不同年代中籼水稻品种的叶片光合性状

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剧成欣, 陶进, 钱希旸, 顾骏飞, 张耗, 赵步洪, 刘立军, 王志琴, 杨建昌. . 不同年代中籼水稻品种的叶片光合性状. 作物学报, 2015, 42(3): 415-426
JU Cheng-Xin, TAO Jin, QIAN Xi-Yang, GU Jun-Fei, ZHANG Hao, ZHAO Bu-Hong, LIU Li-Jun, WANG Zhi-Qin, YANG Jian-Chang. . Leaf Photosynthetic Characteristics of Mid-season Indica Rice Varieties Applied at Different Decades . ACTA AGRONOMICA SINICA, 2015, 42(3): 415-426 复制到剪切板

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作物学报编辑部

不同年代中籼水稻品种的叶片光合性状

剧成欣¹, 陶进¹, 钱希旸¹, 顾骏飞¹, 张耗¹, 赵步洪², 刘立军¹, 王志琴¹, 杨建昌^1,^*

¹扬州大学江苏省作物遗传生理国家重点实验室培育点 / 粮食作物现代产业技术协同创新中心, 江苏扬州225009

²江苏省里下河地区农业科学研究所, 江苏扬州225007

^* 通讯作者(Corresponding author): 杨建昌, E-mail: jcyang@yzu.edu.cn, Tel: 0514-87979317

第一作者联系方式: E-mail: cxju1124@163.com

收稿日期:2015-07-11 接受日期:2015-11-20 网络出版日期:2015-12-18

基金:本研究由国家自然科学基金项目(31271641, 31201155, 31471438), 中央级科研院所基本科研业务费专项(农业)(201103003, 201203079), 国家“十二五”科技计划项目(2014AA10A605, 2011BAD16B14, 2012BAD04B08, 2013BAD07B09), 江苏省农业三新工程项目(SXGC[2014]313), 江苏省普通高校研究生科研创新计划项目(KYZZ15_0364)和江苏高校优势学科建设工程项目(PAPD)资助

摘要

旨在探明中熟籼稻在品种改良过程中籽粒产量和叶片光合性能的变化特点。以江苏省近70年来不同年代在生产上应用的12个代表性中籼水稻品种(含杂交稻组合)为材料, 依据应用年代将其分为20世纪50—60年代、60—70年代、80—90年代和21世纪00—10年代(超级稻) 4种类型, 研究其产量、冠层结构及叶片光合特性的变化。结果表明, 随品种的改良, 中籼水稻品种的产量不断提高。群体总颖花量、面积指数和粒叶比显著增加, 叶基角减小, 群体透光率、光合势(绿叶面积持续期)、抽穗期剑叶光合速率、气孔导度、蒸腾速率和PSII最大和实际光化学效率以及荧光与非荧光淬灭系数增加。最大叶面积指数和全生育期总光合势与籽粒产量呈极显著正相关。灌浆期剑叶光合速率、气孔导度、PSII最大和实际光化学量子效率以及荧光和非荧光淬灭系数与结实率或粒重呈显著相关。表明在品种改良过程中, 株型和叶片光合性能的改善是中籼水稻产量提高的重要原因。

关键词: 中籼水稻; 株型; 光合速率; 叶绿素荧光; 产量

Leaf Photosynthetic Characteristics of Mid-season Indica Rice Varieties Applied at Different Decades

JU Cheng-Xin¹, TAO Jin¹, QIAN Xi-Yang¹, GU Jun-Fei¹, ZHANG Hao¹, ZHAO Bu-Hong², LIU Li-Jun¹, WANG Zhi-Qin¹, YANG Jian-Chang^1,^*

¹Jiangsu Key Laboratory of Crop Genetics and Physiology / Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou 225009, China

² Lixiahe Region Agricultural Research Institute of Jiangsu, Yangzhou 225007, China

Fund:This study was supported by the grants from the National Natural Science Foundation of China (31271641, 31201155, 31471438), China National Public Welfare Industry (Agriculture) Plan (201103003, 201203079), the National Key Technology Support Program of China (2014AA10A605, 2011BAD16B14, 2012BAD04B08, 2013BAD07B09), Jiangsu “Three-innovation” Agricultural Project (SXGC[2014]313), Jiangsu Creation Program for Post-graduation Students (KYZZ15_0364), and the Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD)

Abstract

The objective of this study was to investigate the changes in grain yield and leaf photosynthetic characteristics of mid-season indica rice varieties during their improvement. Twelve typical indica varieties (including hybrid combinations) applied in the production in Jiangsu Province during the last 70 years were used, and classified into four types of 1950-1960s, 1960-1970s, 1980-1990s, and 2000-2010s (super rice) according to their application decades. The grain yield, canopy structure and leaf photosynthetic characteristics were determined. The results showed that the grain yield was progressively increased with the improvement of varieties. With the process of improvement of varieties, the total number of spikelets, leaf area index and grain-leaf ratio were markedly increased, while the leaf base angel was decreased. The improvement of varieties increased the light transmittance ratio, photosynthetic potential (green leaf area duration), net photosynthetic rate, stomatal conductance, transpiration rate, the maximum and the actual photochemical efficiency of PSII, photochemical quenching coefficient and the non-photochemical quenching coefficient of leaves from heading to maturity. The maximum leaf area index and the total photosynthetic potential during the whole growing season were very significantly correlated with grain yield. The photosynthetic rate, stomatal conductance, the maximum and the actual photochemical efficiency of PSII, photochemical quenching coefficient and the non-photochemical quenching coefficient of leaves during grain filling were significantly correlated with filled grain percentage and grain weight. The results indicate that the increase in grain yield in the process of varietal improvement is mainly attributed to the improvement in plant types and leaf photosynthetic characteristics in mid-season indica rice.

Keyword: Middle-season indica rice; Plant type; Photosynthetic rate; Chlorophyll fluorescence; Grain yield

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光合作用是植物生物产量和经济产量形成的基础, 作物产量的90%~95%直接或间接来自光合作用^{[1, 2]}。作物经济产量的高低是由光合生产力和呼吸消耗以及经济系数(又称收获指数)的大小决定的, 其中光合生产力又由光合速率、光合面积和光合功能期决定^{[3, 4]}。叶片作为光合作用的主要器官直接影响作物的光能利用和干物质积累, 尤其是生育后期功能叶光合作用对作物产量起着至关重要的作用^{[5, 6, 7]}。叶绿素荧光作为光合作用研究的探针, 在植物逆境生理研究和光能利用探测等方面得到广泛应用^{[8, 9]}。前人对植物光合作用的机理及其调控途径等作了大量的研究。但有关叶片光合速率与作物产量的关系, 却有着不同的结果: 有的认为两者是正相关^{[10, 11]}, 也有的认为两者无相关性^[12]。

自20世纪50年代末的矮化育种和70年代中期的三系杂交育种以来, 通过株型的改良, 水稻单产获得了大幅度的提高; 20世纪90年代袁隆平提出了中国超级稻“ 形态改良与杂种优势利用相结合” 的育种技术路线, 认为优良的植株形态可以实现水稻超高产的突破^{[2, 13, 14, 15]}。前人从水稻品种改良^{[16, 22]}、亚种间属性^{[17, 18]}以及高产高效协同型品种^[19]等方面对水稻的株型特征进行了深入研究, 认为顶部3叶的着生角度和披垂度减小、有效叶面积和粒叶比提高等形态改良是水稻产量提高的重要原因, 这些结果也为水稻高产高效栽培及品种改良提供了参考依据。

长江中下游地区是我国水稻主产区, 水稻播种面积占全国的51.2%, 稻谷产量占51.3%。该区域稻田种植制度多样, 双季稻和单季中、晚稻都有, 不少地区籼粳并存; 在各品种类型中, 以中籼稻所占比例最大^{[20, 21]}。以往虽然对中籼水稻品种产量与株型的演进特征、根系形态生理特性、氮素利用效率和米质变化等作了较多的研究^{[22, 23, 24, 25]}, 但是关于中籼品种改良过程中叶片光合特性和主要荧光参数的变化特点及其与产量的关系, 缺乏深入研究。本试验以不同年代代表性中籼水稻品种为材料, 研究了水稻主要生育期株型和叶片光合特性的变化特征, 以期进一步阐明水稻品种改良过程中叶片光合作用的变化特点及其与产量的关系, 为水稻高产育种和栽培提供理论依据。

1 材料与方法

1.1 材料与试验设计

试验于2012— 2013年在江苏省扬州市里下河地区农业科学研究所进行。前茬作物为小麦, 土壤为沙质壤土, 耕作层含有机质2.1%、有效氮106.3 mg kg^-1、速效磷30.2 mg kg^-1、速效钾87.6 mg kg^-1。试验选择近70年来在江苏省大面积种植的代表性中籼水稻品种, 依据品种的种植年代将其分为20世纪50— 60年代、60— 70年代、80— 90年代和21世纪00— 10年代(超级稻) 4个类型, 共计12个品种(表1), 各品种在扬州均能正常抽穗结实。其中, 20世纪50— 60年代主要收集整理的是地方优良品种。试验所选用的3个超级稻品种均已通过农业部认定。在田间安装小型气象站记录两年试验生长季节中的主气象因子, 生长期的光照强度、平均温度湿度等数据列于表2。

表1 供试的中籼水稻品种 Table 1 The tested mid-season indica rice varieties

表2 两年试验中的主要气象因子 Table 2 Main meteorological factors in 2012 and 2013

由于不同品种的全生育期相差较大, 本试验采用分期播种来保证各品种抽穗期尽可能一致。20世纪50— 60年代品种于5月24日播种, 60— 70年代品种于5月17日播种, 80— 90年代和21世纪00— 10年代品种(超级稻)于5月8日播种, 6月13日移栽, 株行距15 cm × 20 cm, 杂交籼稻单本移栽, 常规籼稻双本移栽。

采用随机区组设计, 小区面积为16 m², 重复3次。全生育期施用纯氮210 kg hm^-2, 将氮肥折合成尿素(含纯氮46.4%)按基肥(移栽前1 d)∶ 分蘖肥(移栽后7 d)∶ 穗肥(枝梗分化期)=5∶ 1∶ 4施用。移栽前对各小区施用过磷酸钙(含P₂O₅ 13.5%) 300 kg hm^-2和氯化钾(含K₂O 52.0%) 195 kg hm^-2。按照常规高产栽培管理水分等, 严格控制全生育期病虫草害。

