数量相对较少, 对照类型也较单一, 并且缺乏高产栽培条件下的系统比较分析。与以往的超级稻或超高产水稻相比, 高产栽培条件下甬优系列杂交稻的冠层配置究竟有什么不同?群体内部的光照分布情况究竟如何?抽穗后光合系统的机能是如何消退的?迄今尚未见报道。

针对以上问题, 本研究选用甬优系列超高产组合为研究对象, 选用长江下游地区具有代表性的杂交粳稻、常规粳稻和杂交籼稻品种为对照, 在扬州大学试验农场, 采用机插高产种植方式, 配套相应的高产栽培管理技术措施, 进行不同类型水稻品种抽穗期的冠层配置、光照分布及抽穗后的光合与抗衰老特性比较分析, 试图阐明甬优系列杂交稻的株型特征以及冠层结构和光合特性, 从而为籼粳杂交稻的高产栽培及高光效株型育种提供参考依据。

1 材料与方法

1.1 试验地点

扬州大学试验农场(32° 24′ N, 119° 26′ E), 年平均气温14.8℃, 年平均日照2140 h。土壤类型为沙壤土, 20 cm土壤层中含全氮0.13%、碱解氮87.3 mg kg^-1、速效磷32.5 mg kg^-1、速效钾88.5 mg kg^-1。

1.2 供试材料

选用4种类型20份具有代表性的高产水稻品种, 其中籼粳杂交稻(A)为甬优11、甬优12、甬优13、甬优15、甬优17, 杂交粳稻(B)为常优1号、常优2号、常优3号、甬优8号、甬优720, 常规粳稻(C)为扬粳4227、扬粳4038、南粳44、宁粳3号、镇稻11, 杂交籼稻(D)为II优084、新两优6380、丰两优香1号、扬两优6号、两优培九。各品种在江苏扬州均能正常抽穗结实, 2年(2012、2013)抽穗期和成熟期列于表1。

表1 供试水稻品种的抽穗期和成熟期 Table 1 Heading and maturity in different types of rice cultivars (month/day)

1.3 试验设计与栽培管理

2012— 2013年试验采取裂区设计, 品种类型为主区, 品种为裂区, 裂区面积15 m², 重复2次。小区间作埂并用塑料薄膜覆盖, 保证单独排灌。根据不同类型水稻高产特征要求, 应用精确定量栽培技术设计原理分别设置不同类型品种的高产栽培技术管理措施。采用塑盘旱育小苗, 5月20日播种, 6月5日移栽, 株行距为13.3 cm× 30.0 cm, 常规稻每穴4苗, 杂交稻每穴2苗。

总施氮量, 籼粳杂交稻、杂交粳稻和常规粳稻为300 kg hm^-2, 杂交籼稻为225 kg hm^-2, 均按为基蘖肥∶ 穗肥=5∶ 5, 基肥∶ 分蘖肥=6∶ 4, N∶ P₂O₅∶ K₂O=1.0∶ 0.5∶ 1.0。分蘖肥于移栽后1个叶龄一次性施入, 穗肥于倒四叶和倒二叶分2次施用, 磷肥全作基肥, 钾肥分基肥和拔节肥等量施用。在有效分蘖临界叶龄的前1个叶龄, 当茎蘖数达到预期穗数的80%时, 排水搁田, 坚持轻搁、多次搁原则; 拔节至成熟期实行湿润灌溉, 干干湿湿。其他栽培管理措施均按高产栽培要求实施。

1.4 测定项目与方法

1.4.1 株型指标于抽穗期, 从每小区选取生长基本一致的植株10穴, 测定叶长、叶宽、叶披垂度、叶基角、叶张开角、总叶面积(所有茎蘖的叶面积)、有效叶面积(有效茎蘖的叶面积)和高效叶面积(有效茎蘖顶三叶的叶面积)。

高效叶面积率(%) = 高效LAI/最大LAI× 100

有效叶面积率(%) = 有效LAI/最大LAI× 100 (其中: LAI为叶面积指数)

1.4.2 冠层叶面积分布采用上述同一样本, 10 cm间隔切片方法测定分层叶面积。自然状态下测定株高后, 每隔10 cm用自制装置固定植株的自然状态, 自上而下水平分割各层的绿叶、枯叶、鞘、茎、穗等5个部分。其中, 绿叶经扫描仪扫描后, 由Photoshop软件判读叶面积, 再根据群体密度, 计算分层叶面积指数。

1.4.3 冠层内光照强度分布于抽穗期的晴天11:30至12:30, 采用GL-1棍式照度计从冠层顶部向下每隔10 cm测定一次行间光照强度。为消除时间误差, 每次均采用往返观测法。然后将测定数据按层次求取平均值, 作为该处理特征值。

1.4.4 叶片光合特性相关指标于抽穗期、乳熟期、蜡熟期和成熟期, 随机取各品种剑叶10片混合测定光合色素、丙二醛含量(MDA)及过氧化氢酶(CAT)、超氧化物歧化酶(SOD)和过氧化物酶(POD)的活性。选择晴朗无风的上午, 从每品种随机选取10穴, 每穴1个主茎, 采用美国LI-COR公司LI-6400便携式光合测定仪测定剑叶的净光合速率, 叶室CO₂的浓度为350 μ mol mol^-1, 使用红蓝光源, 光量子通量密度(PFD)为1400 μ mol m^-2 s^-1, 温度为28~ 30℃。同时, 选取1个主茎, 采用FMS-2型便携脉冲调制式荧光仪测定叶绿素荧光参数, 光系统II (PSII)的实际光化学效率Ф _PSII = (F_m° - F_s)/F_m° 。

1.4.5 干物质量、叶面积和产量于成熟期每小区随机普查50穴, 计算单位面积有效穗数; 取5穴测定干物质、收获指数、每穗粒数、结实率和千粒重, 计算理论产量, 并实收计产。

1.5 数据计算与统计分析

采用Microsoft Excel 2003处理数据及制表, Origin 8.5绘图, DPS 8.0软件统计分析。

1.5.1 叶面积密度函数的拟合采用Logistic方程[方程(1)]对水稻群体向上累计叶面积指数, 随着相对高度的分布进行拟合, 求得方程参数a、b和c。为方便进行品种类型间比较, 群体叶面积归一化为100, 株高为1。

F(h) = (1)

A(h) = (2)

1.5.2 光照分布函数的拟合采用Richards方程[方程(3)]对冠层光照强度进行拟合分析。为方便进行品种类型间比较, 本文以相对光照强度进行拟合分析, 最大光强归一化为100, 株高为1。

T(h)= (3)

1.5.3 消光系数的测定

采用公式计算消光系数, 其中LAI为群体叶面积指数, I为测定位置的光照强度, I₀为冠层顶部的光照强度, ε 为群体消光系数。

2 结果与分析

2.1 主要冠层结构与光合特性性状的方差分析

方差分析表明(表2), 剑叶长度、剑叶叶基角、叶面积指数、净光合速率、生物学产量和稻谷产量等性状在品种类型间存在极显著差异, 但在年份及年份和类型互作上均无显著差异。说明年度间重复性较好, 本文主要以2012年的数据进行系统分析。

表2 不同类型水稻品种主要冠层结构与光合性状的方差分析 Table 2 Analysis of variance of F-value of main canopy structure and physiological traits in different types of rice cultivars

2.2 不同类型水稻冠层结构特征

2.2.1 叶片大小由表3可知, 上三叶的长度呈A> B> D> C趋势, 其中A倒一、倒二和倒三叶平均为61.19、67.26、64.55 cm, 较B分别高34.87%、31.88%、28.92%, 较C分别高86.50%、78.79%、83.28%, 较D分别高22.53、9.49%、15.60%; 上三叶的宽度呈A> D> B> C趋势, 其中A倒一、倒二和倒三叶平均为2.71、2.24、1.96 cm, 较B分别高18.15%、19.13%和8.29%, 较C分别高40.31%、36.35%和26.78%, 较D分别高14.26%、8.95%和8.29%。

2.2.2 受光姿态表3表明, 上三叶的叶基角均表现为D> A> B> C, 其中A倒一、倒二和倒三叶平均为11.05° 、16.72° 和22.02° , 较B高38.41%、31.98%和24.78%, 较C高130.18%、89.09%和47.49%, 较D低23.08%、20.46%和17.10%; 进一步分析披垂度, 上三叶均表现为D> B> C> A, 其中A倒一、倒二和倒三叶平均为1.29° 、1.87° 和2.76° , 较B分别低55.98%、76.79%和79.20%, 较C分别低46.48%、51.79%和67.01%, 较D分别低75.95%、82.73%和82.85%。

表3 不同类型水稻品种上三叶的配置 Table 3 Characteristics of top three leaves in different types of rice cultivars

2.2.3 不同类型水稻品种群体叶面积密度分布特征

采用Logistic方程对相对累计向上叶面积指数进行模拟, 并对其求导, 得到叶面积密度分布方程(表4), 拟合度较好, 均在0.98以上。从图1-a可以看出, 不同类型水稻冠层上部叶面积密度、冠层下部叶面积密度、最大叶面积密度及其出现的相对高度存在差异, 其中冠层上部的叶面积密度呈D> A> B> C趋势, 冠层下部的叶面积密度呈C> B> A> D趋势。根据方程可以求出的最大叶面积密度出现的相对高度(h_max)及最大叶面积密度(ρ _max)表现为D> A> B> C和A> D> B> C (表4)。

	Figure Option View Download New Window
	图1 不同类型水稻品种抽穗期群体相对叶面积密度和透光率分布特征A: 籼粳杂交稻; B: 杂交粳稻; C: 常规粳稻; D: 杂交籼稻。Fig. 1 The distribution of relative leaf area density and light transmission in different types of rice cultivarsA: indica-japonica hybrid rice; B: japonica hybrid rice; C: conventional japonica rice; D: indicahybrid rice.

