灌浆结实期半深水灌溉对水稻产量、干物质转运及籽粒灌浆的影响
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Influence of Semi-Deep Water Irrigation during Grain-Filling Period on Rice Yield, Dry Matter Transportation and Grain-Filling
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通讯作者:
收稿日期: 2025-12-12 修回日期: 2026-03-25 网络出版日期: 2026-04-10
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Received: 2025-12-12 Revised: 2026-03-25 Online: 2026-04-10
作者简介 About authors
郭威,主要从事水稻栽培研究,E-mail:
近年来,稻渔综合种养扩增迅速,成为一种重要的水稻种植制度。不少稻渔综合种养模式需在水稻种植的一定时期内形成半深水环境以满足水产动物的生长活动需求,这与水稻单作常规浅湿灌溉方式存在较大差异。以粳型常规水稻南粳5718和籼型两系杂交水稻徽两优898为试验材料,设置3种灌溉模式,分别为常规高产栽培浅湿灌溉模式(CK)、抽穗期至抽穗后40 d实施20 cm(WD20)和40 cm深的灌溉(WD40),研究了结实期不同灌溉深度对水稻产量、干物质积累与转运及籽粒灌浆的影响。结果表明,与CK相比,WD20和WD40处理造成2个试验品种结实率和千粒重均显著下降,导致水稻减产,且WD40的减产幅度明显大于WD20处理,徽两优898的产量损失大于南粳5718。经分析发现,与CK相比,半深水灌溉下水稻营养器官呈现出明显的“滞绿”现象,叶片衰老变慢,但光合作用生产的光合物质无法被充分转运利用,茎鞘干物质转运和籽粒灌浆明显差于CK处理,收获指数呈显著下降趋势。半深水灌溉下水稻强势粒和弱势粒的灌浆速率明显降低,导致水稻粒重显著下降,且弱势粒灌浆受半深水灌溉的影响更大。
关键词:
In recent years, the rapid expansion of integrated rice-fish farming has become an important rice planting system. Many integrated rice-fish farming systems need to form a semi-deep water environment during a certain period of rice planting to meet the needs of aquatic animal growth activities, which is quite different from the conventional shallow wet irrigation method of rice monoculture. In this study, japonica conventional rice Nangeng 5718 (NG5718) and indica two-line hybrid rice Huiliangyou 898 (HLY898) were used as experimental materials, and three irrigation modes were conducted: conventional shallow wet irrigation (CK); 20 (WD20) and 40 cm (WD40) irrigation depth from heading stage to 40 days after heading. This research investigated the effects of different irrigation depths during grain period on rice yield, dry matter translocation and accumulation, and grain filling. The results showed that, compared with CK, WD20 and WD40 treatments significantly decreased the seed-setting rate and 1000-grain weight of the two test varieties, resulting in a decrease in rice yield. The yield reduction effect of WD40 was significantly greater than that of WD20 treatment, and the yield loss of Huiliangyou 898 was greater than that of Nanjing 5718. The analysis showed that, compared with the CK, the vegetative organs of rice under semi-deep water irrigation showed obvious 'stay green' phenomenon. The leaf senescence slowed down but the photosynthetic substances could not be fully transported and utilized. The dry matter transport and grain filling of stem and sheath were significantly worse than those of CK treatment, and the harvest index showed a significant downward trend. The grain filling rate of superior and inferior grains of rice under semi-deep water irrigation was significantly reduced, resulting in a significant decrease in grain weight, and the grain filling of inferior grains was more severely affected.
Keywords:
本文引用格式
郭威, 孙冬, 闫治霖, 丁蓓, 杜跃辰, 吴慧扬, 徐强, 窦志, 高辉.
Guo Wei, Sun Dong, Yan Zhilin, Ding Bei, Du Yuechen, Wu Huiyang, Xu Qiang, Dou Zhi, Gao Hui.