1.2 测定项目及方法

1.2.1 植株叶面积和氮含量

记录各品种的抽穗日期, 于抽穗期从各小区取代表性植株10株, 使用量角器测定植株顶三叶的叶基角; 使用SPAD仪分别测定倒数3张叶片的叶绿素含量。使用叶面积仪分别扫描剑叶、倒二叶和倒三叶的叶面积, 并烘干称重, 计算比叶面积和比叶重, 比叶面积(SLA)=叶面积/干物重。绿叶面积持续期即光合势分别于移栽期、穗分化始期、抽穗期和成熟期取每品种代表性植株6穴的所有绿叶, 用叶面积测定仪测定叶面积。光合势(m² d m^-2)=1/2(L₁+L₂)× (t₂-t₁)。式中L₁和L₂为前后两次测定的叶面积 (m² m^-2), t₂-t₁表示前后两次测定的时间间隔(d)。根据成熟期总实粒数(总颖花数× 结实率)和抽穗期绿叶面积计算粒叶比, 即粒叶比=总实粒数/抽穗期叶面积。将烘干后叶片粉碎过筛, 使用凯氏定氮法测定单位重量叶片氮素含量, 单位面积叶片氮素含量(SLNC)用单位重量叶片氮素含量与比叶面积的比值表示。

1.2.2 植株冠层透光率

于抽穗期, 用英国DELTA-T公司生产的SunScan冠层分析仪Type SS1分别测定各小区株间和行间不同高度的光合有效辐射(PAR), 离地面高度分别为0 cm、20 cm、40 cm、60 cm、80 cm和冠层顶部, 重复3次。由于不同处理PAR的测定并不同步, 存在一定的测量时间差, 入射PAR会在一定的范围内上下波动; 因此, 本文采用相对值透光率来反应水稻冠层光合有效辐射的空间分布。各高度的透光率LT=各高度PAR/冠层顶部PAR× 100。参照程占慧等^{[9, 26, 27, 28]}方法计算: 光合作用效率(PE)=光合速率(P_n)/光合有效辐射(PAR); 光合氮素利用效率(PNUE)=光合速率(P_n)/单位面积叶片氮素含量(SLNC)。

1.2.3 叶片光合速率与荧光动力学参数

在抽穗期, 用带有荧光叶室(LI-6400-40, Li-Cor Inc.)的便携式光合仪(Li-6400, Li-Cor Inc., Lincoln, NE, USA)测定植株剑叶的光合速率和叶绿素荧光参数^{[29, 30]}。光量子通量密度(PFD)为1000 μ mol m^-2 s^-1, 使用CO₂钢瓶, 叶室CO₂浓度为400 μ mol mol^-1。叶片暗适应20 min后测定初始荧光(F_o), 然后给定一个饱和光脉冲测定最大荧光(F_m); 将叶片置叶室内300 s, 待荧光恒定时记录净光合速率(P_n)、胞间CO₂浓度(C_i)、气孔导度(G_s)、蒸腾速率(T_r)和稳态荧光(F_s)等气体交换数据; 给定一个饱和光脉冲( > 8500 μ mol m^-2 s^-1持续0.8 s)来测定能化类囊体最大荧光(F_m'); 关闭作用光供应一个暗脉冲(使用远红光优先激发PSI并迫使电子从PSII流出)来测定能化类囊体最小荧光(F_o')。并从以上数据计算以下叶绿素荧光参数, PSII最大光化学量子效率(F_v'/F_m')=(F_m'-F_o')/F_m', PSII实际光化学量子效率(Φ _PSII)=(F_m-F_s)/F_m', 光化学猝灭系数(q_P)= (F_m'-F_s)/(F_m'-F_o')、非光化学猝灭系数(q_N)=(F_m-F_m′ )/(F_m-F_o)。

1.2.4 考种与计产

收获前1 d, 取每小区1 m²考查穗数, 并按照平均穗数取样10株考种, 采用水漂法测定结实率, 每小区实收2 m²计产。

1.3 数据分析

用Microsoft Excel 2003软件整理数据, SAS9.2软件统计分析数据, SigmaPlot10.0绘图。

2 结果与分析

2.1 产量与主要光合特性的方差分析

由表3可知, 产量、叶面积指数、比叶面积、透光率、剑叶光合速率和PSII实际光化学效率等在不同年代中籼水稻品种间存在极显著差异。但年度(两年试验)及年度与品种、年度与年代(品种改良过程中的4个年代)的互作均无显著差异(表3)。因两年的试验结果趋势一致, 故本文中除产量用两年的数据表示外, 其余数据用两年的平均数表示。

2.2 产量及其构成因素

随着品种改良, 中籼水稻品种的产量显著增加。1950— 1960s、1960— 1970s、1980— 1990s、2000— 2010s的品种在两年(2012年和2013年)平均产量分别为4.77、6.83、8.47、9.16 t hm^-2, 增幅依次为43.19%、24.01%和8.15% (表4)。从产量构成因素分析, 品种改良使产量增加的主要原因是总颖花量的增加, 每穗粒数和总颖花量表现出相同的趋势。穗数随品种改良有所减少。20世纪80年代以来的品种的千粒重显著高于20世纪80年代之前的品种。随品种改良, 结实率表现出先增加后减小的趋势。与早期中籼稻相比, 超级稻的结实率表现出低而不稳(表4)。较低的结实率制约了超级稻品种产量潜力的发挥。

2.3 株高和叶面积指数

随品种改良, 不同年代中籼水稻品种的株高表现先减小后增加的趋势(表5)。叶基角随着品种改良逐渐降低。超级稻品种的叶基角与20世纪50— 60年代品种相比降低了61.39° 。最大叶面积指数和粒叶比均随着品种应用年代的推移而增加(表5)。表明在品种改良过程中源(叶面积)和库(总颖花量)均增加, 但库的增加超过了源的增长。随着生育期的延长, 各品种的绿叶面积持续期(光合势)为2000— 2010s > 1980— 1990s > 1960— 1970s > 1950— 1960s (图1)。

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	图1 中籼水稻品种绿叶面积持续期的变化 Fig. 1 Changes in green leaf area duration of mid-season indica rice varieties

2.4 比叶面积与叶绿素含量

在抽穗期, 中籼水稻品种植株顶部3叶的比叶面积和叶绿素含量随品种改良而增加(表6)。与20世纪80— 90年代品种相比, 超级稻品种的比叶面积和叶绿素含量没有显著差异, 但显著高于早期年代的品种。自20世纪80年代以来, 剑叶和倒二叶的比叶面积显著增加, 倒三叶的比叶面积增加不显著; 倒二叶和倒三叶的叶片叶绿素含量显著增加, 剑叶的叶绿素含量增加不显著(表6)。

表6 不同年代中籼水稻品种比叶面积和叶绿素含量的变化 Table 6 Changes in specific leaf area and chlorophyll content (SPAD value) of mid-season indica rice varieties

2.5 透光率与光合作用效率

抽穗期不同年代中籼水稻品种冠层各高度的透光率均随品种改良表现出增加的趋势(图2)。在0~40 cm, 超级稻品种的透光率与20世纪80— 90年代品种相比差异不显著, 但显著高于1950— 1960s的品种。在离地面40~80 cm的群体冠层, 不同年代品种的透光率表现出2000— 2010s > 1980— 1990s > 1960— 1970s > 1950— 1960s (图2)。

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	图2 中籼水稻品种抽穗期冠层不同高度透光率的变化Fig. 2 Changes in light transmission rate at different height at the heading stage of mid-season indica rice varieties

表3 中籼水稻品种产量和主要光合荧光特性的方差分析 Table 3 Analysis of variance F-value of grain yield and main photosynthetic and chlorophyll fluorescence parameters of mid-season indica rice varieties

表4 中籼水稻品种产量及其构成因素的变化 Table 4 Changes in grain yield and its components of mid-season indica rice varieties

品种和年代 Cultivar and decade	产量 Yield (t hm^-2)	穗数 No. of panicles (× 10⁴ hm^-2)	穗粒数 Spikelets per panicle	总颖花量 Total spikelets (× 10⁶ hm^-2)	结实率 Filled grains (%)	千粒重 1000-grain weight (g)
2012
黄瓜籼 Huangguaxian	4.53 d	261.37 a	118.22 e	308.99 e	60.47 d	24.26 cd
银条籼 Yintiaoxian	5.01 d	268.66 a	129.24 e	347.22 d	57.95 d	24.91 c
南京1号 Nanjing 1	4.80 d	242.10 bc	132.83 de	321.58 de	60.26 d	24.79 c
1950-1960s平均 Average	4.78	257.38	126.76	325.93	59.56	24.65
台中籼 Taizhongxian	6.89 c	252.96 ab	138.36 cd	350.00 cd	80.34 bc	24.49 cd
珍珠矮 Zhenzhu’ ai	6.70 c	247.85 ab	140.85 cd	349.10 cd	79.51 bc	24.13 d
南京11 Nanjing 11	6.75 c	241.31 bc	154.50 bc	372.82 bc	75.62 c	23.95 d
1960-1970s平均 Average	6.78	247.37	144.57	357.31	78.49	24.19
扬稻2号 Yangdao 2	8.69 ab	235.30 d	158.85 bc	373.77 bc	89.24 a	26.05 ab
扬稻6号 Yangdao 6	8.19 b	235.54 d	166.06 b	391.14 b	81.75 bc	25.62 bc
汕优63 Shanyou 63	8.70 ab	230.68 d	166.89 b	384.98 bc	85.18 ab	26.52 a
1980-1990s平均 Average	8.53	233.84	163.93	383.30	85.39	26.06
两优培九 Liangyoupeijiu	9.20 a	212.95 e	217.38 a	462.91 a	78.13 bc	25.44 bc
扬两优6号 Yangliangyou 6	9.41 a	226.77 de	193.31 ab	438.37 a	80.99 bc	26.50 a
II优084 II you 084	8.78 ab	212.48 e	209.47 a	445.08 a	75.95 c	25.96 bc
2000-2010s平均 Average	9.13	217.40	206.72	448.79	78.36	25.97
2013
黄瓜籼 Huangguaxian	4.51 e	228.58 b	136.63 ef	312.31 e	57.96 e	24.90 ab
银条籼 Yintiaoxian	4.73 e	262.21 a	127.56 f	334.48 e	57.61 e	24.55 ab
南京1号 Nanjing 1	4.96 e	249.48 ab	137.66 ef	343.43 de	60.32 e	23.92 bc
1950-1960s平均 Average	4.73	246.76	133.95	330.07	58.63	24.46
台中籼 Taizhongxian	7.01 d	255.49 a	145.93 de	372.84 cd	82.72 bc	22.74 d
珍珠矮 Zhenzhu’ ai	6.91 d	241.36 ab	144.17 de	347.97 de	80.58 bc	24.66 ab
南京11 Nanjing 11	6.66 d	236.28 ab	141.13 de	333.46 e	82.20 bc	24.30 b
1960-1970s平均 Average	6.86	244.38	143.74	351.42	81.83	23.90
扬稻2号 Yangdao 2	8.36 bc	258.36 a	161.12 cd	408.52 bc	85.65 ab	23.90 bc
扬稻6号 Yangdao 6	8.02 c	211.97 c	164.34 cd	351.61 de	84.93 ab	26.86 a
汕优63 Shanyou 63	8.74 bc	213.28 bc	173.94 bc	370.98 cd	87.39 a	26.97 a
1980-1990s平均 Average	8.37	227.87	166.47	377.04	85.99	25.91
两优培九 Liangyoupeijiu	8.96 ab	221.16 bc	195.53 b	432.43 ab	78.29 cd	26.46 a
扬两优6号 Yangliangyou 6	9.13 a	216.96 bc	219.49 a	477.42 a	74.52 d	25.66 ab
II优084 II you 084	9.46 a	211.77 c	221.53 a	469.13 a	77.34 cd	26.06 a
2000-2010s平均 Average	9.18	216.63	212.18	459.66	76.72	26.06