表4 不同类型水稻品种抽穗期群体叶面积密度分布特征 Table 4 Distribution of leave area density in different types of rice cultivars

2.2.4 不同类型水稻品种群体内光照分布特征

采用Richards方程模拟冠层内相对光照强度(表5), 拟合度较好, 均在0.98以上。从图1-b可以看出, 不同类型水稻品种冠层相对光照强度存在差异, 冠层上部呈A> B> C> D趋势, 冠层下部呈B> A> C> D趋势。根据方程求得冠层的平均相对光照强度(I_mean)和消光系数(ε ), 分别表现为B> A> C> D和C> D> B> A (表5)。

表5 不同类型水稻品种群体光照分布特征 Table 5 Distribution of light distribution in different types of rice cultivars

2.3 不同类型水稻品种光合生产特征

2.3.1 群体叶面积指数、有效叶面积率及高效叶面积率从表6可以看出, 不同类型水稻品种抽穗期叶面积指数及其组成等存在显著差异, 且年份间趋势一致, 其中抽穗期群体叶面积指数呈D> A> B> C趋势, A平均为7.94, 较B、C分别高1.66%和3.57%, 较D低1.28%; 有效叶面积率呈A> B> C> D趋势, A平均为97.11%, 较B、C、D分别提高1.53%、3.22%和6.45%; 高效叶面积率趋势与群体叶面积指数一致, 较B、C分别高3.09%、5.37%, 较D低2.59%。

2.3.2 颖花/叶、实粒/叶不同类型水稻品种粒叶比存在显著差异(表6), 且年份间趋势一致。颖花/叶、实粒/叶均呈A> B> C> D趋势, 其中A颖花/叶平均为0.75, 较B、C、D分别高16.14%、29.15%和31.39%; A实粒/叶平均为0.63, 较B、C、D分别高13.04%、19.01%和34.19%。

表6 不同类型水稻抽穗期LAI组成及粒叶比 Table 6 LAI component and grain-leaf ratio in different types of rice cultivars

2.3.3 生物产量、经济产量与经济系数不同类型水稻品种生物产量、经济产量和经济系数存在显著差异(表7)。生物产量和经济产量均呈A> B> C> D趋势, A生物产量平均为24.69 t hm^-2, 较B、C、D分别高6.19%、12.65%和23.45%; A经济产量平均为12.15 t hm^-2, 较B、C、D分别高5.41%、11.48%和19.84%。收获指数类型间差异较小, 由高到低顺序为D、C、B、A。

表7 不同类型水稻品种产量与收获指数 Table 7 Yield and harvest index in different types of rice cultivars

2.3.4 生物产量、经济产量与经济系数不同类型水稻品种生物产量、经济产量和经济系数存在显著差异(表7)。生物产量和经济产量均呈A> B> C> D趋势, A生物产量平均为21.25 t hm^-2, 较B、C、D分别高6.89%、13.39%和22.83%; A经济产量平均为12.15 t hm^-2, 较B、C、D分别高5.41%、11.48%和19.84%。收获指数类型间差异较小, 由高到低顺序为D、C、B和A。

2.3.5 叶绿素含量、PSII实际光化学效率及净光合速率叶绿素含量、类胡萝卜素含量、PSII实际光化学效率及净光合速率均随灌浆进程呈逐渐下降趋势且存在品种间差异, 抽穗期和乳熟期类型间表现为D> C> B> A趋势, 蜡熟期和成熟期类型间均呈A> B> C> D趋势且存在品种间差异(图2)。

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	图2 不同类型水稻品种不同生育时期剑叶光合性状的变化A: 籼粳杂交稻; B: 杂交粳稻; C: 常规粳稻; D: 杂交籼稻。Fig. 2 Changes of photosynthetic traits in different types of rice cultivars during different growth periodsA: indica-japonica hybrid rice; B: japonica hybrid rice; C: conventional japonica rice; D: indicahybrid rice.

2.3.6 MDA含量与膜质过氧化酶活性不同类型水稻品种MDA含量和SOD、POD、CAT活性存在差异, 且年份间趋势一致(图3)。其中MDA含量随灌浆进程呈逐渐上升趋势, 抽穗期、乳熟期、蜡熟期和成熟期类型间表现为D> C> B> A趋势。SOD、POD和CAT的活性随灌浆进程呈逐渐下降趋势, 类型间趋势一致, 为A> B> C> D。

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	图3 不同类型水稻品种不同生育时期剑叶膜质过氧化的变化A: 籼粳杂交稻; B: 杂交粳稻; C: 常规粳稻; D: 杂交籼稻。Fig. 3 Changes of membrane lipid peroxidation in different types of rice cultivars during different growth periodsA: indica-japonica hybrid rice; B: japonica hybrid rice; C: conventional japonica rice; D: indicahybrid rice.

3 讨论

3.1 关于甬优系列籼粳杂交稻的冠层结构特征

光能是作物转化利用的对象, 群体光截获量和光能利用率是决定作物产量的重要因素^[23]。因此, 通过改善作物冠层结构, 使更多的光能到达冠层中下部, 进而提高群体光能利用率, 对提高单位面积产量具有重要意义。已有研究表明, 水稻群体光照分布受到栽培条件的显著影响^{[24, 25]}, 同时在基因型或品种类型间存在很大差异^{[26, 27, 28]}。陈温福等^[26]研究表明, 水稻穗型对群体冠层内光分布有较大影响, 直立穗型群体消光系数低, 群体内部光分布较均匀, 更利于群体光能利用和物质生产。吕川根等^[27]研究认为, 在相同高产栽培条件下, 紧凑型品种两优培九基部透光率明显高于松散型品种汕优63。张运波等^[28]研究表明, 太阳辐射利用率、冠层截获的有效辐射量呈超级杂交稻、常规稻> 普通杂交稻趋势。马荣荣等^[21]研究表明, 超高产栽培条件下, 籼粳杂交稻甬优6号群体内部透光率明显优于杂交籼稻汕优63。本研究结果表明, 不同类型水稻品种冠层内光照分布存在差异, 在冠层相对高度0~0.7范围内, 类型间光照呈杂交粳稻(甬优8号)> 籼粳杂交稻(甬优12) > 常规粳稻(镇稻11)> 杂交籼稻(扬两优6号)趋势, 在0.7~1.0范围内, 类型间光照呈甬优12 > 甬优8号> 镇稻11> 扬两优6号趋势, 冠层的平均光照和消光系数表现为甬优8号> 甬优12> 镇稻11> 扬两优6号和镇稻11> 扬两优6号> 甬优8号> 甬优12, 说明甬优系列籼粳杂交稻受光条件好, 光能截获率高, 为其光合系统持续产出及形成高生物学产量提供了生态基础, 这与马荣荣等^[21]在甬优6号上得出的结论基本一致。

水稻群体光分布与其冠层结构配置密切相关, 群体大小、叶片姿态及群体叶面积分布均影响光照在群体内的分布^[29]。根据本研究结合前期研究结果, 我们认为甬优系列籼粳杂交稻冠层结构优势的形成有以下3点。(1)在大穗少蘖品种特性以及合理基本苗起点基础上, 通过全程精确栽培调控使得群体发展平稳, 最终群体质量高(高峰苗低于杂交粳稻、常规粳稻和杂交籼稻; 成穗率高于杂交籼稻, 略低于杂交粳稻、常规粳稻; 有效叶面积率高于杂交粳稻、常规粳稻和杂交籼稻; 高效叶面积率低于杂交籼稻, 高于杂交粳稻和常规粳稻); (2)虽然亚种间强大的杂种优势使得个体繁茂性强(高效叶长度、宽度均高于杂交粳稻、常规粳稻和杂交籼稻), 但受卷叶基因的影响使得肥大的高效叶反而能保持直立(叶披垂度几乎为零, 显著低于杂交粳稻、常规粳稻和杂交籼稻), 塑造了良好的受光姿态; (3)较小的群体规模与较大的个体繁茂性平衡作用使得中期形成适宜的群体叶面积指数(抽穗期LAI表现为杂交籼稻> 籼粳杂交稻> 杂交粳稻> 常规粳稻), 同时有效率高, 平衡了足源与遮光的矛盾, 在此基础上, 通过优化群体相对叶面积密度的分布(冠层上部叶面积密度表现为杂交籼稻> 籼粳杂交稻> 杂交粳稻> 常规粳稻, 冠层下部叶面积密度表现为常规粳稻> 杂交粳稻> 籼粳杂交稻> 杂交籼稻), 使得群体透光性得到提高。综上所述, 甬优系列杂交稻具有显著的光合生态优势, 良好的冠层结构是其基础。

3.2 关于甬优系列籼粳杂交稻的光合特征

籽粒产量是生物学产量和收获指数的乘积, 提高产量是通过增加生物学产量还是通过提高收获指数, 不同学者提出了不同的观点^{[30, 31]}。本研究观察到, 与其他3种类型品种相比, 甬优系列籼粳杂交稻经济系数低于杂交籼稻, 与杂交粳稻和常规粳稻无显著差异, 说明其经济系数相对稳定, 但无论与杂交粳稻、常规粳稻还是杂交籼稻相比, 其生物学产量和籽粒产量均有大幅提高, 这与许德海等在甬优6号上的研究结果基本一致^[22], 表明增加生物学产量, 稳定收获指数是甬优系列籼粳杂交稻较其他类型品种增产的主要原因, 也可能是进一步提高其籽粒产量的可靠途径。籽粒产量可分解为叶面积指数与粒叶比两个构成因素。理论上讲, 提高叶面积指数和提高粒叶比中任何一个均可以提高产量, 但凌启鸿、杨建昌等^{[31, 32]}研究认为, 在适宜的叶面积指数基础上, 大幅提高粒叶比是提高水稻产量的有效途径。本研究结果表明, 甬优系列籼粳杂交稻抽穗期的群体叶面积指数低于杂交籼稻, 但高于杂交粳稻和常规粳稻, 说明其群体叶面积指数适中; 粒叶比表现为甬优系列籼粳杂交稻明显高于其他3类品种, 可见甬优系列籼粳杂交稻主要通过大幅提高粒叶比来提高产量, 这一结果符合凌启鸿等^{[32, 33]}的观点。