2024年,我国稻渔综合种养面积达307万hm2[1],约占全国水稻种植总面积的10%,表明稻渔综合种养已成为我国主要稻作制度之一。稻渔综合种养与水稻单作的关键环境差异在于,为满足鱼、虾、蟹和鳖等水产品大规格、高品质生长的需求,在水稻特定生长阶段需建立并维持较深水层,一般灌溉深度在20~40 cm[2-3],既高于常规稻田淹灌水层深度(3~6 cm),但又明显低于东南亚所谓“深水稻”的淹水深度(50 cm以上)[4],因此本文暂将其定义为“半深水灌溉”。例如,在长江流域的稻虾共作模式中,待水稻返青、植株增高后,田间逐步建立15 cm以上水层并投放克氏原螯虾苗种,最终维持在20~40 cm水层以满足虾类生长需求[5-6]。再如,浙江地区的稻鱼共作模式中,水稻有效分蘖期后稻田水层会逐渐加深并长期保持在20~35 cm[7],为田鱼生长活动提供适宜水深,直至乳熟期才排水捕鱼。
1 材料与方法
1.1 试验地概况
试验于2023年在安徽省天长市永丰镇稻渔综合种养创新试验基地进行。试验地点位于高邮湖西岸,属季风性湿润气候区,光照充足,雨水充沛。试验田土壤为黏土,其基本理化性质为pH 6.0、有机质27.5 g/kg、全氮1.83 g/kg、有效磷17.8 mg/kg、速效钾123 mg/kg。
1.2 试验设计
试验在专门为稻渔综合种养试验建设的水泥池小区里进行。每个水泥池长10.0 m、宽5.0 m、深0.5 m,可实现不同灌溉处理方式。供试水稻品种为粳型常规水稻南粳5718和籼型两系杂交水稻徽两优898。5月15日采用塑料软盘育秧,6月16日人工栽插,南粳5718栽插行株距为30 cm×13 cm,每穴栽3苗,徽两优898栽插行株距为30 cm×17 cm,每穴栽2苗。试验设置3种灌溉模式,分别为常规高产栽培浅湿灌溉模式(CK);结实期20 cm水层灌溉,即自水稻抽穗至抽穗后40 d田间保持20 cm水层(WD20);结实期40 cm水层灌溉,即自水稻抽穗至抽穗后40 d田间保持40 cm水层(WD40)。3个处理具体水分管理方式见表1。每个处理3次重复,采用随机区组排列。
表1 各处理全生育期水分管理方式
Table 1
| 生育时期Growth period | CK | WD20 | WD40 |
|---|---|---|---|
| 缓苗期Seedling recovering | 0~2 cm水层 | 0~2 cm水层 | 0~2 cm水层 |
| 分蘖期Tillering | 2~4 cm水层 | 2~4 cm水层 | 2~4 cm水层 |
| 搁田期Field drying | 排干水,搁至土表有鸡爪纹 | 排干水,搁至土表有鸡爪纹 | 排干水,搁至土表有鸡爪纹 |
| 拔节孕穗期Jointing and booting | 浅湿交替 | 浅湿交替 | 浅湿交替 |
| 抽穗期-抽穗后40 d Heading - 40 days after heading | 浅湿交替 | 保持20 cm水层 | 保持40 cm水层 |
| 抽穗后40 d-成熟期40 days after heading - maturity | 浅湿交替 | 浅湿交替 | 浅湿交替 |
| 成熟期Maturity | 收获前7 d左右断水 | 收获前7 d左右断水 | 收获前7 d左右断水 |
同一品种不同灌溉处理下的养分管理方案保持一致。南粳5718总施氮量为300 kg/hm2,徽两优898总施氮量225 kg/hm2,2个品种氮肥施用比例关系均为基肥:分蘖肥:穗肥=4:3:3,磷钾肥的施用量均按照N:P2O5:K2O=1:0.5:0.8的比例换算,磷肥一次性基施,钾肥分基肥和穗肥等量施用,基肥于移栽前2 d施用,分蘖肥于移栽后第7天施用,穗肥于水稻基部第一节间定长、第二节间伸长1~2 cm时施用。水稻病虫害防控参照当地稻渔综合种养常规方法进行,采用人工拔草方式控除杂草。
1.3 测定项目与方法
1.3.1 产量及其构成因素
于水稻成熟期每小区选取连续30穴植株,普查有效穗数,按照平均穗数取代表性3穴,考查每穗颖花数、结实率和千粒重,计算理论产量。每个小区另取50穴水稻植株进行收割、脱粒、称重,用于计算实际产量。
1.3.2 干物质积累与转运相关指标
各小区分别于抽穗期和成熟期取样,每次每小区取长势具有代表性的水稻植株3穴,分成茎鞘、叶和穗,在105 ℃下杀青30 min后,在80 ℃下烘干至恒重,称重。
相关指标计算方法[15]:
成熟期单茎地上部干重=成熟期单茎茎鞘干重+成熟期单茎叶干重+成熟期单茎穗干重;