Values within the same column and year followed by a different letter are significantly different at the 0.05 probability level.

同栏同年内比较, 标以不同字母的值在P=0.05水平上差异显著。

表4 中籼水稻品种产量及其构成因素的变化 Table 4 Changes in grain yield and its components of mid-season indica rice varieties

表5 中籼水稻品种株高、叶面积指数和粒叶比的变化 Table 5 Changes in plant height, leaf area index and filled grain-leaf ratio of mid-season indica rice varieties

叶片氮含量随品种改良逐渐增加(表7)。不同年代中籼水稻品种的光合作用效率、光合氮素利用效率和氮肥偏生产力均表现为2000— 2010s > 1980— 1990s > 1960— 1970s > 1950— 1960s (表7)。

表7 中籼水稻品种光能利用率和光合氮素利用效率的变化 Table 7 Changes in photosynthetic efficiency and photosynthetic nitrogen utilization efficiency of mid-season indica rice varieties

2.6 光合速率与叶绿素荧光

自抽穗至成熟, 叶片的净光合速率在抽穗期达到最大, 此后逐渐下降(图3-A)。气孔导度和蒸腾速率与净光合速率表现出相似的变化趋势(图3-B, C)。胞间CO₂浓度表现出先增加后降低的趋势(图3-D)。在抽穗期和抽穗后10 d, 不同年代中籼水稻品种的净光合速率随品种改良不断升高, 表现出2000— 2010s > 1980— 1990s > 1960— 1970s > 1950— 1960s。在抽穗后10 d和抽穗后20 d, 超级稻品种的光合速率的降幅较大, 表现出1980— 1990s > 2000— 2010s > 1960— 1970s > 1950— 1960s (图3-A), 气孔导度和蒸腾速率的变化与净光合速率的变化趋势相一致(图3-B, C)。

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	图3 中籼水稻品种叶片光合速率的变化Fig. 3 Changes in leaf photosynthetic traits of the mid-season indica rice varieties at different growth stages

自抽穗至成熟, 不同年代中籼水稻品种的PSII最大光化学量子效率(F_v'/F_m')和实际光化学量子效率(Φ _PSII)均随品种改良而逐渐增加, 随灌浆进程而逐步降低(图4-A, B)。在抽穗期, 超级稻的F_v'/F_m'和Φ _PSII在各类品种中最大, 但抽穗后超级稻的F_v'/F_m'和Φ _PSII下降幅度较大。在抽穗后20 d, 超级稻的F_v'/F_m'和Φ _PSII显著低于20世纪80— 90年代品种(图4-A, B)。这可能是超级稻结实率较低的一个重要生理原因。不同年代中籼水稻品种的光化学淬灭系数(q_P)和非光化学淬灭系数(q_N)与F_v'/F_m'及Φ _PSII表现出类似的变化趋势(图4-C, D)。

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	图4 抽穗灌浆期中籼水稻品种叶绿素荧光参数的变化Fig. 4 Changes in leaf chlorophyll fluorescence parameters in tested rice varieties during the heading and grain filling

2.7 叶片光合性状与产量的相关

水稻籽粒产量与各品种的最大叶面积指数、抽穗期剑叶光合速率和全生育期总光合势呈极显著正相关(图5-A~C)。抽穗后剑叶光合速率、气孔导度、PSII最大和实际光化学量子效率以及荧光和非荧光淬灭系数与结实率或粒重的相关达到显著或极显著水平(表8)。说明提高叶片光合性能有利于增加水稻产量。

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	图5 中籼水稻品种叶片光合性状与产量的相关性Fig. 5 Correlations of leaf photosynthetic characteristics with grain yield of mid-season indica rice varieties

表8 抽穗后叶片光合荧光参数与结实率和粒重的相关关系 Table 8 Correlations of leaf photosynthetic chlorophyll fluorescence traits with filled grain rate and grain weight of mid-seasonindica rice varieties

3 讨论

本研究表明, 随着品种改良, 中籼水稻品种的产量逐渐增加。从产量构成因素分析, 产量的增加主要在于总颖花量的增加, 而总颖花量的增加主要在于每穗颖花数的增加。说明在长江中下游地区, 增加每穗颖花数是中籼水稻品种获取高产的一条重要途径。

在作物品种改良过程中, “ 源” 即光合作用是如何适应库(籽粒总库容)的增大?相关研究报道甚少。本研究观察到, 中籼水稻品种的改良既增加了源的数量(叶面积指数、绿叶面积持续期等), 又提高了源的质量(叶片光合速率、叶绿素荧光参数等), 使得在库容量增加的同时保证了光合同化物的供应, 从而实现了源库协调和产量的提高。

本研究结果显示, 中籼水稻品种改良提高了群体最大叶面积指数。通常, 群体叶面积指数的增加会减少群体的透光率, 影响基部叶片的光合作用, 进而影响根系活力^{[31, 32]}。但本研究观察到, 随品种改良, 水稻植株中下部透光率增加, 群体中下部光合有效辐射空间分布改善。这主要得益于稻株顶部3叶叶基角减小, 改善了群体的受光姿态。不仅如此, 品种改良还改善了叶面积的组成, 即增加了植株顶部3叶面积占总叶面积的比例。有研究表明, 稻株顶部3叶的生长与穗分化同步, 与抽穗后的籽粒灌浆充实有着密切的关系, 因而被称为高效叶面积^{[5, 32]}。因此, 植株顶部3叶面积占总叶面积的比例的增加和受光姿态的改善, 这是在中籼水稻品种改良过程中叶片光合性状得以改善的一个重要方面。

中籼水稻品种改良还突出地表现在“ 源” 质量的改善。本研究表明, 随着品种改良, 各品种的光合速率、气孔导度和蒸腾速率都显著提高。PSII最大光化学量子效率(F_v'/F_m')、实际光化学量子效率(Φ _PSII)、荧光光化学淬灭效率(q_P)和非光化学淬灭系数(q_N)随品种改良逐渐增加。F_v'/F_m'指类囊体能化时PSII固有效率, 反映开放的PSII反应中心原初光能转化效率; Φ _PSII反映了PSII的实际光化学量子效率, 它和非循环式电子传递的量子产量密切相关, 表示部分反应中心关闭情况下的实际原初光能转化效率。q_P在一定程度上反映了PSII反应中心的开放程度; q_N则反映了PSII反应中心对天线色素吸收过量光能后的热耗散能力及光合机构的损伤程度^[34]。因此, F_v'/F_m'、Φ _PSII、q_P和q_N值大, 是叶片光合性能好和对环境适应性强的指标。品种改良增强了叶片光合速率, 改善了叶绿素荧光参数, 这是中籼水稻品种改良过程中叶片光合性状得以改善的另一个重要方面。

以往有关叶片光合性能与作物产量的关系, 存在着不同的研究结论^{[10, 11, 12]}。本研究表明, 抽穗后剑叶光合速率、气孔导度、PSII最大和实际光化学量子效率以及荧光和非荧光淬灭系数与结实率或粒重均呈显著或极显著正相关。说明通过遗传改良或栽培措施提高叶片光合性能, 有利于提高水稻产量。

本研究还观察到, 随品种改良, 叶片氮含量和光合氮素利用效率也显著提高。较高的光合氮素利用效率不仅有利于提高产量, 而且还可以提高水稻氮肥利用效率^{[27, 28]}。说明品种改良不仅改善了叶片光合性能, 而且提高了中籼水稻品种的氮肥利用效率。至于品种改良提高光合氮素利用效率的原因和机理还有待深入研究。

值得注意的是, 现代超级稻品种虽然表现出较大的库容和较高的产量, 但与1960— 1970s或1980— 1990s的品种相比, 超级稻的结实率较低, 这已成为限制其产量潜力发挥的重要因子^{[35, 36, 37, 38]}。从本研究结果分析, 超级稻结实率低可能与其灌浆期叶片光合速率及F_v'/F_m'、Φ _PSII、q_P和q_N值下降快有密切关系。深入研究超级稻结实率较低的原因和提高其结实率的途径, 对于实现超级稻的增产潜力有重要意义。