经济产量与生物学产量的协同优势以及适宜叶面积指数与高粒叶比的形成共同说明了甬优系列籼粳杂交稻在抽穗至成熟阶段存在着显著的光合优势。众所周知, 水稻叶片光合能力体现在光合面积、单位面积的光合生产能力(光合速率)和光合能力的持续等3个方面^[34]。根据本研究结合前期研究可以确定甬优系列籼粳杂交稻在光合面积和光合时间上均具有明显优势, 但与其他3种类型品种相比, 其单位面积光合生产能力是否也存在优势?随灌浆进程, 光合系统的机能是如何消退的?迄今尚未见报道。本研究在品种灌浆的关键生理时期, 对不同类型水稻的光合色素、叶绿素荧光动力学参数、净光合速率等光合指标系统测定和分析发现, 在灌浆前期(抽穗期至乳熟期), 甬优系列籼粳杂交稻叶绿素含量、类胡萝卜素含量、PSII的实际光化学效率、净光合速率均低于杂交粳稻、常规粳稻和杂交籼稻, 但灌浆后期(蜡熟期至成熟期)相关光合指标呈相反趋势, 可见, 甬优系列杂交稻的光合强度优势集中表现在灌浆后期。进一步测定MDA含量和膜质抗氧化酶系统发现, 甬优系列杂交稻的MDA含量在整个灌浆期均低于其他3类品种, SOD、POD和CAT酶活性均高于杂交粳稻、常规粳稻和杂交籼稻, 说明甬优系列杂交稻抽穗后功能叶片衰老缓慢, 灌浆后期仍能维持较好的光合机能, 这可能是其光合产物更多产出、籽粒充实良好以及产量优势形成的重要生理原因之一。

4 结论

机插高产栽培条件下, 甬优系列籼粳杂交稻(A)、杂交粳稻(B)、常规粳稻(C)和杂交籼稻(D)在叶片受光姿态、群体冠层光分布、光合特性等方面存在明显差异, 具体表现为: (1)上3张叶片的长度、宽度表现为A> B> D> C, 叶基角表现为D> A> B> C, 披垂度表现为D> B> C> A。冠层上部叶面积密度表现为D> A> B> C, 冠层下部叶面积密度表现为C> B> A> D, 最大叶面积密度表现为A> D> B> C, 最大叶面积密度出现的相对高度表现为D> A> B> C。冠层上部相对光照表现为A> B> C> D, 冠层下部相对光照表现为B> A> C> D, 冠层平均相对光照表现为B> A> C> D, 冠层消光系数表现为C> D> B> A。(2)抽穗期群体叶面积指数、高效叶面积率均呈D> A> B> C趋势, 有效叶面积率呈A> B> C> D趋势; 颖花/叶和实粒/叶均表现为A> B> C> D。经济产量、生物产量均表现为A> B> C> D, 经济系数呈D> C> B> A趋势; 蜡熟期和成熟期剑叶的叶绿素含量、类胡萝卜素含量、PSII的光化学效率及净光合速率呈A> B> C> D趋势; 抽穗至成熟期剑叶的MDA含量呈D> C> B> A趋势, SOD、POD、CAT活性呈A> B> C> D趋势。说明, 与其他3种类型水稻相比, 甬优系列籼粳杂交稻的冠层结构与光合特性具有显著优势, 这是甬优系列杂交稻产量潜力正常发挥的生态生理基础, 也是进一步提高亚种间杂交稻群体生产力的重要途径。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

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[16]	熊洁, 陈功磊, 王绍华, 丁艳锋. 江苏省不同年代典型粳稻品种的产量及株型差异. 南京农业大学学报, 2011, 34(5): 1-6 Xiong J, Chen G L, Wang S H, Ding Y F. The difference in grain yield and plant type among typical japonica varieties in different years in Jiangsu Province. J Nanjing Agric Univ, 2011, 34(5): 1-6 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.916]
[17]	张庆, 殷春渊, 张洪程, 魏海燕, 马群, 杭杰, 李敏, 李国业. 水稻氮高产高效与低产低效两类品种株型特征差异研究. 作物学报, 2010, 36: 1011-1021 Zhang Q, Yin C Y, Zhang H C, Wei H Y, Ma Q H J, Li M, Li G Y. Differences of plant-type characteristics between rice cultivars with high and low levels in yield and nitrogen use efficiency. Acta Agron Sin, 2010, 36: 1011-1021 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.667]
[18]	李敏, 张洪程, 杨雄, 葛梦婕, 马群, 魏海燕, 戴其根, 霍中洋, 许轲. 不同氮利用效率基因型水稻茎秆特性比较. 作物学报, 2012, 38: 1277-1285 Li M, Zhang H C, Yang X, Ge M J, Ma Q, Wei H Y, Dai Q G, Huo Z Y, Xu K. Comparison of culm characteristics with different nitrogen use efficiencies for rice cultivars. Acta Agron Sin, 2012, 38: 1277-1285 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
[19]	袁隆平, 武小金, 颜应成, 罗孝和. 水稻广谱广亲和系的选育策略. 中国农业科学, 1997, 30(4): 1-8 Yuan L P, Wu X J, Yan Y C, Luo X H. A strategy for developing wide spectrum compatibility rice line. Sci Agric Sin, 1997, 30(4): 1-8 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]
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[21]	马荣荣, 许德海, 王晓燕, 禹盛苗, 金千瑜, 欧阳由男, 朱练峰. 籼粳亚种间杂交稻甬优6号超高产株形特征与竞争优势分析. 中国水稻科学, 2007, 21: 282-286 Ma R R, Xu D H, Wang X Y, Yu S M, Jin Q Y, Yu S M, Ou-Yang Y N, Zhu L F. Competition advantage on plant morphology of Yongyou 6: an yield indica-japonica super hybrid rice. Chin J Rice Sci, 2007, 21: 282-286 (in Chinese with English abstract) [本文引用:3] [CJCR: 1.494]
[22]	许德海, 王晓燕, 马荣荣, 禹盛苗, 朱练峰, 欧阳由男, 金千瑜. 重穗型籼粳杂交稻甬优6号超高产生理特性. 中国农业科学, 2007, 21: 281-286 Xu D H, Wang X Y, Ma R R, Yu S M, Zhu L F, Ou-Yang Y N, Jin Q Y. Analysis on physiological properties of the heavy panicle type of indica-japonica inter-subspecific hybrid rice Yongyou6. Sci Agric Sin, 2007, 21: 281-286 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.889]
[23]	Campbell C S, Heilman J L, Mclnnes K J, Wilson L T, Medley J C, Wu G W, Cobos D R. Seasonal variation in radiation use efficiency of irrigated rice. Agric For Meteorol, 2001, 110: 45-54 [本文引用:1]
[24]	Zhang L, van der Werf W, Bastiaans L, Zhang S, Li B, Spiertz J H J. Light interception and utilization in relay intercrops of wheat and cotton. Field Crops Res, 2008, 107: 29-42 [本文引用:1] [JCR: 2.474]
[25]	陈雨海, 余松烈, 于振文. 小麦生长后期群体光截获量及其分布与产量的关系. 作物学报, 2003, 29: 730-734 Chen Y H, Yu S L, Yu Z W. Relationship between amount or distribution of PAR interception and grain output of wheat communities. Acta Agron Sin, 2003, 29: 730-734 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
[26]	Chen W F, Xu Z J, Zhang L B, Zhou H F, Yang S R. Effects of different panicle type on canopy properties light distribution and dry matter production of rice population. Acta Agron Sin, 1995, 21: 83-89 [本文引用:2] [CJCR: 1.667]
[27]	吕川根, 邹江石, 胡凝, 姚克敏. 水稻叶片形态对冠层特性和光合有效辐射传输的影响. 江苏农业科学, 2007, 23: 501-508 Lü C G, Zou J S, Hu N, Yao K M. Effects of morphological factors on canopy structure and transmission of photosynthetically active radiation in rice. Jiangsu J Agric Sci, 2007, 23: 501-508 (in Chinese with English abstract) [本文引用:2] [CJCR: 0.895]
[28]	李迪秦, 唐启源, 秦建权, 张运波, 郑华英, 杨胜海, 陈立军, 邹应斌. 施氮与氮管理模式对超级稻产量和辐射利用率的影响. 核农学报, 2010, 24: 809-814 Li D Q, Tang Q Y, Qin J Q, Zhang Y B, Zheng H Y, Yang S H, Chen L J, Zou Y B. Effects of nitrogen management patterns on grain yield, solar radiant use efficiency in super hybrid rice. J Nucl Agric Sci, 2010, 24: 809-814 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.237]
[29]	李绪孟, 王小卉, 郑华斌, 黄璜. 水稻群体冠层结构及其光分布研究进展. 作物研究, 2012, 26: 591-595 Li X M, Wang X H, Zheng H B, Huang H. Research progress on canopy structure of and its light distribution in rice. Crops Res, 2012, 26: 591-595 (in Chinese with English abstract) [本文引用:1] [JCR: 2.474]
[30]	Peng S, Gassman K G, Virmani S S, Sheehy J, Khush G S. Yield potential trends of tropical rice since release of IR8 and the challenge of increasing rice yield potential. Crop Sci, 1999, 39: 1552-1559 [本文引用:1] [JCR: 1.513]
[31]	Yang J, Peng S, Zhang Z, Wang Z, Visperas R M, Zhu Q. Grain yield and dry matter yields and partitioning of assimilates in japonica/indica hybrid rice. Crop Sci, 2002, 42: 766-772 [本文引用:2] [JCR: 1.513]
[32]	凌启鸿, 杨建昌. 水稻群体“粒叶比”与高产栽培途径的研究. 中国农业科学, 1986, 19(3): 1-8 Ling C H, Yang J C. Studies on “grain-leaf ratio” of population and cultural approaches of high yield in rice plants. Sci Agric Sin, 1986, 19(3): 1-8 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.889]
[33]	杨建昌, 杜永, 吴长付, 刘立军, 王志琴, 朱庆森. 超高产粳型水稻生长发育特性的研究. 中国农业科学, 2006, 39: 1336-1345 Yang J C, Du Y, Wu C F, Liu L J, Wang Z Q, Zhu Q S. Growth and development characteristics of super-high-yielding mid-season japonica rice. Sci Agric Sin, 2006, 39: 1336-1345 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]
[34]	李敏, 张洪程, 杨雄, 葛梦婕, 魏海燕, 戴其根, 霍中洋, 许轲. 高产氮高效型粳稻品种的叶片光合及衰老特性研究. 中国水稻科学, 2013, 27: 168-176 Li M, Zhang H C, Yang X, Ge M J, Wei H Y, Dai Q G, Huo Z Y, Xu K. Leaf photosynthesis and senescence characteristics of japonica rice cultivars with high yield and high N-efficiency. Chin J Rice Sci, 2013, 27: 168-176 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.494]

2000

0.0

... 回顾我国水稻单产的两次重大飞跃(20世纪50年代末的矮化育种增产及70年代中期的三系杂交育种增产), 归结其根本的原因是水稻株型得到了改良^[1,2,3] ...