抽穗―成熟期单茎穗干重积累量=成熟期单茎穗干重-抽穗期单茎穗干重;
抽穗―成熟期单茎茎鞘干物质输出量=抽穗期单茎茎鞘干重-成熟期单茎茎鞘干重;
茎鞘干物质输出率(%)=抽穗―成熟期单茎茎鞘干物质输出量/抽穗期单茎茎鞘干重×100;
茎鞘干物质贡献率(%)=抽穗―成熟期单茎茎鞘干物质输出量/抽穗―成熟期单茎穗干重积累量×100;
收获指数(%)=成熟期单茎穗干重/成熟期单茎地上部干重×100。
1.3.3 水稻籽粒灌浆相关指标
各小区于抽穗期一次标记同日始花、生长整齐的穗子250个。自抽穗期开始,每10 d每小区随机取标记穗20个。从田间取回样本后,取直接着生于穗顶部4个一次枝梗上的颖花,作为强势粒样本,取穗基部4个二次枝梗上的颖花,作为弱势粒样本。取样结束后,先在105 ℃下烘30 min杀青,再于75 ℃下连续烘60 h,测定并计算粒重。使用Richards方程拟合籽粒增重过程,参照朱庆森等[16]方法由拟合方程导出一系列一级和次级参数,对不同处理水稻籽粒灌浆过程进行生长分析。
Richards方程:
式中,W为各期生长量,即千粒重,g;t为齐穗后天数,d;A、B、K、N为方程参数,并用决定系数(r2)(W依t的回归平方和占总平方和的比率)表示其配合适度。
各次级参数计算公式如下:
式中,R0为起始生长势,g;Tmax为达到最大灌浆速率的时间,d;GRmean为平均灌浆速率,mg/d;GRmax为最大灌浆速率,mg/d;D为活跃灌浆期,d。
1.3.4 剑叶光合参数、SPAD值
于水稻抽穗后15(D15)和35 d(D35),在各小区选择8株长势一致的主茎,用SPAD-502叶绿素仪(Minolta,日本)测定水稻剑叶上、中、下部的叶绿素相对含量(SPAD值),取平均值。
在抽穗后15和35 d各小区选择5株长势一致的水稻主茎植株,用Li-6400便携式光合仪(Li- Cor,美国)测定剑叶的净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和胞间CO2浓度(Ci)。
1.4 数据处理
使用Microsoft Excel 2019处理数据和制作表格,运用IBM SPSS Statistics 23进行统计分析,根据研究设计和数据特点采用单因素方差分析并用LSD法进行多重比较,使用Origin 2021b作图。
2 结果与分析
2.1 不同灌溉深度对水稻产量及其构成因素的影响
方差分析结果(表2)表明,品种和灌溉深度对每穗颖花数、结实率、千粒重、理论产量、实际产量和收获指数均有显著或极显著影响,同时品种和灌溉深度的互作对所有产量及其构成因素均无显著影响。
表2 结实期不同灌溉深度对水稻产量及收获指数的影响
Table 2
| 品种 Variety | 处理 Treatment | 单位面积有效穗数 Effective panicles per unit area (×104/ hm2) | 每穗颖花数 Spikelets per panicle | 结实率 Seed-setting rate (%) | 千粒重 1000-grain weight (g) | 理论产量 Theoretical yield (t/hm2) | 实际产量 Actual yield (t/hm2) | 收获指数 Harvest index (%) |
|---|---|---|---|---|---|---|---|---|
| 南粳5718 NG5718 | CK | 316.30a | 168.86a | 91.91a | 26.44a | 12.25a | 12.19a | 58.71a |
| WD20 | 311.60a | 159.92ab | 88.60b | 25.54b | 11.02b | 10.98b | 56.93ab | |
| WD40 | 309.13a | 153.73b | 83.22c | 25.18c | 9.41c | 9.31c | 52.86b | |
| 徽两优898 HLY898 | CK | 234.64a | 242.98a | 88.73a | 25.80a | 12.08a | 12.39a | 60.33a |
| WD20 | 232.22a | 234.78ab | 84.19b | 24.42b | 10.42b | 10.63b | 58.16ab | |