4 结论

品种改良显著提高了中籼水稻品种的产量, 每穗颖花数的增加是产量提高的主要原因。品种改良显著提高了叶面积指数、绿叶面积持续期、叶片光合速率和PSII光化学效率, 这是产量提高的重要物质生产基础。超级稻结实率较低与其灌浆期叶片光合速率及PSII光化学效率参数值下降较快有密切关系。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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[24]	剧成欣, 陶进, 钱夕旸, 顾骏飞, 赵步洪, 杨凯鹏, 王志琴, 杨建昌. 不同年代中籼水稻品种的产量与氮肥利用效率. 作物学报, 2015, 41: 422-431 Ju C X, Tao J, Qian X Y, Gu J F, Zhao B H, Yang K P, Wang Z Q, Yang J C. Grain yield and nitrogen use efficiency of mid-season indica rice cultivars applied at different decades. Acta Agron Sin, 2015, 41: 422-431 (in Chinese with English abstract) [本文引用:1]
[25]	张耗, 谈桂露, 孙小淋, 刘立军, 杨建昌. 江苏省中籼水稻品种演进过程中米质的变化. 作物学报, 2009, 35: 2037-2044 Zhang H, Tan G L, Sun X L, Liu L J, Yang J C. Changes in grain quality during the evolution of mid-season indica rice cultivars in Jiangsu Province. Acta Agron Sin, 2009, 35: 2037-2044 (in Chinese with English abstract) [本文引用:1]
[26]	冯大兰, 刘芸, 钟章成, 杨娟, 谢君. 三峡库区消落带芦苇(Phragmites communis (Reed))的光合生理响应和叶绿素荧光特性. 生态学报, 2008, 28: 2013-2021 Feng D L, Liu Y, Zhong Z C, Yang J, Xie J. Photosynthesis and chlorophyll fluorescence parameters of the reed (Phragmites communis) grown in the hydro-fluctuation belt of Three Gorges Reservoir Area. Acta Ecol Sin, 2008, 28: 2013-2021 (in Chinese with English abstract) [本文引用:1]
[27]	Ju C X, Roland J B, Wang Z Q, Zhang H, Liu L J, Yang J C, Zhang J H. Root and shoot traits for rice varieties with higher grain yield and higher nitrogen use efficiency at lower nitrogen rates application. Field Crops Res, 2015, 175: 47-55 [本文引用:2]
[28]	Pang J Y, Palta J A, Rebetzke G J, Milroy S P. Wheat genotypes with high early vigour accumulate more nitrogen and have higher photosynthetic nitrogen use efficiency during early growth. Funct Plant Biol, 2014, 41: 215-222 [本文引用:2]
[29]	Gu J F, Yin X Y, Tjeerd J S, Wang H Q, Paul C S. Physiological basis of genetic variation in leaf photosynthesis among rice (Oryza sativa L. ) introgression lines under drought and well-watered conditions. J Exp Bot, 2012, 63: 5137-5153 [本文引用:1]
[30]	Gu J F, Yin X Y, Paul C S, Tjeerd J S, Wang H Q. Using chromosome introgression lines to map quantitative trait loci for photosynthesis parameters in rice (Oryza sativa L. ) leaves under drought and well-watered field conditions. J Exp Bot, 2012, 63: 455-469 [本文引用:1]
[31]	Maddonni G A, Otegui M E, Cirilo A G. Plant population density, row spacing and hybrid effects on maize canopy architecture and light attenuation. Field Crops Res, 2001, 71: 183-193 [本文引用:1]
[32]	吕丽华, 赵明, 赵久然, 陶洪斌, 王璞. 不同施氮量下夏玉米冠层结构及光合特性的变化. 中国农业科学, 2008, 41: 2624-2632 Lü L H, Zhao M, Zhao J R, Tao H B, Wang P. Canopy structure and photosynthesis of summer maize under different nitrogen fertilizer application rates. Sci Agric Sin, 2008, 41: 2624-2632 (in Chinese with English abstract) [本文引用:2]
[33]	赵国臣, 姜楠, 徐克章, 凌凤楼, 武志海, 邸玉婷. 吉林省1958—2005年间育成推广水稻品种部分叶片特征的变化. 作物学报, 2011, 37: 1101-1108 Zhao G C, Jiang N, Xu K Z, Ling F L, Wu Z H, Di Y T. Changes of some leaf characteristics in rice (Oryza sativa L. ) cultivars released from 1958 to 2005 in Jilin province. Acta Agron Sin, 2011, 37: 1101-1108 (in Chinese with English abstract) [本文引用:1]
[34]	王建波, 孙国荣, 陈刚, 曹文钟, 梁建生, 余政哲, 陆兆华. Na₂CO₃胁迫下星星草幼苗叶片PSII光能利用和耗散与培养基质渗透势的关系. 生态学报, 2006, 26: 115-121 Wang J B, Sun G R, Chen G, Cao W Z, Liang J S, Yu Z Z, Lu Z H. The relationship between light energy utilization and dissipation of PSII of Puccinellia tenuiflora seedlings and osmotic potential of culture solution under Na₂CO₃ stress. Acta Ecol Sin, 2006, 26: 115-121 (in Chinese with English abstract) [本文引用:1]
[35]	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 Fu J, Yang J C. Research advances in physiology of super rice under high-yield cultivation. Chin J Rice Sci, 2011, 25: 343-348 (in Chinese with English abstract) [本文引用:1]
[36]	Peng S B, Tang Q Y, Zou Y B. Current status and challenges of rice production in China. Plant Prod Sci, 2009, 12: 3-8 [本文引用:1]
[37]	Cheng S H, Zhuang J Y, Fan Y Y, Du J H, Cao L Y. Progress in research and development on hybrid rice: a super-domesticate in China. Ann Bot, 2007, 100: 959-966 [本文引用:1]
[38]	Kato T, Shinmura D, Taniguchi A. Activities of enzymes for sucrose-starch conversion in developing endosperm of rice and their association with grain filling in extra-heavy panicle types. Plant Prod Sci, 2007, 10: 442-450 [本文引用:1]

2014

0.0

赵黎明, 李明, 郑殿峰, 顾春梅, 那永光, 解保胜. 水稻光合作用研究进展及其影响因素分析. 北方水稻, 2014, 44(5): 66-71

Zhao L

, Li

, Zheng D

, Gu C

, Na Y

, Xie B

. Analysis of the factors and their effect on the photosynthesis of rice. North Rice, 2014, 44(5): 66-71 (in Chinese with English abstract)

摘　要：本文从两方面出发,一方面以光合作用为中心,详细的介绍了国内外关于水稻光合作用的研究进展;另一方面以光合作用的影响因素为中心,较全面地介绍了影响水稻光合作用的主要外界环境因素,并着重介绍了矿质养分和不同基因型对水稻光合作用的影响, 为今后如何提高水稻光合效率和利用率提供参考。

... 光合作用是植物生物产量和经济产量形成的基础, 作物产量的90%~95%直接或间接来自光合作用^[1,2] ...

2006

0.0

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

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

... 光合作用是植物生物产量和经济产量形成的基础, 作物产量的90%~95%直接或间接来自光合作用^[1,2] ...

... 的育种技术路线, 认为优良的植株形态可以实现水稻超高产的突破^[2,13,14,15] ...

1999

0.0

许大全. 光合速率、光合效率与作物产量. 生物学通报, 1999, 34(8): 8-10

Xu D

. Photosynthetic rate, photosynthetic efficiency and crop yield. Bull Biol, 1999, 34(8): 8-10 (in Chinese)

摘　要：简要论述了光合速率，光合效率与作物产量的关系及光合效率研究的重要性，预计到２０３０年，要满足我国１６亿人口的需求，粮食单产必须比现在增加５０％左右，为此，要充分挖掘作物的生产潜力，不断选育光合效率更高，产量更高的作物新品种，这需要作物光合效率调节控制理论研究上的突破，深入研究作物光合人用过程中光能转换和利用及耗散的分子机理与调控规律，确定光合效率改善的主要限制位点与相应对策，就可为通过其因工程定向

... 作物经济产量的高低是由光合生产力和呼吸消耗以及经济系数(又称收获指数)的大小决定的, 其中光合生产力又由光合速率、光合面积和光合功能期决定^[3,4] ...

2013

0.0

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.

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。

2005

0.0

... 叶片作为光合作用的主要器官直接影响作物的光能利用和干物质积累, 尤其是生育后期功能叶光合作用对作物产量起着至关重要的作用^[5,6,7] ...

... 有研究表明, 稻株顶部3叶的生长与穗分化同步, 与抽穗后的籽粒灌浆充实有着密切的关系, 因而被称为高效叶面积^[5,32] ...

2002

0.0

翟虎渠, 曹树青, 万建民, 陆巍, 张荣铣. 超高产杂交稻灌浆期光合功能与产量的关系. 中国科学(C辑), 2002, 32: 211-217

Zhai H

, Cao S

, Wan J

, Lu

, Zhang R

. The relationship between photosynthetic function and grain yield at filling stage of super-high-yield hybrid rice. Sci China (Ser C), 2002, 32: 211-217 (in Chinese)

以杂交水稻协优63为对照,对亩产800kg以上的超高产杂交水稻组合协优9308抽穗前、后的物质生产特性及剑叶光合作用与穗部物质积累的关系进行研究.结果表明:（i）与协优63 相比,协优9308抽穗前、后的物质生产能力无论生物量还是日干物质生产量,均极显著高于协优63,尤以抽穗后更为明显.（ii）协优9308剑叶的光合碳同化能力（叶源量）极显著高于协优63,且能够切合籽粒灌浆需求,在单株水平上其净同化产物基本满足籽粒灌浆的需求,而协优63抽穗后20d左右剑叶光合功能快速衰退,单株净同化产物不能满足籽粒灌浆的需求.结果证明,灌浆后期仍能保持高效光合功能、并且能够切合籽粒灌浆需求是实现水稻超高产的关键

... 叶片作为光合作用的主要器官直接影响作物的光能利用和干物质积累, 尤其是生育后期功能叶光合作用对作物产量起着至关重要的作用^[5,6,7] ...

2009

0.0

... 叶片作为光合作用的主要器官直接影响作物的光能利用和干物质积累, 尤其是生育后期功能叶光合作用对作物产量起着至关重要的作用^[5,6,7] ...

2010

0.0

... 叶绿素荧光作为光合作用研究的探针, 在植物逆境生理研究和光能利用探测等方面得到广泛应用^[8,9] ...