2006

0.0

1.667

邓启云, 袁隆平, 蔡义东, 刘建丰, 赵炳然, 陈立云. 超级杂交稻模式株型的光合优势. 作物学报, 2006, 32: 1287-1293

Deng Q

, Yuan L

, Cai Y

, Liu J

, Zhao B

, Chen L

Photosynthetic advantages of model plant-type in super hybrid rice.

Acta Agron Sin, 2006, 32: 1287-1293 (in Chinese with English abstract)

Looking back at the history of rice breeding, two breakthroughs resulted largely from identification and utilization of dwarfing genes in 1960s and the wild abortive cytoplasmic male sterility (CMS) in 1970s had significantly raised rice yield potential. But the yield potential of both inbred and hy

1 China National Hybrid Rice Research & Development Center, Changsha 410125, Hunan; 2 Hunan Agricultural University, Changsha 410128, Hunan, China

以三系杂交稻汕优63为对照，比较研究表明，具有超级杂交稻模式株型的培矮64S/E32有以下生理优势。（1）叶面积指数前期较小，中期稳健，后期衰减缓慢、维持较高水平；（2）抽穗后剑叶和倒2叶的比叶重显著或极显著大于对照；（3）后期上3叶叶绿素含量衰减缓慢，变化平稳；（4）上3叶单位叶面积气孔数目比对照多19.2%，达极显

2013

0.0

0.451

2011

0.0

1.494

付景, 杨建昌. 超级稻高产栽培生理研究进展. 中国水稻科学, 2011, 25: 343-348

, Yang J

Research advances in physiology of super rice under high-yielding cultivation.

Chin J Rice Sci, 2011, 25: 343-348 (in Chinese with English abstract)

China launched a program for breeding of super rice or super hybrid rice by combining intersubspecific heterosis with ideal plant type in 1996. Today, about 80 super rice varieties have been released and some of them have achieved grain yields of 12-21 tons per hectare in field experiments. Compared with conventional cultivars, main reasons for high yield of super rice can be summarized as follows: more spikelets per panicle and larger sink size (number of spikelets per square meter); larger leaf area index, longer duration of green leaf area, higher photosynthetic rate and higher lodging resistance of culms; more matter accumulation before the heading stage, more remobilization of prestored carbohydrates from stems and leaves to grains during the grain filling period; and larger root system and greater root activity. However, super rice also has two major problems in production: poor grainfilling of the laterflowering inferior spikelets (in contrast to earlierflowering superior spikelets), and low and unstable seedsetting rate. The research advances in cultivation physiology of super rice in respects of biological features, formation of yield components, population quality and physiological mechanisms were reviewed, and suggestion on further studies on super rice cultivation physiology were put forward.

Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China

我国于1996年启动了理想株型与亚种间杂种优势相结合为技术路线的超级稻育种计划，目前已育成约80个超级稻品种。这些品种产量高，兼顾品质与抗性，在试验示范区或特定气候条件下产量可达到12～21 t/hm2，展示了超级稻的巨大增产潜力。对于超级稻品种高产的原因，可概括为以下几个方面：与常规高产品种相比，超级稻品种每穗颖花数多、库容量大(即单位面积颖花数多)；叶面积指数大、绿叶面积持续期长、光合速率高、茎秆抗倒性强；抽穗前干物质累积量高, 结实期茎叶中碳水化合物转运到籽粒中的量大; 根量大、根系活性强。但是，超级稻品种在生产上也存在一些问题，突出地表现在两个方面：一是强势粒(着生在穗子中上部、开花较早的籽粒)、弱势粒(着生在穗子基部、开花较迟的籽粒)充实整齐性差；二是结实率低且不稳定。综述了超级稻的生物特性、产量结构、群体质量特征以及相关生理机理方面的研究进展，提出深入开展超级稻栽培生理研究的建议。

... 众多水稻学者一致认为, 优良的植株形态是实现水稻超高产第三次突破的骨架^[4,5,6] ...

2008

2009

1968

1.643

0.0

Donald C

The breeding of crop ideotypes.

Euphytica, 1968, 17: 385-403

1. Waite Agricultural Research Institute, The University of Adelaide, South Australia

Most plant breeding is based on “defect elimination” or “selection for yield”. A valuable additional approach is available through the breeding of crop ideotypes, plants with model characteristics known to influence photosynthesis, growth and (in cereals) grain production. Some instances of the successful use of model characters of this kind are quoted.

... 前人围绕水稻高产、超高产的理想株型及其机能开展了大量研究, 并根据地域和品种特性等提出了诸多理想株型模式^{[7,8,9,10,11,12]}, 也从水稻亚种属性^[13,14]、品种演进^[15,16]以及高产高效协同型品种^[17,18]等角度对水稻的株型特征进行了深入研究, 如张洪程等^[13]研究认为与籼稻相比, 粳稻生育中期能形成适宜的叶面积指数, 较紧凑的株型以及较挺拔的叶片, 群体内部相对通风透光, 并且抽穗期群体干物质量积累适中, 较高的粒叶比保证最终库源关系较协调 ...

2005

0.0

0.3579

1996

0.0

1.494

杨守仁, 张龙步, 陈温福, 徐正进, 王进民. 水稻超高产育种理论和方法, 中国水稻科学, 1996, 10: 115-120

Yang S

, Zhang L

, Chen W

, Xu Z

, Wang J

Theories and methods of rice breeding for maximum yield.

Chin J Rice Sci, 1996, 10: 115-120 (in Chinese with English abstract)

Rice breeding for maximum yield is a hot topic today in the world and a tremendous challenge as well. China's population is now approaching 1. 2 billion people. It is a country where rice constitutes a lion's share in grain production. The higher and higher the yield we seek, the more and more difficult the task will be. It is a breeding for maximum yield. Since there is no ready theory and method for us to follow, we have to grope along through trial and error. In this paper, the author attempts to make a comprehensive analysis of and systemize the valuable experience acquired by so many people through decades of painstaking labor, in order to further refine the theories and methods for combining ideal plant morphology with the utilization of vigor, as well as for optimized conbination of traits and criteria for selecting hybrid progeny put forward in the past, for reference and trial application by colleagues at home and abroad.

水稻超高产育种是当前世界性的热门课题，也是难题。现在我国人口已近十二亿，水稻是我国粮食生产的大头。高产更高产，难度越来越大。它的超高产育种，既无现成的理论和方法可循，就只能摸索前进。本文试图将几十年来多少人的汗水心血所凝成的宝贵经验，全面加以分析整理，藉以进一步完善过去所提出的理想株形与优势利用相结合的大方向，优化性状组配以及杂交后代选择标准等理论和方法，以供国内外同行参考试用。

1997

0.0

0.605

1997

0.0

1.889

周开达, 汪旭东, 李仕贵, 李平, 黎汉云, 黄国寿, 刘太清, 沈茂松. 亚种间重穗型杂交稻研究. 中国农业科学, 1997, 30(5): 91-93

Zhou K

, Wang X

, Li S

, Li

, Li H

, Huang G

, Liu T

, Shen M

The study on heavy panicle type of inter-subspecific hybrid rice (Oryza sativa L. ).

Sci Agric Sin, 1997, 30(5): 91-93 (in Chinese with English abstract)

四川农业大学水稻研究所，温江 611130 重庆市作物研究所

1994

0.0

0.6672

2013

0.0

1.889

张洪程, 张军, 龚金龙, 常勇, 李敏, 高辉, 戴其根, 霍中洋, 许轲, 魏海燕. “籼改粳”的生产优势及其形成机理. 中国农业科学, 2013, 46: 686-704

Zhang H

, Zhang

, Gong J

, Chang

, Li

, Gao

, Dai Q

, Huo Z

, Xu

, Wei H

The productive advantages and formation mechanisms of “indica rice to japonica rice”.