| WD40 | 234.96a | 223.82b | 80.07c | 23.87c | 9.12c | 9.05c | 53.58b | |
| 方差分析 Analysis of variance | V | 2453.09** | 2723.53** | 90.91** | 46.41** | 8.63* | 28.82** | 23.28** |
| ID | 2.16 | 82.01** | 178.20** | 39.41** | 186.48** | 33.34** | 104.70** | |
| V×ID | 1.96 | 2.99 | 1.21 | 1.71 | 1.09 | 0.05 | 2.27 |
同一列不同小写字母表示在P < 0.05水平上差异显著。“*”和“**”分别表示在P < 0.05和P < 0.01水平上差异显著。V表示品种,ID表示灌溉深度,V×ID表示品种与灌溉深度互作。下同。
Different lowercase letters in the same column indicate significant differences at P < 0.05 level.“*”and“**”indicate significant differences at P < 0.05 and P < 0.01 levels, respectively. V represents variety, ID represents irrigation depth, V×ID represents the interaction between variety and irrigation depth. The same below.
由表2可知,2个品种不同处理下的理论产量均表现出CK>WD20>WD40的趋势。对水稻产量构成因素分析发现,2个品种单位面积有效穗数在不同处理间无显著差异。与CK处理相比,WD20处理下南粳5718的每穗颖花数、结实率和千粒重分别下降了5.3%、3.6%和3.4%,并减产了10.1%,WD40处理每穗颖花数、结实率和千粒重分别下降了10.1%、9.6%和4.8%,同时理论产量下降了23.2%。与CK相比,WD20处理使徽两优898每穗颖花数、结实率和千粒重分别下降了5.1%、5.1%和5.3%,理论产量下降了13.7%;WD40处理使徽两优898每穗颖花数、结实率和千粒重分别下降了10.4%、9.8%和7.5%,理论产量下降了24.5%。与CK处理相比,WD20和WD40处理使南粳5718的实际产量分别下降了9.9%和23.6%,使徽两优898的实际产量分别下降了14.2%和26.9%,这一结果与理论产量变化趋势完全一致。与CK相比,2个品种的收获指数在WD20和WD40处理均表现出下降趋势,且收获指数在CK和WD40之间差异达到显著水平。
2.2 不同灌溉深度对水稻干物质积累与转运的影响
方差分析的结果(表3)表明,品种对不同时期水稻各器官干物质重均有极显著影响,灌溉深度对除水稻抽穗期单茎茎鞘干重、单茎叶干重和单茎穗干重以外的干物质积累与转运的各项指标均具有极显著影响,两因素互作仅对成熟期单茎穗干重和成熟期单茎地上部干重有显著影响。
表3 结实期不同灌溉深度对水稻干物质积累与转运的影响
Table 3
| 品种 Variety | 处理 Treatment | 抽穗期 单茎茎鞘干重 SDWPS-H (g) | 抽穗期 单茎叶干重 LDWPS-H (g) | 抽穗期 单茎穗干重 PDWPS-H (g) | 抽穗期 单茎地上部干重 ADWPS-H (g) | 成熟期 单茎茎鞘干重 SDWPS-M (g) | 成熟期 单茎叶干重 LDWPS-M (g) |
|---|---|---|---|---|---|---|---|
| 南粳5718 NG5718 | CK | 3.27a | 1.17a | 0.56a | 5.00a | 2.56b | 0.49b |
| WD20 | 3.28a | 1.07a | 0.58a | 4.93a | 2.65b | 0.55ab | |
| WD40 | 3.24a | 1.05a | 0.57a | 4.86a | 2.81a | 0.60a | |
| 徽两优898 HLY898 | CK | 3.88a | 0.93a | 0.81a | 5.62a | 3.03b | 0.59b |
| WD20 | 3.92a | 0.94a | 0.79a | 5.65a | 3.10b | 0.64b | |