2010

0.0

程占慧, 刘良云. 基于叶绿素荧光发射光谱的光能利用率探测. 农业工程学报, 2010, 26(增刊-2): 74-80

Cheng Z

, Liu L

. Detection of vegetation light use efficiency based on chlorophyll fluorescence spectrum. Trans CSAE, 2010, 26(suppl-2): 74-80 (in Chinese with English abstract)

In this paper, a new method was presented to detect LUE based on the remotely sensed chlorophyll fluorescence (ChlF) signals. A diurnal change experiment was carried out to acquire the vegetation photosynthesis, canopy spectra and chlorophyll fluorescence parameters. The solar-induced chlorophyll fluorescence signals were separated from the canopy spectra at 688 and 760 nm based on the Fraunhofer Line Depth algorithm. The LUE model (fluo-LUE) was established based on the ChlF signals at the two oxygen absorption bands, and it was also compared with the LUE model (PRI-LUE) based on photochemical reflectance index (PRI). The result showed that LUE was highly significantly correlated with the PSII (photosystemⅡ) Yield of quantum efficiency(ΦPSII) (R2=0.802), which proved the possibility to detect LUE based on the remotely sensed ChlF signals. And LUE was significantly correlated with the ChlF signals at 688 and 760 nm. Compared with PRI-LUE model (R2=0.646), the correlation coefficient of fluo-LUE model was more significant (R2=0.799). Therefore, Chlf Signals could track the diurnal changes of LUE much better than the PRI. These research results show the solar-induced Chl signals might provide a prospective method for detecting LUE.

该文从光合作用光谱探测角度出发，研究基于自然光下植被叶绿素荧光光谱的植被光能利用率遥感探测的可行性。设计了日变化试验，获取植被光合、光谱及荧光参数，分析光能利用率的影响因素和光谱探测方法；基于叶绿素荧光夫琅和费（fraunhofer）线探测原理，计算植被冠层在2个氧吸收波段（688 nm和760 nm）叶绿素荧光发射光谱；并在此基础上建立了基于荧光发射光谱的光能利用率统计模型，并与基于光化学植被指数的光能利用率统计模型进行了对比。研究结果表明：PSII（photosystemⅡ，光系统II）光下量子产量与光能利用率显著正相关（R2=0.802），证实了通过叶绿素荧光来定量探测光能利用率是可行的；688 nm 和760 nm荧光均与植被光能利用率显著相关，相比光化学植被指数光能利用率统计模型（R2=0.646），基于荧光的光能利用率的统计模型精度更高，决定系数R2达到了0.799。因此，叶绿素荧光信号更能用于跟踪短期植被光能利用率变化。植被自然光下荧光光谱信号能够为光能利用率遥感探测提供一条新的途径。

... 叶绿素荧光作为光合作用研究的探针, 在植物逆境生理研究和光能利用探测等方面得到广泛应用^[8,9] ...

... 参照程占慧等^[9,26,27,28]方法计算: 光合作用效率(PE)=光合速率(P_n)/光合有效辐射(PAR) ...

2012

0.0

付景, 陈露, 黄钻华, 王志琴, 杨建昌. 超级稻叶片光合特性和根系生理性状与产量的关系. 作物学报, 2012, 38: 1264-1276

, Chen

, Huang Z

, Wang Z

, Yang J

. Relationship of leaf photosynthetic characteristics and root physiological traits with grain yield in super rice. Acta Agron Sin, 2012, 38: 1264-1276 (in Chinese with English abstract)

The success in super rice breeding has been considered as a great progress in rice production in China. This study aimed to understand the mechanism underlying the yield formation of super rice. Four super rice cultivars, Liangyoupeijiu and IIyou 084 ( indica hybrids), Huaidao 9 and Wujing 15 ( japonica ), and two elite check cultivars, Shanyou 63 ( indica hybrids) and Yangfujing 8 ( japonica ), were grown in field. Leaf photosynthetic characteristics and root physiological traits were investigated at different growth stages. The results showed that average total spikelet number and grain yield of the four super rice cultivars were 43.5% and 16.1%, respectively, more than those of two elite check cultivars, but the seed-setting rate of the four super rice cultivars was low 15.3% than that of the two elite check cultivars. The contents of chlorophyll a and b, total chlorophyll (a+b) and carotenoid, photosynthetic rate, transpiration rate, stomatal conductance of leaves, and specific root oxidation activity, root oxidation activity per plant, total root absorption area per plant, root active absorption area, specific surface area were higher in super rice than in elite check cultivars at earlier growth stages, while those traits decreased more quickly in super rice cultivars than in check cultivars at latter growth stages, and lower in super rice than in check cultivars up to 20 days after full heading. The results suggest that stronger leaf photosynthetic capacity and root activity during the early and mid-growth periods contribute to greater yield sink capacity of super rice, while the rapid decline of leaf photosynthetic capacity and root physiological activity at the late growth stage results in poor grain filling, and consequently, limits yield potential of super rice. Enhancing root physiological activity at late growth stage, especially during the grain filling period would be an important approach for super rice to further increase grain yield.

为阐明超级稻产量形成机理, 以4个超级稻品种[两优培九和II优084 (杂交籼稻)、淮稻9号和武粳15 (粳稻)]为材料, 2个高产品种[汕优63 (杂交籼稻)和扬辐粳8号(粳稻)]为对照, 观测了不同生育时期叶片光合性状和根系生理性状的变化特点。结果表明, 4个超级稻品种的平均总颖花量和产量较两个对照品种分别高出43.5%和16.1%, 但超级稻的结实率较对照品种低15.3个百分点。超级稻品种在生育前期叶片中叶绿素a含量、叶绿素b含量、总叶绿素含量、类胡萝卜素含量、光合速率、蒸腾速率、气孔导度和根系中单位干重根系活力、每株根系活力、总根系吸收面积、活跃吸收面积和比表面积均高于对照品种, 而在生育后期以上性状下降速率大于各自对照品种, 直至齐穗后20 d前以上性状指标均小于各自对照品种。说明超级稻强大的产量库容与其生育前中期较强的叶片光合能力和较好的根系生理性状密切相关, 生育后期叶片光合能力和根系生理活性下降快导致其结实率下降, 从而限制了其产量潜力的发挥。提高生育后期特别是结实后期根系生理活性是进一步提高超级稻产量的重要途径。

... 但有关叶片光合速率与作物产量的关系, 却有着不同的结果: 有的认为两者是正相关^[10,11], 也有的认为两者无相关性^[12] ...

... 以往有关叶片光合性能与作物产量的关系, 存在着不同的研究结论^[10,11,12] ...

2007

0.0

陈悦, 袁隆平, 王学华, 张道允, 陈娟, 邓启云, 赵炳然, 许大全. 超级杂交水稻谷粒产量与叶光合速率的关系. 植物生理与分子生物学学报, 2007, 33: 235-243

Chen

, Yuan L

, Wang X

, Zhang D

, Chen

, Deng Q

, Zhao B

, Xu D

. Relationship between grain yield and leaf photosynthetic rate in super hybrid rice. J Plant Physiol Mol Biol, 2007, 33: 235-243 (in English with Chinese abstract)

在2000～2005年期间,通过测定几种超级杂交水稻与普通杂交水稻'汕优63'的产量构成和叶片光合作用探讨了谷粒产量与光合作用的关系.结果表明:(1)4种超级杂交水稻'培矮64S/E32'、'P88S/0293'、'金23A/611'和'GD-1S/RB207'的产量水平显著高于'汕优63',是对照的108%～120%.(2)与'汕优63'相比,这些超级杂交水稻的株型好,上层叶片直立,穗大即每穗粒数多,是对照的125%～177%.(3)与'汕优63'相比,这些组合第二叶的净光合速率显著提高,但第一叶即剑叶的未必都较高.(4)去半叶处理降低了'GD-1S/RB207'的结实率,而去半穗处理显著提高了结实率.因此,这些超级杂交水稻的高产原因在于穗大、株型好以及群体光能利用效率高.增加单叶特别是剑叶的光合能力是克服谷粒产量的光合产物源限制和在未来的超级杂交水稻育种中实现产量潜力新突破的关键. Abstract： In order to explore the relationship between grain yield and photosynthesis, the yield composition and leaf photosynthetic rate in some super hybrid rices and ordinary hybrid rice 'Shanyou 63' as control were measured in 2000-2005. The results were as follows. (1) The yield levels of the four super hybrid rices, 'Pei'ai 64S/E32', 'P88S/0293', 'Jin23A/611' and 'GD-1S/RB207',were significantly higher, being 108%-120% of 'Shanyou 63'. (2) These super hybrid rices had a better plant type with more erect upper layer leaves and bigger panicles or more spikelets per panicle, being 125%-177% of 'Shanyou 63'. (3) Net photosynthetic rates of these super hybrid rices were significantly higher in the second leaf but not necessarily in the first leaf or flag leaf than those of 'Shanyou 63'. (4) The removal of half flag leaf led to a decline in the seed-setting rate, while the removal of half panicle induced its increase in 'GD-1S/RB207'. Hence, higher yield in these super hybrid rices can be attributed to their bigger panicles, better plant type and higher light use efficiency of their canopies. Raising the photosynthetic capacity of each leaf, especially flag leaf, is the key to overcome the photosynthate-source restriction on grain yield and to make a new breakthrough of yield potential in future breeding of super hybrid rice.

... 但有关叶片光合速率与作物产量的关系, 却有着不同的结果: 有的认为两者是正相关^[10,11], 也有的认为两者无相关性^[12] ...

... 以往有关叶片光合性能与作物产量的关系, 存在着不同的研究结论^[10,11,12] ...

2004

0.0

唐文邦, 陈立云, 肖应辉, 蔡义东, 兰海. 水稻功能叶形态及光合速率与产量构成因素的相关研究. 湖南农业科学, 2004, (2): 29-31

Tang W

, Chen L

, Xiao Y

, Cai Y

, Lan

. Relations of photosynthetic ability and functional leaves features to rice yield and yield components. Hunan Agric Sci, 2004, (2): 29-31 (in Chinese with English abstract)

为探索水稻功能叶光合速率及叶片性状与产量的关系,2002年对以玉米、高粱等外源DNA导入水稻的后代及其它水稻品种(组合)共17份材料的光合速率、功能叶的叶长、叶宽、叶面积与产量构成因素进行了相关分析.结果表明:功能叶的长度、宽度、叶面积及光合速率与水稻千粒质量呈负相关,但相关不显著.功能叶面积与光合速率及单穗质量呈显著正相关;光合速率与单穗质量呈正相关,但相关不显著.供试材料间的光合速率变幅较大,变异系数达23.1%.