Sci Agric Sin, 2013, 46: 686-704 (in Chinese with English abstract)

【Objective】 The objective of this study was to reveal the productive advantages of “indica rice to japonica rice” and summarize its physiological ecological mechanisms systematically from the viewpoint of cultivation. 【Method】 A field experiment was conducted with representative japonica rice varieties (Wuyunjing24, Xudao3, Lianjing7, Xuyou733, etc.) and indica rice varieties (Liangyoupeijiu, Ⅱyou084, Xinliangyou6380, Yangliangyou6, etc.) as materials in rice-wheat cropping areas (Yangzhou, Xinghua, Donghai of Jiangsu) in the lower reaches of the Yangtze River in 2010-2011. Difference of growth security, grain yield, quality, utilization of temperature and solar radiation, dynamics of population growth, plant type and lodging characteristics between japonica rice and indica rice were studied systematically. 【Result】 Under the simplified cultivation conditions of mechanical and swing transplanting, the growth security of japonica rice was better than indica rice, for japonica rice possessed longer milky stage and the resistance to low temperature at the later period compared to indica rice. The average yields of japonica rice in both years were, 10.90 and 10.87 t•hm-2 in Yangzhou, 10.93 and 10.65 t•hm-2 in Xinghua, 10.49 and 10.41 t•hm-2 in Donghai, higher than indica rice significantly. Grain filling characteristics of japonica rice exceeded indica rice, which was the key point of harvesting high yield of japonica rice, with the roughly same level of population spikelets or slightly less. Milling quality and cooking and eating quality of japonica rice were better than indica rice significantly, while the appearance and nutritional quality followed an opposite tendency accordingly. The physiological ecological characteristics of productive advantages of “indica rice to japonica rice” were as follows. Firstly, growth duration of japonica rice was longer than indica rice significantly and japonica rice was more adapted to cool weather at the later period of grain filling, increasing utilization of temperature and solar radiation and ensuring japonica rice maturing safely. Secondly, strong photosynthetic capacity of japonica rice at the later period increased dry matter weight and total filling. Thirdly, japonica rice improved the resistance to cold and premature senescence, and lodging due to strong root, stem and sheath, guaranteeing the enrichment and support of large sink. 【Conclusion】 Under the mechanizing simplified cultivation conditions, japonica rice, not only matured safely with strong growth advantages at the later period, but also achieved high grain yield, quality, efficiency and comprehensive productivity because of making full use of temperature and solar radiation in comparison to indica rice.

College of Agriculture, Yangzhou University/Innovation Center of Rice Cultivation Technology in the Yangtze Valley, Ministry of Agriculture/Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou 225009, Jiangsu

【目的】旨在阐明“籼改粳”的生产优势，并从栽培角度系统研究其优势形成的生理生态机理。【方法】2010—2011年，在长江下游稻麦两熟制地区（江苏扬州、兴化、东海），以当地代表性的粳稻品种（武运粳24号、徐稻3号、连粳7号、徐优733等）和籼稻品种（两优培九、Ⅱ优084、新两优6380、扬两优6号等）为材料，系统比较研究籼粳稻的生育安全性、产量、品质、温光资源利用、群体生长发育动态、株型和倒伏性状等方面的差异。【结果】在机插和摆栽两种轻简栽培条件下，粳稻生育安全性优于籼稻，后期耐低温，灌浆持续时间长。两年粳稻的平均产量，扬州分别为10.90和10.87 t•hm-2，兴化分别为10.93和10.65 t•hm-2，东海分别为10.49和10.41 t•hm-2，显著或极显著高于籼稻。粳稻群体颖花量与籼稻相当或略少于籼稻，但籽粒的充实性状优于籼稻，这也是粳稻获得较高产量的关键。粳稻加工品质和蒸煮食味品质显著或极显著优于籼稻，外观品质和营养品质稍逊于籼稻。“籼改粳”生产优势形成的生理生态特征为：粳稻全生育期较籼稻明显延长，灌浆后期粳稻更能适应温凉天气，增加水稻对温光资源的利用，使得粳稻能够安全成熟；粳稻后期具有较高的光合生产能力，能够增加群体光合物质积累量，提高群体库容总充实量；同时，粳稻后期能够适应低温天气而不早衰，维持强壮根系和较高的茎鞘强度，增强群体抗倒伏能力，保证较大库容的安全充实与支撑。【结论】在机械化轻简化栽培条件下，粳稻更能充分利用温光资源，生育后期仍保持强劲生长优势，不仅能够安全成熟，而且产量高、品质优、效益好，综合生产力高。

2014

0.0

1.667

龚金龙, 邢志鹏, 胡雅杰, 张洪程, 戴其根, 霍中洋, 许轲, 魏海燕, 高辉. 籼、粳超级稻光合物质生产与转运特征的差异. 作物学报, 2014, 40: 497-510

Gong J

, Xing Z

, Hu Y

, Zhang H

, Dai Q

, Huo Z

, Xu

, Wei H

, Gao

Difference of characteristics of photosynthesis, matter production and translocation between indica and japonica super rice.

Acta Agron Sin, 2014, 40: 497-510 (in Chinese with English abstract)

This study was to reveal the difference of characteristics of dry matter accumulation and photosynthetic production between indica and japonica super rice. Field experiments were conducted by using five representative super hybrid indica combinations and five conventional japonica super rice varieties planted on a large scale in Jiangsu region with rice-wheat double cropping rotation. Dry matter accumulation, distribution and translocation, leaf area, leaf area duration (LAD), crop growth rate (CGR), net assimilation rate (NAR), seedling quality and leaf type were analyzed systematically. Results showed that, dry matter accumulation, LAD, CGR and NAR at the early growth stage (from transplanting to jointing), leaf length, leaf base angle, angle between stem and leaf and drooping angle in top three leaves, leaf area decreasing per day after heading and harvest index of japonica rice were less than those of indica rice significantly, while dry matter accumulation, LAD, CGR and NAR at the middle and later growth stages (from jointing to maturity), ratio of leaf area of productive tillers, ratio of leaf area of top three leaves, grain-leaf ratio (spikelets per cm 2 leaf area, filled grains per cm 2 leaf area and grain weight per cm 2 leaf area), maximum leaf area index, total filling, grain yield, biological yield, maximum output and apparent output and their ratio were larger than those of indica rice significantly. Although dry matter weight per stem of japonica rice at the main growth stages was lower than that of indica rice, the superiority of population quantity suggested that japonica rice would possess higher dry matter accumulation and leaf area. With the advancement of growth and development, the superiority of photosynthesis and matter production of japonica rice was increasing, with a higher dry matter accumulation more than that of indica rice 25 d after heading approximately. The growth potential of japonica rice remained strong at the late period of grain-filling (from milky stage to maturity), while stored matter kept in the stem and sheath exported appropriately at the early period of grain-filling (from heading to milky stage), ensuring the security of efficient photosynthetic layer and the enrichment of high accumulation yield sink. Therefore, balanced formation of high biological yield, stable-increasing-slow-decreasing tendency of leaf area, and the large amounts of high efficient photosynthate production from jointing to maturity are the important characteristics and causes for the efficient and sustainable output of photosynthetic systems, more grain-filling and high yield formation in japonica rice.

为阐明籼、粳超级稻干物质积累及光合生产特征的差异，以江苏地区大面积推广种植的 5 个超级杂交籼稻组合和 5 个常规粳型超级稻品种为试验材料，对稻麦两熟制条件下籼、粳超级稻干物质积累、分配、运转及叶面积、光合势、群体生长率、净同化率、秧苗素质、叶型等方面进行了系统的比较研究。结果表明，粳稻生育前期 ( 移栽至拔节期 ) 干物质积累量、光合势、群体生长率、净同化率及上三叶叶长、叶基角、叶开角、披垂度和叶面积衰减率、收获指数均小于籼稻，而生育中后期 ( 拔节至成熟期 ) 干物质积累量、光合势、群体生长率、净同化率及有效叶面积率、高效叶面积率、粒叶比 ( 颖花 / 叶、实粒 / 叶、粒重 / 叶 ) 、最大叶面积指数、总充实量、实收产量、生物产量、茎鞘最大输出量和表观输出量及比率均大于籼稻，差异显著或极显著。虽然粳稻主要生育期单茎干物重均不及籼稻，但群体数量优势保证粳稻具有较高的群体干物质积累量和叶面积，且随着生长发育的持续，群体光合物质生产优势不断加大，群体干物质积累量于抽穗后 25 d 前后超过籼稻。粳稻灌浆后期 ( 乳熟至成熟期 ) 仍保持强劲生长优势，而灌浆初期 ( 抽穗至乳熟期 ) 茎鞘贮存物质合理输出，有效保障了高效光合层的安全支撑及高积累产量库的流畅充实。高生物学产量的稳定形成和叶面积 “ 稳升缓降 ” 态势以及拔节至成熟期较强的高效光合物质生产，是粳稻光合系统高效持续产出、灌浆充实多及高产形成的重要特征和原因。

2006

0.0

1.667

杨建昌, 王朋, 刘立军, 王志琴, 朱庆森. 中籼水稻品种产量与株型演进特征研究. 作物学报, 2006, 32: 949-955

Yang J

, Wang

, Liu L

, Wang Z

, Zhu Q

Evolution characteristics of grain yield and plant type for mid-season indica rice cultivars.

Acta Agron Sin, 2006, 32: 949-955 (in Chinese with English abstract)

Rice is a most important food crop in China and its grain yield per hectare has been over 6 ton in recent years, which is 65% higher than that of the world, on average. Improvement in rice cultivars plays an important role in the increase of the grain yield. However, little is known about physiological and agronomical features responsible for the improvement. In this study, 16 typical mid-season indica rice cultivars (including hybrid combinations) applied in the production in Jiangsu Province during the last 60 years were used, and classified into six types of early tall (ET), improved tall (IT), dwarf cultivars (DC), semi-dwarf cultivars (SDC), semi-dwarf hybrid (SDH), and super rice (SR), according to their application times, plant types and genotypes. All the tested cultivars were grown in the field and the evolution characteristics of grain yield and its components, source-sink relationships and plant types were investigated. Results showed that grain yield was progressively increased with the evolution of the cultivars. Increase in grain yield was attributed mainly to the increase in total number of spikelets, which resulted mainly from large panicles. DC had the greatest, whereas SR had the least, percentage of filled grains. There was no significant difference in 1000-grain weight among the six types of cultivars. From ET to DC, plant height was reduced and biomass and harvest index were simultaneously increased, especially the harvest index increased more. From DC to SR, plant height was a little increased, and biomass dramatically increased, especially after heading, and harvest index a little changed. Improvement of rice cultivars significantly reduced the leaf angle of the top three leaves on stems and increased leaf area at heading. Increase in spikelets, however, was more than that in leaf area, which resulted in the increase of grain-leaf ratio (the ratio of total spikelet number to leaf area at heading). With the improvement of rice cultivars, root dry weight and the ratio of root to shoot were increased, but the amount of root bleeding sap was reduced. The speed of reduction in the concentration of zeatin and zeatin riboside in root bleeding sap and leaves from heading to maturity was SR>SDH>SDC. The results suggest that enlargement of sink resulted from large panicles, increase in biomass especially from heading to maturity and improvement of plant type contribute to the increase of grain yield for modern rice cultivars. Low root activity may account for the low percentage of filled grains for SR. Approaches to further increase in grain yield of rice were discussed based on the evolution characteristics of the cultivars.

Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, Jiangsu, China

以江苏省近60年生产上应用的16个代表性中籼品种(含杂交稻组合)为材料，依品种应用时期结合株型和基因型，将供试品种分为早期高秆（ET）、改良高秆（IT）、矮秆（DC）、半矮秆常规品种（SDC）、半矮秆杂交稻（SDH）和超级稻（SR）6个类型，研究其产量、源库关系和株型变化特性。结果表明，随着品种改良，籽粒产量逐步提高。这主要是由于总颖花量的增加，其关键在于每穗粒数的增多；结实率以DC最高，SR最低，千粒重变化不大。由ET到DC，植株高度降低，生物产量与收获指数同步提高，收获指数的提高更为明显。由DC到SR，植株高度略有增加，生物产量明显提高，抽穗后增加的量尤为显著，收获指数变化较小。品种改良明显减小了顶部3叶的着生角度，增加了抽穗期的叶面积，但颖花量的增加超过叶面积的增加，导致粒叶比（颖花数与叶面积之比）提高。随品种的演进，抽穗期根重以及根冠比提高，但根系伤流液量减少。抽穗至成熟根系伤流液和叶片中玉米素＋玉米素核苷浓度下降的速度表现为SR>SDH>SDC。依据品种演进特征，对进一步提高产量的途径进行了讨论。

... 杨建昌等^[15]研究发现, 品种改良明显减小了顶部3叶的着生角度, 增加了抽穗期的叶面积, 但颖花量的增加超过叶面积的增加, 导致粒叶比(颖花数与叶面积之比)提高 ...

2011

0.0

0.916

2010

0.0

1.667

张庆, 殷春渊, 张洪程, 魏海燕, 马群, 杭杰, 李敏, 李国业. 水稻氮高产高效与低产低效两类品种株型特征差异研究. 作物学报, 2010, 36: 1011-1021

Zhang

, Yin C

, Zhang H

, Wei H

, Ma

Q H J

, Li

G Y

. Differences of plant-type characteristics between rice cultivars with high and low levels in yield and nitrogen use efficiency. Acta Agron Sin, 2010, 36: 1011-1021 (in Chinese with English abstract)

A field experiment with 225 kg ha -1 N fertilizer application was carried out in 2006 and 2007 on the farm of Yangzhou university, Jiangsu province, China. Using 12 rice genotypes (six N-efficient and six N-inefficient) selected from 120 rice cultivars grown in Yangzhou during 2004 and 2005 to investigate the relationship among the plant-type characteristics, N use efficiency and yield performance. The results showed that the top three leaves length and drop-angle of high yield and N-efficient rice genotypes were 9.29%, 65.3% lower than those of the low yield and N-inefficient ones, and the width of the top three leaves, population LAI, valid LAI, high valid LAI and LW/LA(leaves weight/leaves area) of the former of were 29.25%, 8.27%, 13.32%, 6.66%, and 5.82% higher than those of the latter on average respectively. The high yield and N-efficient genotypes had higher lodging resistance capability and more reasonable distribution for the length of internodes on stem than the low yield and N-inefficient ones. Yield and N use efficiency negatively correlated (significantly at P P P P

1Key Laboratory of Crop Genetic and Physiology of Jiangsu Province,Yangzhou University;2Key Laboratory of Crop Physiology,Ecology and Cultivation in Middle and Lower Reaches of Yangtze River,Ministry of Agriculture Yangzhou 225009,China

以氮利用高产高效型和低产低效型的12个粳稻品种的大田试验研究了225 kg hm -2 纯氮水平下株型特征的类型间差异，以及株型特征、倒伏性状、产量和氮肥利用率的相互关系。结果表明，水稻上三叶的叶长、叶披垂度，氮高产高效型品种显著低于氮低产低效型，平均分别低9.29%、65.3%；上三叶的叶宽、群体LAI、有效LAI、高效LAI和比叶重，氮高产高效型平均分别比氮低产低效型高29.25%、8.27%、13.32%、6.66%和5.82%。相对于氮低产低效型品种，氮高产高效型品种植株茎秆节间配置合理，抗倒能力增强。水稻的产量、氮肥利用效率与植株上三叶的叶长、叶披垂度均呈显著或极显著负相关，与上三叶的叶宽、群体LAI、有效LAI、高效LAI和比叶重呈显著或极显著正相关，与抗折力、弯曲力矩和秆型指数呈显著或极显著正相关。尤其是剑叶、倒三叶的叶长和叶宽对氮素利用效率的作用较大。说明良好的株型结构是氮肥利用效率提高的前提。

... 张庆等^[17]研究表明, 与低产低效型品种相比, 高产高效型品种水稻上三叶的叶长和叶披垂度较低, 上三叶的叶宽、群体LAI、有效LAI和比叶重等均较高 ...

2012

0.0

1.667

李敏, 张洪程, 杨雄, 葛梦婕, 马群, 魏海燕, 戴其根, 霍中洋, 许轲. 不同氮利用效率基因型水稻茎秆特性比较. 作物学报, 2012, 38: 1277-1285

, Zhang H

, Yang

, Ge M

, Ma

, Wei H

, Dai Q

, Huo Z

, Xu

Comparison of culm characteristics with different nitrogen use efficiencies for rice cultivars.

Acta Agron Sin, 2012, 38: 1277-1285 (in Chinese with English abstract)

The morphological and physiological characteristics of low-yielding and low N-efficiency, high-yielding and medium N-efficiency, high-yielding and high N-efficiency of rice cultivars were investigated using six representative japonica varieties under their respective optimal N application levels. The results showed that compared with low-yielding varieties, high-yielding varieties showed shorter basal 1st internodes, longer basal 6th internodes, increased plant height, larger culm diameter and wall thickness, significantly higher dry weight of stem and sheath, obviously increased K and Si content in culm and sheath, and a higher ratio of Si to N. As a result of their improved stalk physical characteristics and the different chemical compositions, the comprehensive breaking resistance of the culm of high-yielding cultivars was improved evidently, and the lodging index reduced as well. The cultivars with different N-efficiencies displayed diverse culm morphological and physiological characteristics even though they were all in one high-yielding level. Comparing with medium N-efficiency genotypes, the high N-efficiency ones had longer 4th and 5th internode lengths, increased culm wall thickness, in spite of slightly decreased culm diameter, resulted in higher dry weight and strengthened plumpness status of culm, significantly decreased content of Si in stem and sheath, decreased N content, and remarkably unchanged K content. Given these results, for the purpose of further increasing the nitrogen fertilizer use efficiency of the high-yielding varieties, it would be favorable to properly increase 4th and 5th internode length, thus improving top three leaves configuration, to appropriately reduce the culm diameter while increasing wall thickness and filling degree for the sake of stalk breaking resistance and transporting capability, and to properly reduce the content of Si in culm and sheath to facilitate the translocation of nitrogen.

1 Innovation Center of Rice Technology in Yangtze Rice Valley, Ministry of Agriculture / Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China; 2 Rice Research Institute of Guizhou Province, Guiyang 550006, China

选用低产氮低效型、高产氮中效型和高产氮高效型具有代表性的6个粳稻品种，在各基因型各自最适氮素水平下，研究了茎秆力学特性、物理特性和化学成分含量的差异及其与氮效率的关系。结果表明：(1)较之低产类型品种，高产类型品种茎秆基部N 1 节间变短、N 6 节间变长，株高有所增加；茎粗、茎壁厚、茎鞘干重均极显著增加；茎鞘的K、Si含量极显著增加，N含量显著降低。由于茎秆物理性状的改善及化学成分的差异导致茎秆综合抗折力明显提高，倒伏指数降低。(2)同为高生产力类型品种，因氮效率的差异茎秆形态生理特征表现不同。较之高产氮中效类型，高产氮高效类型水稻品种茎秆N 4 、N 5 节间变长；茎秆粗度略有降低，但茎壁厚增加，表现茎秆干重增加，充实度加强；茎鞘的K含量无明显变化，但Si含量显著降低，N含量也呈降低趋势。对于高产品种，适当增加N 4 、N 5 节间长度以改善叶片配置，适当降低茎粗而提高壁厚和充实度以保证茎秆抗折力和输导能力，适当降低茎鞘Si含量以促进氮素的转移，有利于进一步提高氮肥利用率。

1997

0.0

1.889

袁隆平, 武小金, 颜应成, 罗孝和. 水稻广谱广亲和系的选育策略. 中国农业科学, 1997, 30(4): 1-8

Yuan L

, Wu X

, Yan Y

, Luo X

A strategy for developing wide spectrum compatibility rice line.