| WD40 | 3.86a | 0.93a | 0.83a | 5.60a | 3.35a | 0.74a | |
| 方差分析 Analysis of variance | V | 87.38** | 35.97** | 161.88** | 16.32* | 62.42** | 3.68** |
| ID | 0.83 | 0.82 | 0.45 | 0.47 | 24.91** | 2.64** | |
| V×ID | 0.12 | 0.77 | 1.36 | 0.38 | 0.05 | 3.71 | |
| 品种 Variety | 处理 Treatment | 成熟期 单茎穗干重 PDWPS-M (g) | 成熟期 单茎地上部干重 ADWPS-M (g) | 抽穗―成熟期 单茎穗干重积累量 PDWAPS (g) | 抽穗―成熟期 单茎茎鞘干物质输出量 SDMOPS (g) | 茎鞘干物 质转运率 SDMTR (%) | 茎鞘干物 质贡献率 SDMCR (%) |
| 南粳5718 NG5718 | CK | 4.34a | 7.39a | 3.78a | 0.71a | 21.80a | 21.81a |
| WD20 | 4.08b | 7.29a | 3.49ab | 0.62a | 19.04b | 17.84a | |
| WD40 | 3.82c | 7.22a | 3.25b | 0.42b | 13.15c | 13.10b | |
| 徽两优898 HLY898 | CK | 5.82a | 9.49a | 5.01a | 0.85a | 21.91a | 17.01a |
| WD20 | 5.39b | 9.27a | 4.60b | 0.78a | 19.81b | 16.89a | |
| WD40 | 4.52c | 8.44b | 3.69c | 0.53b | 13.66c | 14.29b | |
| 方差分析 Analysis of variance | V | 162.47** | 266.42** | 74.39** | 30.98** | 1.93 | 2.29 |
| ID | 73.18** | 28.87** | 60.01** | 172.96** | 217.86** | 58.08** | |
| V×ID | 12.34* | 15.40* | 10.61** | 0.52 | 0.31 | 6.67* |
SDWPS-H:抽穗期单茎茎鞘干重;LDWPS-H:抽穗期单茎叶干重;PDWPS-H:抽穗期单茎穗干重;ADWPS-H:抽穗期单茎地上部干重;SDWPS-M:成熟期单茎茎鞘干重;LDWPS-M:成熟期单茎叶干重;PDWPS-M:成熟期单茎穗干重;ADWPS-M:成熟期单茎地上部干重;PDWAPS:抽穗―成熟期单茎穗干重积累量;SDMOPS:抽穗―成熟期单茎茎鞘干物质输出量;SDMTR:茎鞘干物质转运率;SDMCR:茎鞘干物质贡献率。
SDWPS-H: stem-sheath dry weight per stem at heading; LDWPS-H: leaf dry weight per stem at heading; PDWPS-H: panicle dry weight per stem at heading; ADWPS-H: aboveground dry weight per stem at heading; SDWPS-M: stem-sheath dry weight per stem at maturity; LDWPS-M: leaf dry weight per stem at maturity; PDWPS-M: panicle dry weight per stem at maturity; ADWPS-M: aboveground dry weight per stem at maturity; PDWAPS: panicle dry weight accumulation per stem at heading-maturity; SDMOPS: stem-sheath dry matter output per stem at heading-maturity; SDMTR: translocation rate of stem-sheath dry matter; SDMCR: contribution rate of stem-sheath dry matter.