... 但有关叶片光合速率与作物产量的关系, 却有着不同的结果: 有的认为两者是正相关^[10,11], 也有的认为两者无相关性^[12] ...

... 以往有关叶片光合性能与作物产量的关系, 存在着不同的研究结论^[10,11,12] ...

2013

0.0

王昆, 罗琼, 蔡庆红, 蒋敏明, 匡建业, 丁秋凡, 唐启源. 水稻株型的研究进展. 湖南农业科学, 2013, (17): 1-4

Wang

, Luo

, Cai Q

, Jiang M

, Kuang J

, Ding Q

, Tang Q

. Study progresses of rice plant type. Hunan Agric Sci, 2013, (17): 1-4 (in Chinese with English abstract)

综述了水稻株型的国内外研究现状及各构成因子包括根部、茎部、叶型及穗型性状的研究进展，指出了现阶段水稻株型研究中存在的主要问题，探讨了适宜湖南省稻区的理想株型，并提出应从株型育种和株型栽培调控等方面进行协同研究，这样才能达到形态与机能兼顾、理想株型与优势利用相结合，实现水稻超高产第三次突破。

... 的育种技术路线, 认为优良的植株形态可以实现水稻超高产的突破^[2,13,14,15] ...

2015

0.0

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. 作物学报, 2015, 41: 286-296

Jiang Y

, Xu

, Zhao

, Sun J

, Wei H

, Xu J

, Wei H

, Guo B

, Huo Z

, Dai Q

, Zhang H

. Canopy structure and photosynthetic characteristics of Yongyou series of indica- japonica hybrid rice under high-yielding cultivation condition. Acta Agron Sin, 2015, 41: 286-296 (in Chinese with English 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 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.

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MAD含量呈D>C>B>A趋势，SOD、POD、CAT酶活性呈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。

... 的育种技术路线, 认为优良的植株形态可以实现水稻超高产的突破^[2,13,14,15] ...

2008

0.0

... 的育种技术路线, 认为优良的植株形态可以实现水稻超高产的突破^[2,13,14,15] ...

2011

0.0

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

, Chen G

, Wang S

, Ding Y

. 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)

以江苏省近50年来在生产上大面积种植的10个代表性粳稻品种为材料,研究不同年代水稻产量形成、干物质生产、源库关系和株型的差异。结果表明:从20世纪60年代至90年代水稻单产显著提高,之后提高的幅度趋于平缓。在品种演进过程中,结实率、千粒重的遗传改良效果不显著,但每穗粒数显著增加,这是产量提高的关键。从20世纪60年代至80年代水稻的干物质量和收获指数同时增加,20世纪80年代以后收获指数略有下降,但干物质生产能力继续增加。品种演进使株型得到改良,顶3叶的叶长、叶角明显减小,穗部着粒密度、二次枝梗数增加,粒叶比也得到提高。从20世纪70年代至21世纪的品种株高明显增加,基部节间长度逐渐减小,直径逐渐增加。

... 前人从水稻品种改良^[16,22]、亚种间属性^[17,18]以及高产高效协同型品种^[19]等方面对水稻的株型特征进行了深入研究, 认为顶部3叶的着生角度和披垂度减小、有效叶面积和粒叶比提高等形态改良是水稻产量提高的重要原因, 这些结果也为水稻高产高效栽培及品种改良提供了参考依据 ...

2013

0.0

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

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)

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。

2012

0.0

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.

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。

1988

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梅方权, 吴宪章, 姚长溪, 李路平, 王磊, 陈秋云. 中国水稻种植区划. 中国水稻科学, 1988, 2: 97-110

Mei F

, Wu X

, Yao C

, Li L

, Wang

, Chen Q

. Rice cropping regionalization in China. Chin J Rice Sci, 1988, 2: 97-110 (in Chinese with English abstract)

According to the ecological environments, social and economic backgrounds and rice cropping characteristics in different areas of China, the Chinese rice planting area is divided into 6 rice cropping regions(the first-grade regions) and 16 subregions (the second-grade regions). Their general information, natural and ecological environments, paddy field cropping systems, types of rice varieties and their distribution and major diseases and insect pests are respectively described. Orientations, approaches and measures for developing rice production are raised and meanwhile the suggestions are put forward on the layout of production, paddy field crops combination and selection and construction of commercial rice production bases.

根据各地生态环境、社会经济条件和水稻种植特点,将中国稻区划分为6个稻作区(一级区)和16个稻作亚区(二级区)。对各个稻作区和亚区的基本情况、自然生态条件、稻田种植制度、品种类型、主要病虫害等作了评述,並提出水稻生产发展方向、途径和措施。

... 在各品种类型中, 以中籼稻所占比例最大^[20,21] ...

2003

0.0

郑景生, 黄育民. 中国稻作超高产的追求与实践. 分子植物育种, 2003, 1: 585-596

Zheng J

, Huang Y

. Thrust and practice of super high yielding rice production in China. Mol Plant Breed, 2003, 1: 585-596 (in Chinese with English abstract)

追求水稻高产再高产一直是中国人的夙愿.经过半个世纪长期不懈的努力,育种技术和栽培方法不断开拓创新,中国的水稻单产增长了2.37倍,居世界高产大国,总产增长3.13倍,占世界总产的35%,并取得了许多辉煌的科技成就.其中杂交水稻育种技术居世界领先水平,水稻最高单产屡屡突破世界纪录.本文列举了中国对水稻超高产执着追求的实例,回顾了中国水稻产量的提高历程,综述了水稻产量的理论极限及提高水稻产量的育种、栽培途径及高新技术的应用,展望了中国稻作未来的发展.

... 在各品种类型中, 以中籼稻所占比例最大^[20,21] ...

2006

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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 characteristic 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.

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。

... 以往虽然对中籼水稻品种产量与株型的演进特征、根系形态生理特性、氮素利用效率和米质变化等作了较多的研究^{[22,23,24,25]}, 但是关于中籼品种改良过程中叶片光合特性和主要荧光参数的变化特点及其与产量的关系, 缺乏深入研究 ...

2011

0.0

张耗, 黄钻华, 王静超, 王志琴, 杨建昌. 江苏中籼水稻品种演进过程中根系形态生理性状的变化及其与产量的关系. 作物学报, 2011, 37: 1020-1030

Zhang

, Huang Z

, Wang J

, Wang Z

, Yang J

. Changes in morphological and physiological traits of roots and their relationships with grain yield during the evolution of mid-season indica rice cultivars in Jiangsu Province. Acta Agron Sin, 2011, 37: 1020-1030 (in Chinese with English abstract)

Roots are involved in acquisition of nutrients and water, synthesis of plant hormones, and anchorage of plants. However, little is known what changes in root physiology and morphology during the evolution of rice cultivars. In this study, 13 typical mid-season indica rice cultivars (including hybrid combinations) applied in the production in Jiangsu Province during the last 60 years were used, which were classified into four types of early tall cultivars, dwarf cultivars, semi-dwarf cultivars, and super rice cultivars according to their application times, plant types and genotypes. All the tested cultivars were grown in the field and changes in morphological and physiological traits of roots and their relationships with grain yield during the evolution of cultivars were investigated. Results showed that the root dry weight, root weight density, root length, root length density and root diameter were increased with the evolution of the cultivars at main growth stages. From the heading stage, the shoot dry weight was increased with the evolution of the cultivars. Compared with other cultivars, the root-shoot ratio of super rice cultivars was increased at the mid-tillering and panicle initiation stages, but showed no significant differences among the four types of cultivars at latter growth stages. Specific root length was decreased with the evolution of the cultivars at the mid-tillering stage, and showed no significant differences among the four types of cultivars at other growth stages. The root oxidation activity, leaf photosynthetic rate, total absorbing surface area and active absorbing surface area of root, and the content of cytokinins (zeatin + zeatin riboside) in root bleeding and grain yield wereincreased 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 a large panicle. Regression analysis showed that the root dry weight, root length, root diameter, root oxidation activity, total absorbing surface area and active absorbing surface area of root very significantly correlated with grain yield. The results suggest that the improved root and shoot growth increases grain yield of the modern cultivars, especially super rice cultivars.

以江苏省近60年来各阶段具有代表性的13个中熟籼稻品种(含杂交稻组合)为材料，依据品种种植年代结合株型特点将其分为早期高秆(ET)、矮秆(DC)、半矮秆 (SDC)和超级稻(SR) 4个类型，研究了中籼水稻品种演进过程中根系形态生理性状的变化及其与产量的关系。结果表明，在各主要生育期，根干重、根重密度、根长、根长密度和根直径随品种演进增加或显著增加。自抽穗期，地上部干物重随品种演进显著增加。在分蘖中期和穗分化始期，超级稻品种的根冠比显著大于其他类型品种，在生育进程中，各类型间无显著差异。在分蘖中期，随品种演进，比根长显著降低，但在生育进程中，各类型间无显著差异。在生长早期和中期，根系氧化力、叶片光合速率、根系总吸收表面积和活跃吸收表面积以及根系伤流液中细胞分裂素(玉米素+玉米素核苷)浓度随品种演进增加或显著增加。随着品种演进产量逐步提高，其原因主要是每穗粒数的增多导致总颖花量的增加。回归分析表明，根干重、根长、根直径、根系氧化力、根系总吸收表面积和根系活跃吸收表面积与产量呈极显著线性正相关关系。说明改善的根系和地上部的生长，促进了现代品种特别是超级稻品种产量的提高。

2015

0.0

剧成欣, 陶进, 钱夕旸, 顾骏飞, 赵步洪, 杨凯鹏, 王志琴, 杨建昌. 不同年代中籼水稻品种的产量与氮肥利用效率. 作物学报, 2015, 41: 422-431

Ju C

, Tao

, Qian X

, Gu J

, Zhao B

, Yang K

, Wang Z

, Yang J

. Grain yield and nitrogen use efficiency of mid-season indica rice cultivars applied at different decades. Acta Agron Sin, 2015, 41: 422-431 (in Chinese with English abstract)

Improvement in rice cultivars plays an important role in increasing grain yield. However, little is known about the relationship between yield and nitrogen use efficiency in mid-season indica rice cultivar improvement process. In this study, 12 typical cultivars (including hybrid combinations) applied in the production in Jiangsu Province during the last 70 years were used, and classified into four types of 1940–1950s, 1960–1970s, 1980–1990s, and after 2000 (super rice) according to their application times. Three treatments of zero N (0N), 210 kg ha -1 N (MN) and 300 kg ha -1 N (HN) were designed. The results showed that grain yield and nitrogen use efficiency were progressively increased with the improvement of cultivars under each nitrogen rate. The super rice cultivars had a higher biomass and nitrogen accumulation, higher activities of root oxidation and nitrogen metabolic enzyme and higher leaf photosynthetic rate resulting in higher grain yield and nitrogen use efficiency when compared with any other types of cultivars. The root oxidation activity and photosynthetic characteristics of super rice were the peak at the heading stage, but sharp declined from heading to maturity, which could account for an important physiological reason for a lower filled-grain percentage of super rice. It would be an important approach to further increase grain yield of super rice through increasing root activity and leaf photosynthetic rate during grain filling.