Sci Agric Sin, 1997, 30(4): 1-8 (in Chinese with English abstract)

湖南杂交水稻研究中心,长沙 410125

选育广谱广亲和系对于利用水稻亚种间杂种优势具有极其重要的意义。根据现有的研究结果，初步假设水稻亚种间亲和性基本上可以分为四大类：广谱广亲和性、部分广亲和性、弱亲和性和非亲和性。决定亚种间亲和性的基因可能有两类：一类是广亲和基因Wc，另一类是辅助亲和性基因Sc。这种辅助亲和性基因既有可能是几个主基因，也有可能是一系列微效多基因。广谱广亲和系可能同时完全具备Wc和Sc基因；部分广亲和系可能只具备Wc基因；弱亲和系可能只具备Sc基因；非亲和系则两者都不具备。选育广谱广亲和系的途径有二个，一是利用现有的广谱广亲和种质直接转育，二是利用部分广亲和系与弱亲和系杂交，组合广亲和基因与辅助亲和性基因。筛选广谱广亲和系可以分三步进行：初选，即在低世代用两个具有代表性的品系进行测交鉴定；复选，即在中世代增加测交品系个数，进一步作亲和性鉴定；决选，即用更多的非亲和品系与当选株系测交，在进一步确定广谱广亲和性的同时筛选优势组合。

... 籼粳亚种间杂交稻蕴藏着巨大的杂种优势, 其应用一直被认为是进一步提高水稻产量的有效途径^[19,20] ...

2000

1.643

0.0

Li R

, Jiang T

, Xu C

, Li X

, Wang X

Relationship between morphological and genetic differentiation in rice (Oryza sativa L. ).

Euphytica, 2000, 114: 1-8

Indica and Japonica differentiation under the species of Oryza sativa L. has been documented using morphological characters, isozyme and DNA markers. We illustrate marker assisted analysis of genetic differentiation with a sample of RFLP and isozyme data from 111 accessions of rice lines and compare it to morphological differentiation based on Cheng and Wang' morphological character index. The results lead to three main conclusions: (1) Among the 92 polymorphic cloned probes deliberately selected, 41 were highly associated with Indica-Japonica differentiation, suggesting such differentiation was an accumulative process of gene change at multiple loci; (2) There exist two main types of relationships between morphological and genetic differentiation. For well differentiated Indica and Japonica (as the case in the control subset), morphology and genomic constitutions are synchronously specialized. For incompletely differentiated rice lines (as the case in the subset of DH lines), morphological differentiation and genetic differentiation are asynchronous. (3)Synthetic or incompletely differentiated rice lines with different genomic constitutions are often adapted to specific environments. These results indicate that marker alleles provide applied breeders with effective ways to identify, track, and incorporate regions of chromosomes governing adaptedness to specific target environments into improved cultivars.

1.China Agricultural University Department of Plant Genetics and Breeding Beijing 100094 China Beijing 100094 China 2.Huazhong Agricultural University National Key Laboratory of Crop Genetic Improvement Wuhan 430070 China Wuhan 430070 China

... 籼粳亚种间杂交稻蕴藏着巨大的杂种优势, 其应用一直被认为是进一步提高水稻产量的有效途径^[19,20] ...

2007

0.0

1.494

... 目前, 国内已有一些关于该系列组合的形态生理特征研究的报道^[21,22], 但供试材料数量相对较少, 对照类型也较单一, 并且缺乏高产栽培条件下的系统比较分析 ...

... 马荣荣等^[21]研究表明, 超高产栽培条件下, 籼粳杂交稻甬优6号群体内部透光率明显优于杂交籼稻汕优63 ...

... 甬优12, 说明甬优系列籼粳杂交稻受光条件好, 光能截获率高, 为其光合系统持续产出及形成高生物学产量提供了生态基础, 这与马荣荣等^[21]在甬优6号上得出的结论基本一致 ...

2007

0.0

1.889

... 本研究观察到, 与其他3种类型品种相比, 甬优系列籼粳杂交稻经济系数低于杂交籼稻, 与杂交粳稻和常规粳稻无显著差异, 说明其经济系数相对稳定, 但无论与杂交粳稻、常规粳稻还是杂交籼稻相比, 其生物学产量和籽粒产量均有大幅提高, 这与许德海等在甬优6号上的研究结果基本一致^[22], 表明增加生物学产量, 稳定收获指数是甬优系列籼粳杂交稻较其他类型品种增产的主要原因, 也可能是进一步提高其籽粒产量的可靠途径 ...

2001

0.0

Campbell C

, Heilman J

, Mclnnes K

, Wilson L

, Medley J

, Wu G

, Cobos D

Seasonal variation in radiation use efficiency of irrigated rice.

Agric For Meteorol, 2001, 110: 45-54

... 1 关于甬优系列籼粳杂交稻的冠层结构特征光能是作物转化利用的对象, 群体光截获量和光能利用率是决定作物产量的重要因素^[23] ...

2008

2.474

0.0

Zhang

, van der Werf

, Bastiaans

, Zhang

, Li

, Spiertz J H

. Light interception and utilization in relay intercrops of wheat and cotton. Field Crops Res, 2008, 107: 29-42

... 已有研究表明, 水稻群体光照分布受到栽培条件的显著影响^[24,25], 同时在基因型或品种类型间存在很大差异^[26,27,28] ...

2003

0.0

1.667

陈雨海, 余松烈, 于振文. 小麦生长后期群体光截获量及其分布与产量的关系. 作物学报, 2003, 29: 730-734

Chen Y

, Yu S

, Yu Z

Relationship between amount or distribution of PAR interception and grain output of wheat communities.

Acta Agron Sin, 2003, 29: 730-734 (in Chinese with English abstract)

在高产条件下,通过设计群体数量大小和植株分布不同的小麦群体类型,测定了小麦生长后期群体光合有效辐射(PAR)截获量及其分布、小麦产量和干物质生产量,研究表明:群体分布状况明显影响群体的光截获量.均匀分布群体比大小行种植的不均匀群体具有较高的光截获率.种植密度对均匀分布群体的光截获量影响较小;随着密度增加,不均匀

... 已有研究表明, 水稻群体光照分布受到栽培条件的显著影响^[24,25], 同时在基因型或品种类型间存在很大差异^[26,27,28] ...

1995

0.0

1.667

Chen W

, Xu Z

, Zhang L

, Zhou H

, Yang S

Effects of different panicle type on canopy properties light distribution and dry matter production of rice population.

Acta Agron Sin, 1995, 21: 83-89

Shenyang Agricultural University, Shenyang,110161

... 已有研究表明, 水稻群体光照分布受到栽培条件的显著影响^[24,25], 同时在基因型或品种类型间存在很大差异^[26,27,28] ...

... 陈温福等^[26]研究表明, 水稻穗型对群体冠层内光分布有较大影响, 直立穗型群体消光系数低, 群体内部光分布较均匀, 更利于群体光能利用和物质生产 ...

2007

0.0

0.895

吕川根, 邹江石, 胡凝, 姚克敏. 水稻叶片形态对冠层特性和光合有效辐射传输的影响. 江苏农业科学, 2007, 23: 501-508

Lü C

, Zou J

, Hu

, Yao K

Effects of morphological factors on canopy structure and transmission of photosynthetically active radiation in rice.

Jiangsu J Agric Sci, 2007, 23: 501-508 (in Chinese with English abstract)

以4个叶片形态差异明显的高产水稻为材料,测定了5个主要叶片形态因子,分析了其对冠层结构、光合有效辐射传输和不同叶层净光合速率的影响.结果表明:品种间叶角和叶片卷曲度的变异系数分别为31.92%和17.15%,而叶长及叶宽的变异系数则分别为5.86%和6.94%,品种间叶角和叶片卷曲度的差异明显大于叶长及叶宽;不同形态叶片的冠层,其叶面积分布也不相同;在水稻冠层辐射传输特征上,光合有效辐射透过率(Tr)、光合有效辐射直接透过率(Dr)和消光系数(K)的大小及变化都与叶片形态因子有密切关系;叶片形态对净光合速率(Pr)和叶面积衰减率(DLAI)有重要作用,影响途径主要是Tr的改变.以稻田地面的Tr=0.03为指标,估算出4个品种可容纳的最大叶面积指数为6.88～8.22.

... 已有研究表明, 水稻群体光照分布受到栽培条件的显著影响^[24,25], 同时在基因型或品种类型间存在很大差异^[26,27,28] ...

... 吕川根等^[27]研究认为, 在相同高产栽培条件下, 紧凑型品种两优培九基部透光率明显高于松散型品种汕优63 ...

2010

0.0

1.237

李迪秦, 唐启源, 秦建权, 张运波, 郑华英, 杨胜海, 陈立军, 邹应斌. 施氮与氮管理模式对超级稻产量和辐射利用率的影响. 核农学报, 2010, 24: 809-814

Li D

, Tang Q

, Qin J

, Zhang Y

, Zheng H

, Yang S

, Chen L

, Zou Y

Effects of nitrogen management patterns on grain yield, solar radiant use efficiency in super hybrid rice.

J Nucl Agric Sci, 2010, 24: 809-814 (in Chinese with English abstract)

Abstract：In order to explore the effects of different nitrogen management patterns on the solar radiation use efficiency(RUE) and yield of super rice, we used super hybrid rice Liangyou 293(LY293) as the experimental material, and three nitrogen management patterns(seven applied nitrogen levels, N0~N6)were designed as the treatments. The result showed that leaf area index,total dry matter, solar radiation use efficiency and yield of super rice were increased as applied nitrogen rate increased. Grain yield of N6 is 99270kg/hm2, about 48% to 335% higher than that of other treatments.But yield and interception photosynthetically active radiation (IPAR)amount of different treatments used different nitrogen management patterns decreased if applied N more than 210kg/hm2. There is an extreme significant difference of total dry weight and RUE between treatment N6 and other treatments.Total dry weight and RUE by N6treatments were are 18537g/m2 and 159g/MJ, respectively, which were higher than that of the other treatments.