由表3可知,2个品种各处理下抽穗期单茎茎鞘干重和单茎叶干重差异不显著,而成熟期单茎地上部干重和抽穗―成熟期单茎穗干重积累量在CK和WD40处理之间均存在显著差异,且均表现出CK>WD20>WD40的趋势。与CK处理相比,南粳5718在WD20处理下成熟期单茎穗干重和抽穗―成熟期单茎穗干重积累量分别降低了6.0%和7.7%,WD40处理下则分别下降了11.2%和14.0%;徽两优898在WD20处理下成熟期单茎穗干重和抽穗―成熟期单茎穗干重积累量分别下降了7.4%和8.2%,WD40处理下则分别下降了22.3%和26.4%。
不同品种水稻各处理的抽穗―成熟期单茎茎鞘干物质输出量、茎鞘干物质转运率和茎鞘干物质贡献率均呈现为CK>WD20>WD40,且任意2个处理之间茎鞘干物质转运率的差异均达到显著水平;抽穗―成熟期单茎茎鞘干物质输出量和茎鞘干物质贡献率在CK与WD40处理之间、WD20与WD40处理之间均存在显著差异,而在CK与WD20处理之间无显著差异。与CK处理相比,南粳5718在WD20处理下茎鞘干物质转运率和贡献率分别下降了12.7%和18.2%,WD40处理下则分别下降了39.7%和40.0%;徽两优898在WD20处理下茎鞘干物质转运率和贡献率分别下降了9.6%和0.7%,WD40处理分别下降了37.7%和15.9%。
2.3 不同灌溉浓度对水稻籽粒灌浆相关指标的影响
对结实期间粒重变化用Richards方程进行了拟合,参数估计值和决定系数见表4。强势粒和弱势粒的拟合度均在0.9900以上,说明不同粒位的籽粒灌浆过程均可用Richards模型描述。2个品种各处理强、弱势粒方程形状参数N均大于1.00,说明速率曲线右偏,强势粒的N值表现为CK>WD20>WD40,而弱势粒则相反。
表4 籽粒灌浆过程的Richards方程参数估计值
Table 4
| 品种 Variety | 处理 Treatment | 粒位 Grain position | A | B | K | N | 生长量 W(g) | 标准差 S | 决定系数 r2 |
|---|---|---|---|---|---|---|---|---|---|
| 南粳5718 NG5718 | CK | 强势粒 | 28.75 | 75.89 | 0.25 | 2.05 | 28.74 | 0.57 | 0.9994 |
| 弱势粒 | 23.79 | 542.92 | 0.25 | 2.86 | 23.74 | 0.56 | 0.9992 | ||
| WD20 | 强势粒 | 26.48 | 54.17 | 0.25 | 1.80 | 26.47 | 1.33 | 0.9964 | |
| 弱势粒 | 21.53 | 2404.84 | 0.27 | 3.94 | 21.45 | 0.44 | 0.9993 | ||
| WD40 | 强势粒 | 24.76 | 14.34 | 0.18 | 1.33 | 24.69 | 0.96 | 0.9976 | |
| 弱势粒 | 19.91 | 5440.38 | 0.24 | 4.03 | 19.31 | 1.45 | 0.9917 | ||
| 徽两优898 HLY898 | CK | 强势粒 | 29.46 | 129.16 | 0.27 | 2.87 | 29.45 | 0.9 | 0.9983 |
| 弱势粒 | 25.42 | 9970.80 | 0.30 | 4.67 | 25.36 | 1.36 | 0.9958 | ||
| WD20 | 强势粒 | 27.59 | 20.10 | 0.20 | 1.82 | 27.54 | 1.16 | 0.9967 | |
| 弱势粒 | 24.60 | 3532.45 | 0.24 | 4.76 | 24.22 | 1.23 | 0.9961 | ||