本研究旨在探明中籼水稻在品种改良过程中产量与氮肥利用效率的变化特点。以江苏省近70年来不同年代在生产上广泛应用的12个代表性中籼水稻品种(含杂交稻组合)为材料，依据应用年代将其分为20世纪40—50年代、60—70年代、80—90年代和2000年以后4个类型，设置0 N (全生育期不施氮)、MN (全生育期施氮210 kg hm -2 ) 和HN (全生育期施氮300 kg hm -2 ) 3个施氮量处理，研究其产量、氮肥利用效率及其生理特性。结果表明，随品种应用年代的演进，不同年代中籼水稻品种的产量和氮肥利用效率均获得较大提高。2000年以后的品种(超级稻)产量和氮肥利用效率较高，根系性状和叶片光合特性以及氮代谢相关酶活性强是其重要生理基础。超级稻抽穗后根系氧化力和剑叶光合速率下降的幅度较大可能是导致超级稻结实率较低的一个重要原因。提高灌浆中后期超级稻的根系氧化力和剑叶光合速率，有望提高超级稻的结实率。

2009

0.0

张耗, 谈桂露, 孙小淋, 刘立军, 杨建昌. 江苏省中籼水稻品种演进过程中米质的变化. 作物学报, 2009, 35: 2037-2044

Zhang

, Tan G

, Sun X

, Liu L

, Yang J

. Changes in grain quality during the evolution of mid-season indica rice cultivars in Jiangsu Province. Acta Agron Sin, 2009, 35: 2037-2044 (in Chinese with English abstract)

Improvement in rice quality is very important in rice production. However, little is known about changes in rice quality during the evolution of cultivars. In this study, 13 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 five types of early tall cultivars, dwarf cultivars, semi-dwarf cultivars, semi-dwarf hybrids, and super rice according to their application times, plant types and genotypes. All the tested cultivars were grown in the field and the evolution characteristics of the quality and yield of rice were investigated. Results showed that the milling quality (brown rice rate, milled rice rate, and head milled rice rate) differed within the same type of cultivars, but showed no significant differences between the types. Compared with those of old cultivars (early tall and dwarf cultivars), the chalkiness, setback values and prolamin protein content in grains were significantly decreased, whereas the breakdown values and the contents of albumin and glutelin proteins, lysine, glycine and arginine were significantly increased in grains of modern cultivars (semi-dwarf hybrids, and super rice). 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. The results suggest that both quality and yield are improved during the evolution for mid-season indica rice cultivars.

以江苏省近 60 年来各阶段具有代表性的 13 个中熟籼稻品种 ( 含杂交稻组合 ) 为材料，依据品种种植年代结合株型特点将其分为早期高秆 (ET) 、矮秆 (DC) 、半矮秆常规稻 (SDC) 、半矮秆杂交稻 (SDH) 和超级稻 (SR) 5 个类型，研究了品种演进过程中米质的变化。结果表明，稻米的碾磨品质 ( 糙米率、精米率和整精米率 ) 在同一类品种间有较大差异，在品种演进过程中无明显变化。与早期品种相比，现代品种 ( 半矮秆杂交稻和超级稻 ) 稻米的垩白度显著降低，崩解值变大，消减值变小，稻米中的清蛋白、谷蛋白、赖氨酸、甘氨酸和精氨酸含量显著增加，醇溶蛋白含量显著降低。随着品种演进，产量逐步提高，其原因主要是总颖花量的增加及每穗粒数的显著增多。说明中籼稻品种演进过程中，米质和产量协同提高。

2008

0.0

冯大兰, 刘芸, 钟章成, 杨娟, 谢君. 三峡库区消落带芦苇(Phragmites communis (Reed))的光合生理响应和叶绿素荧光特性. 生态学报, 2008, 28: 2013-2021

Feng D

, Liu

, Zhong Z

, Yang

, Xie

. Photosynthesis and chlorophyll fluorescence parameters of the reed (Phragmites communis) grown in the hydro-fluctuation belt of Three Gorges Reservoir Area. Acta Ecol Sin, 2008, 28: 2013-2021 (in Chinese with English abstract)

After research on soil moisture contents of the hydro——fluctuation belt in Three Gorges Reservoir Area, three different types of water treatment were applied to the reed (Phragmites communis) to determine its photosynthetic characteristics and chlorophyll fluorescence parameters. The treatments T1 (with soil being complete submerged 2cm), T2 (with soil water content being 70 % to 100 % of field capacity), T3 (with soil water content 40% to 60 % of field capacity ) simulated the conditions in the hydro-fluctuation belt. The content of photosynthetic pigment, leaf gas exchange parameters, apparent resources use efficiency and chlorophyll fluorescence parameters of the reed (Phragmites communis) were clearly different for the three treatments. The T1-plants (i.e. the flooded plants) had the lowest content of photosynthetic pigment, while the T3-plants (i.e. the dry-treatment′ plants) had the highest carotenoids. The T1-plants had the lowest apparent light use efficiency(LUE), apparent CO2 use efficiency(CUE), net photosynthetic rate(Pn), electron transport rate(ETR) and lowest photochemical quenching (qP). However, compared to other plant species, known to have high tolerance to inundation, the flooded reed (Phragmites communis) had still a higher net photosynthetic rate(17.067 mol•m2•s 1). This indicates that reed also has a high tolerance to inundation. The T2- and T3-plants had, compared to the T1-plants, a high light use efficiency(LUE), CO2 use efficiency(CUE) and net photosynthetic rate(Pn), electron transport rate (ETR) and photochemical quenching (qP). The water use efficiency (WUE) of T3-plants was the highest, while that of the T2-plants was the lowest; In the T1-treatment the plants had the highest photochemical quantum efficiency of photo-system II (F′v /F′m ),while for the T2-treatment the highest photochemical quenching (qP) was found. The T3-plants had the highest electron transport rate (ETR) and non- photochemical quenching (NPQ), which indicated that under the high temperature and photosynthetical active radiation condition, those plants were able to protect photosynthetic structures by increasing respiration and thermal dissipiation, and hence maintaining a high photosynthetic rate（20.47μmol•m-2•s-1）.It was also demonstrated that the reed (Phragmites communis) could not only tolerate water submersion and wet condition but also endure drought.Therefore，The reed (Phragmites communis) could be a precedent species for restoration and rehabilitation of vegetation in the hydro-fluctuation belt of Three Gorges reservoir region.

模拟三峡库区消落带土壤含水量变化特征，设置了T1(淹水超过土壤表面2cm)、T2(土壤含水量为田间持水量的70%~100%)、T3(土壤含水量为田间持水量的40%~60%)3个不同处理组，用嘉陵江江水灌溉芦苇(Phragmites communis (reed))，研究了三峡库区不同消落带带位土壤不同含水量条件下芦苇穗期的光合生理生态响应机理和适应对策。结果表明，土壤不同含水量对芦苇植株光合色素、叶片气体交换参数、资源利用效率以及叶绿素荧光参数有不同影响。其中，T1的光合色素含量最低，T3的类胡萝卜素含量最高。在T1条件下，芦苇表现出较低的光能利用率(LUE)、CO2利用率(CUE)、净光合速率(Pn)、电子传递速率(ETR)和光化学荧光猝灭系数(qP)，但与其它耐水淹植物相比，T1条件下的穗期芦苇仍具有较高的光合速率(17067μmol•m-2•s-1)，说明芦苇具有较强的耐水淹能力。在T2和T3条件下，芦苇具有较高的光能利用率(LUE)、CO2利用率(CUE)、净光合速率(Pn)、电子传递速率(ETR)和光化学荧光猝灭系数(qP)。水分利用效率(WUE)大小顺序为T3>T1>T2。虽然T1的PSII有效光化学量子产量(F′v/F′m)最高，T2的光化学荧光猝灭系数(qP)最大，而T3的电子传递效率(ETR)和非光化学淬灭系数(NPQ)最高，表明在高光强和高温条件下，T3具有较强的热耗散能力，能有效保护光合机构，因而光合速率最高（20.47μmol•m-2•s-1）。研究证实芦苇不仅具有耐水湿的特点，还具有耐旱性，芦苇适合作为三峡库区消落带植被恢复建设禾本科先锋物种。

... 参照程占慧等^[9,26,27,28]方法计算: 光合作用效率(PE)=光合速率(P_n)/光合有效辐射(PAR) ...

2015

0.0

... 参照程占慧等^[9,26,27,28]方法计算: 光合作用效率(PE)=光合速率(P_n)/光合有效辐射(PAR) ...

... 较高的光合氮素利用效率不仅有利于提高产量, 而且还可以提高水稻氮肥利用效率^[27,28] ...

2014

0.0

... 参照程占慧等^[9,26,27,28]方法计算: 光合作用效率(PE)=光合速率(P_n)/光合有效辐射(PAR) ...

... 较高的光合氮素利用效率不仅有利于提高产量, 而且还可以提高水稻氮肥利用效率^[27,28] ...

2012

0.0

... , Lincoln, NE, USA)测定植株剑叶的光合速率和叶绿素荧光参数^[29,30] ...

2012

0.0

... , Lincoln, NE, USA)测定植株剑叶的光合速率和叶绿素荧光参数^[29,30] ...