College of Agriculture, Hunan Agricultural University, Changsha,Hunan 410128

摘要：通过探讨施氮量和氮管理模式对超级稻产量和辐射利用率的影响，为超级稻超高产栽培和育种提供技术措施和理论依据。以超级稻两优293为材料，于2007年在湖南浏阳进行大田试验，研究了农民管理模式、实地氮肥管理和实时氮肥管理3种氮管理模式、涉及7种氮水平（N0～N6）下水稻的产量及群体辐射利用率。结果表明：随氮肥施用量的增加，群体叶面积指数﹑干物质积累总量﹑产量﹑群体光合有效辐射截获量和辐射利用率都相应增加。处理N6水平(施氮量为210kg/hm2)下，产量最高，为99270kg/hm2，分别比其他处理高48%～335%，当施氮水平超过210kg/hm2时，不同施氮方法，处理产量和群体光合有效截获量反而降低；处理N6干物质积累总量和辐射利用率也最大，分别为18537g/m2和159g/MJ，显著高于其他处理。从而表明，施氮量和氮管理模式对超级稻产量和辐射利用率有重要影响，相同施氮水平下，采用实地氮肥管理更有利于产量和超级稻群体辐射利用率的提高。

... 已有研究表明, 水稻群体光照分布受到栽培条件的显著影响^[24,25], 同时在基因型或品种类型间存在很大差异^[26,27,28] ...

... 张运波等^[28]研究表明, 太阳辐射利用率、冠层截获的有效辐射量呈超级杂交稻、常规稻> ...

2012

2.474

0.0

... 水稻群体光分布与其冠层结构配置密切相关, 群体大小、叶片姿态及群体叶面积分布均影响光照在群体内的分布^[29] ...

1999

1.513

0.0

... 2 关于甬优系列籼粳杂交稻的光合特征籽粒产量是生物学产量和收获指数的乘积, 提高产量是通过增加生物学产量还是通过提高收获指数, 不同学者提出了不同的观点^[30,31] ...

2002

1.513

0.0

... 理论上讲, 提高叶面积指数和提高粒叶比中任何一个均可以提高产量, 但凌启鸿、杨建昌等^[31,32]研究认为, 在适宜的叶面积指数基础上, 大幅提高粒叶比是提高水稻产量的有效途径 ...

1986

0.0

1.889

凌启鸿, 杨建昌. 水稻群体“粒叶比”与高产栽培途径的研究. 中国农业科学, 1986, 19(3): 1-8

Ling C

, Yang J

Studies on “grain-leaf ratio” of population and cultural approaches of high yield in rice plants.

Sci Agric Sin, 1986, 19(3): 1-8 (in Chinese with English abstract)

江苏农学院

... 粒叶比表现为甬优系列籼粳杂交稻明显高于其他3类品种, 可见甬优系列籼粳杂交稻主要通过大幅提高粒叶比来提高产量, 这一结果符合凌启鸿等^[32,33]的观点 ...

2006

0.0

1.889

杨建昌, 杜永, 吴长付, 刘立军, 王志琴, 朱庆森. 超高产粳型水稻生长发育特性的研究. 中国农业科学, 2006, 39: 1336-1345

Yang J

, Du

, Wu C

, Liu L

, Wang Z

, Zhu Q

Growth and development characteristics of super-high-yielding mid-season japonica rice.

Sci Agric Sin, 2006, 39: 1336-1345 (in Chinese with English abstract)

【Objective】Rice is one of the most important food crops in China. The realization of its super-high-yielding has a great significance for food security of the country. This study investigated the growth and development characteristics of the super-high-yielding (grain yield >11 t·ha-1) of rice.【Method】Four mid-season japonica rice cultivars (including lines), Lianjiajing 2, Huajing 5, 0026 and 9823, were grown in a paddy field. Growth analysis was performed during the growth period, and yield and yield components were determined at maturity.【Result】Super-high-yielding rice had more sipkelets per panicle and higher filled-grain percentage than the high-yielding rice (CK, grain yield 8.98～9.16 t·ha-1). There was no significant difference in 1000-grain weight between the super-high-yielding and the CK. Super-high-yield rice exhibited fewer tillers at the early growth stage (from transplanting to jointing), while a higher ratio of productive tillers to total tillers, as compared with the CK. The leaf area index, photosynthetic potential and dry matter accumulation of the super-high-yielding rice were lower than those of the CK at the early growth stage, and their differences were not significant between the two rice types at heading, and were greater for the former than the later after heading. The root-shoot ratio at each growth stage, root bleedings from heading to maturity, grain-leaf ratio, translocation percentage of the matter from stems and sheaths and harvest index of super-high-yielding rice were greater than those of CK. 【Conclusion】 The indexes for the growth and development of super-high-yielding mid-season rice population were suggested, i. e., total spikelets > 4.5 × 104·m-2, filled-grain percentage > 90%, 1000-grain weight > 26 g; ratio of productive tillers > 80%, leaf area index at heading 7.5-8.0, photosynthetic potential during whole growth period > 5×106 m2·d·ha-1, total dry matter weight at maturity >22 t·ha-1, harvest index > 0.51; grain-leaf ratio (No of spikelets per cm2 leaf area) > 0.58; root-shoot ratio at heading stage >0.25 and the amount of root exudates > 5 g·m-2·h-1. The regulation approaches and key cultivation techniques for raising the super-high-yielding population were discussed.

扬州大学农学院

【目的】水稻是中国最重要的粮食作物，实现水稻超高产对保证国家粮食安全有重要作用。本研究旨在探讨超高产水稻（产量>11 t·ha-1）的生长发育特性。【方法】4个中熟粳稻品种（含品系）连嘉粳2号、华粳5号、0026和9823种植于大田，对其物质生产和产量形成进行了观察分析。【结果】与高产水稻（CK，产量为8.98～9.16 t·ha-1）相比，超高产水稻每穗颖花数多，结实率高，千粒重与CK无显著差异；超高产水稻移栽至拔节期的茎蘖数较CK少，但分蘖成穗率较高；超高产水稻的叶面积指数、光合势和干物质积累，生育前期较CK低，抽穗期与CK无显著差异，抽穗后则显著高于CK。超高产水稻各生育时期的根冠比、抽穗至成熟的根系伤流量以及粒叶比、茎鞘物质运转率和收获指数均显著高于CK。【结论】提出了中熟粳稻超高产群体的生育诊断指标：总颖花量>4.5万/m2，结实率>90%，千粒重>26 g；茎蘖成穗率>80%，抽穗期叶面积指数7.5～8.0，全生育期光合势>5×106 m2·d·ha-1，成熟期总干重>22 t·ha-1，收获指数>0.51；抽穗期粒叶比[颖花/叶(cm2)]>0.58，根冠比>0.25，根系伤流量>5 g·m-2·h-1。对培育超高产水稻群体的调控途径与关键栽培技术进行了讨论。

2013

0.0

1.494

李敏, 张洪程, 杨雄, 葛梦婕, 魏海燕, 戴其根, 霍中洋, 许轲. 高产氮高效型粳稻品种的叶片光合及衰老特性研究. 中国水稻科学, 2013, 27: 168-176

, Zhang H

, Yang

, Ge M

, Wei H

, Dai Q

, Huo Z

, Xu

Leaf photosynthesis and senescence characteristics of japonica rice cultivars with high yield and high N-efficiency.

Chin J Rice Sci, 2013, 27: 168-176 (in Chinese with English abstract)

The difference in leaf photosynthesis and senescence characteristics of lowyielding and low Nefficiency, highyielding and medium Nefficiency, highyielding and high Nefficiency rice cultivars was investigated using six representative japonica varieties under their optimum N levels, respectively. The results were as follow: compared with lowyielding varieties, the highyielding ones showed higher population leaf area index (LAI), higher leaf area rates of productive tillers and top 3 leaves, higher flag leaf chlorophyll content (SPAD value)and net photosynthetic rate, and slower senescence of leaves. Among highyielding varieties, as the nitrogen efficiency increased, the population leaf area index (LAI) reduced at the early and middle growth stages (Nn, jointing, and full heading), and increased at maturing. Though with no significant changes during early grain filling stage (0-10 days after full heading), the net photo synthetic rate remarkably increased in middle and late grain filling stages (20-40 days after full heading), which could mainly be attributed to the less breakingdown of chlorophyll, ensuring the higher CO2 assimilation capacity of leaves, the higher SOD activity providing strong antioxidant capacity of the plant, and the lower MDA content implying the less damage degree of membrane lipid. It could be one promising method for further improvement in nitrogen efficiency of highyielding rice varieties to reasonably control the growth of nonproductive and inefficient leaf area before heading, thus properly reducing population leaf area scale before heading and effectively delaying leaves senescence after heading to ensure sustained higher chlorophyll content and net photosynthetic rate of leaves.

1 Rice Research Institute of Guizhou Province, Guiyang 550006, China; 2 Innovation Center of Rice Technology in Yangtze Rice Valley, Ministry of Agriculture, Yangzhou University, Yangzhou 225009, China;

选用6个具代表性的低产氮低效型、高产氮中效型和高产氮高效型粳稻品种，在各自最适氮素水平下，研究了叶片光合、衰老特性的差异及其与氮效率的关系。结果表明，高产类型群体叶面积指数(LAI)、高效叶面积率及有效叶面积率、剑叶叶绿素含量(SPAD值)和净光合速率以及剑叶超氧化物歧化酶(SOD)活性均显著高于低产类型品种，丙二醛(MDA)含量显著低于低产类型。高产品种间比较，随着氮效率提升，生育前中期(够苗、拔节、齐穗期)群体叶面积指数下降，成熟期上升；灌浆前期(齐穗后0 d~10 d)剑叶净光合速率并未明显变化，灌浆中后期(齐穗后20 d~40 d)显著增高，主要原因如下：叶绿素分解少，保证了叶片对CO2的高同化能力；SOD活性高，保证了植株更强的抗氧化能力，MDA含量少，膜脂破坏程度低。说明抽穗前合理控制无效及低效叶面积生长以适当减小群体叶面积规模，抽穗后有效延缓植株衰老，以保证叶片持续较高的叶绿素含量和净光合速率，是促进水稻高产品种进一步提升氮效率的重要途径。

... 众所周知, 水稻叶片光合能力体现在光合面积、单位面积的光合生产能力(光合速率)和光合能力的持续等3个方面^[34] ...