2001

0.0

... 通常, 群体叶面积指数的增加会减少群体的透光率, 影响基部叶片的光合作用,进而影响根系活力^[31,32] ...

2008

0.0

吕丽华, 赵明, 赵久然, 陶洪斌, 王璞. 不同施氮量下夏玉米冠层结构及光合特性的变化. 中国农业科学, 2008, 41: 2624-2632

Lü L

, Zhao

, Zhao J

, Tao H

, Wang

. Canopy structure and photosynthesis of summer maize under different nitrogen fertilizer application rates. Sci Agric Sin, 2008, 41: 2624-2632 (in Chinese with English abstract)

【目的】试验通过氮肥调控,塑造高效冠层,确立不同夏玉米品种高效冠层与氮肥调控的定量化关系指标。【方法】设置4个施氮水平(0、90、180、270kg·ha-1),研究不同施肥量对夏播玉米CF008、郑单958、金海5号冠层结构和光合特性的影响。【结果】3个品种夏玉米在施氮量90或180kg·ha-1下冠层光合性能和产量较高。在灌浆前期,群体光合势(LAD)和叶面积指数(LAI)在施氮180或270kg·ha-1条件下显著较高,之后为90或180kg·ha- 1下较高,说明,高氮条件下生育前期冠层内透光率较低,冠层结构较不合理,导致生育后期叶片提早衰老,使后期光合性能和产量降低。【结论】由此得出在施氮量90或180kg·ha-1下构建的冠层较合理,其中冠层与氮肥调控的定量化关系指标为:群体透光率在吐丝期和灌浆中期分别达 12.56%～17.87%和13.13%～19.73%;最大LAI达5.76～6.75,成熟期仍然保持在2.09～3.68;吐丝期中上层叶片Pn 达35μmol·m2·s-1左右;吐丝后的群体LAD较高,郑单958和金海5号达210m2·d·m-2左右;产量达 10688～11461kg·ha-1。

... 通常, 群体叶面积指数的增加会减少群体的透光率, 影响基部叶片的光合作用,进而影响根系活力^[31,32] ...

... 有研究表明, 稻株顶部3叶的生长与穗分化同步, 与抽穗后的籽粒灌浆充实有着密切的关系, 因而被称为高效叶面积^[5,32] ...

2011

0.0

赵国臣, 姜楠, 徐克章, 凌凤楼, 武志海, 邸玉婷. 吉林省1958—2005年间育成推广水稻品种部分叶片特征的变化. 作物学报, 2011, 37: 1101-1108

Zhao G

, Jiang

, Xu K

, Ling F

, Wu Z

, Di Y

. Changes of some leaf characteristics in rice (Oryza sativa L. ) cultivars released from 1958 to 2005 in Jilin province. Acta Agron Sin, 2011, 37: 1101-1108 (in Chinese with English abstract)

In order to understand the changes of leaf characteristics associated with yield during the genetic improvement of rice, 33 japonica rice varieties released from 1958 to 2005 were planted under same field conditions. Some indexes related to leaf characteristics, such as leaf number, leaf area per plant, leaf area duration (LAD), length, width and area of top three leaves, chlorophyll content index (CCI) and specific leaf weight (SLW) of flag-leaf were measured at 10d after heading. The results showed that the grain yield was increased from 7 756.6 kg ha -1 in 1958 to 12 807.3 kg ha -1 in 2005, with the increase of 65.11% in 47 years and an annual increase rate of 1.39% according to the calculation of regression equation. The increasing of grain yield was mainly attributed to the increase of leaf number, leaf area per plant and LAD with years of release. By studying on the top three leaves of plant, our results showed that the significant decrease of flag-leaf length resulted in the decline of flag-leaf area; the changes of top second leaf area was not obvious; as the increase in the length and width of top 3rd leaf, significantly increased was found in its area with years of release. Leaf number, leaf area per plant and LAD were all positive correlated with years of release significantly. CCI and SLW of flag-leaf were positively correlated with year of release and with yield significantly. Our results showed that the increase of plantleaf area and LAD were resulted from the increase ofleaf number and the area of top 3rd leaf. CCI and SLW of flag-leaf were increased with years of release. We suggested that leaf number, leaf area per plant, LAD, and CCI as well as SLW of flag-leaf at 10d after heading could be as potential indices in the future high yield rice breeding.

1 Agronomy College, Jilin Agricultural University, Changchun 130118, China; 2Institute of Rice Research, Jilin Academy of Agricultural Sciences, Changchun 130124, China

以吉林省1958—2005年间育成并推广的33个水稻品种为材料，研究了植株抽穗后10 d叶片数量、叶面积、光合势(LAD)、上三叶叶片的长度、宽度和叶面积以及剑叶叶绿素含量(CCI)和比叶重(SLW)的变化及其与产量的关系。结果表明，吉林省47年来水稻品种的平均产量由7 756.6 kg hm -2 提高到12 807.3 kg hm -2 ，增加了65.11%，平均每年增加1.39%。随着水稻品种产量的提高，单株叶面积和LAD增加，并与育成年代和产量呈显著正相关。单株叶面积和LAD的增加主要是植株叶片数量和倒三叶叶面积显著增加的结果；倒二叶叶面积变化不大；剑叶长度显著变短，面积呈下降趋势，但剑叶叶绿素含量和SLW与育成年代和产量呈显著正相关。本文结果表明，吉林省水稻品种产量的遗传改良主要是通过增加抽穗后植株叶片数量和倒三叶叶面积，提高了单株叶面积和LAD。水稻抽穗后植株叶片数量、单株叶面积、LAD、CCI值和SLW可以作为高产品种的参考指标。

2006

0.0

王建波, 孙国荣, 陈刚, 曹文钟, 梁建生, 余政哲, 陆兆华. Na₂CO₃胁迫下星星草幼苗叶片PSII光能利用和耗散与培养基质渗透势的关系. 生态学报, 2006, 26: 115-121

Wang J

, Sun G

, Chen

, Cao W

, Liang J

, Yu Z

, Lu Z

. The relationship between light energy utilization and dissipation of PSII of Puccinellia tenuiflora seedlings and osmotic potential of culture solution under Na₂CO₃ stress. Acta Ecol Sin, 2006, 26: 115-121 (in Chinese with English abstract)

Method of fluorescence induction kinetics was used to investigate the relationship between light energy utilization and dissipation of PSⅡ of Puccinellia tenuiflora seedlings and osmotic potential of culture solution under Na 2 CO 3 stress. It was found that the changes of maximal photochemical efficiency of PSⅡ ( Fv/Fm ), potential photochemical efficiency ( Fv/Fo ), actual photochemical eff iciency of PSⅡ in the light (ΦPSⅡ) and photochemical efficiency of PSⅡ in the light ( Fv′/Fm ′) were not obvious in an osmotic potential higher than -4 bar. At the same time, Fv/Fm, Fv/Fo, Fv′/Fm ′ increased along with the osmotic potential lower than -4 bar; nevertheless　ΦPSⅡ, electron transfer rate (ETR), photochemical efficiency, heat dissipation and heat dissipation rate decreased. The results showed that the dissipation mechanism of surplus of light energy of P. tenuiflora seedlings may be various in different osmotic potential induced by Na2CO3 stress (lower than -4 bar and higher than -4 bar), and there may be a refined osmotic regulation mechanism under osmotic potential higher than -4 bar. Furthermore, in high osmotic stress induced by Na2CO3, the safeguard mechanism of P. tenuiflora was different from that of other plants, they dissipated surplus light energy via two approach: increasing light energy absorbed by light-harvesting pigment that used in heat dissipation and heat dissipation rate; increasing ΦPSⅡ, ETR and photochemical efficiency, and building up fake circulate photosynthetic phosphorylation, which resulted in the augment of active oxygen eliminated by antioxidant enzymes system. In this way do P. tenuiflora protect photosynthetic apparatus from the damage of surplus of light energy.

用荧光动力学的方法研究了碱性盐Na 2 CO 3 胁迫下星星草幼苗叶片PSⅡ光能利用和耗散与培养基质渗透势的关系。结果发现在大于-4bar的胁迫下，PSⅡ最大光化学效率（Fv/Fm）、PSⅡ潜在光化学效率（ Fv/Fo ）、PSⅡ实际光化学效率（ΦPSⅡ）以及开放的PSⅡ反应中心有效光化学效率（ Fv′/Fm′ ）的变化不大；然而在小于-4bar Na 2 CO 3 胁迫下， Fv/Fm、Fv/Fo 和 Fv′/Fm ′均随着渗透势的增大而增大，而ΦPSⅡ、电子传递速率（ETR）、光化学速率、捕光色素吸收的光能被用于热耗散的相对份额及热耗散速率则随着渗透势的增大而减小。这些研究结果说明星星草幼苗在Na 2 CO 3 胁迫所导致的不同的渗透胁迫下（小于-4bar和大于-4bar）其过剩光能的耗散机制可能不同，大于-4bar的胁迫下可能存在精细的渗透调节机制，而在高强度的Na2CO3所导致的渗透胁迫下具有与其它植物不同的保护机制，可能通过两条途径耗散过剩的光能，一方面通过增加捕光色素吸收的光能被用于热耗散的相对份额及热耗散速率；另一方面通过增大ΦPSⅡ、光化学速率、ETR，增强假循环式光合磷酸化过程，而由此引起的活性氧的增加则通过体内较高活性的保护酶系统来清除，以保护光合器官免受过剩光能的损伤。

... q_N则反映了PSII反应中心对天线色素吸收过量光能后的热耗散能力及光合机构的损伤程度^[34] ...

2011

0.0

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-yield 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.

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。

... 1990s的品种相比, 超级稻的结实率较低, 这已成为限制其产量潜力发挥的重要因子^{[35,36,37,38]} ...

2009

0.0

... 1990s的品种相比, 超级稻的结实率较低, 这已成为限制其产量潜力发挥的重要因子^{[35,36,37,38]} ...

2007

0.0

... 1990s的品种相比, 超级稻的结实率较低, 这已成为限制其产量潜力发挥的重要因子^{[35,36,37,38]} ...

2007

0.0

... 1990s的品种相比, 超级稻的结实率较低, 这已成为限制其产量潜力发挥的重要因子^{[35,36,37,38]} ...