| WD40 | 强势粒 | 26.42 | 18.74 | 0.18 | 1.80 | 26.33 | 1.06 | 0.9970 | |
| 弱势粒 | 22.65 | 7485.26 | 0.21 | 5.01 | 20.92 | 0.03 | 0.9999 |
由表4可知,2个品种各处理下弱势粒最终粒重(生长终值量A)均明显低于强势粒,且强、弱势粒的生长量W均表现出CK>WD20>WD40的趋势。南粳5718在WD20处理下强势粒和弱势粒粒重分别较CK处理降低了7.9%和9.6%,WD40处理下强势粒和弱势粒的粒重分别较CK降低了13.9%和16.3%;徽两优898在WD20处理下强势粒和弱势粒的粒重分别较CK处理降低了6.4%和3.2%,WD40处理下强势粒和弱势粒的粒重分别较CK处理降低了10.3%和10.9%。
表5 结实期不同灌溉深度对水稻籽粒灌浆特征参数的影响
Table 5
| 品种 Variety | 处理 Treatment | 粒位 Grain position | 起始生长势 R0 | 达到最大灌浆速率的时间 Tmax (d) | 平均灌浆速率 GRmean (mg/d) | 最大灌浆速率 GRmax (mg/d) | 活跃灌浆期 D (d) |
|---|---|---|---|---|---|---|---|
| 南粳5718 NG5718 | CK | 强势粒 | 0.1241 | 14.21 | 0.9025 | 1.8051 | 31.85 |
| 弱势粒 | 0.0884 | 20.72 | 0.6189 | 1.2379 | 38.43 | ||
| WD20 | 强势粒 | 0.1367 | 13.83 | 0.8575 | 1.7151 | 30.88 | |
| 弱势粒 | 0.0678 | 24.01 | 0.4843 | 0.9686 | 44.45 | ||
| WD40 | 强势粒 | 0.1361 | 13.17 | 0.6721 | 1.3443 | 36.84 | |
| 弱势粒 | 0.0587 | 30.50 | 0.3903 | 0.7806 | 51.01 | ||
| 徽两优898 HLY898 | CK | 强势粒 | 0.0950 | 13.93 | 0.8255 | 1.6510 | 35.68 |
| 弱势粒 | 0.0653 | 25.14 | 0.5807 | 1.1614 | 43.77 | ||
| WD20 | 强势粒 | 0.1079 | 12.26 | 0.7084 | 1.4168 | 38.94 | |
| 弱势粒 | 0.0597 | 27.94 | 0.4921 | 0.9842 | 49.99 | ||
| WD40 | 强势粒 | 0.0988 | 13.18 | 0.6179 | 1.2358 | 42.75 | |
| 弱势粒 | 0.0427 | 34.12 | 0.3460 | 0.6919 | 65.46 |
图1
图1
结实期不同灌溉深度下水稻籽粒粒重和灌浆速率动态
Fig.1
Grain weight and grain filling rate dynamic of rice under different irrigation depths during the grain filling stage
2个品种不同灌溉处理下强势粒达到GRmax的时间差异较小,但弱势粒达到GRmax的时间在不同处理间存在明显差异,CK处理明显早于WD20处理,WD20处理又明显早于WD40处理,且WD40处理与WD20之间的差异较WD20与CK处理之间的差异更大。2个品种强、弱势粒GRmax和GRmean均表现出CK>WD20>WD40的趋势。南粳5718强势粒活跃灌浆期在CK和WD20处理之间无明显差异,但WD40处理下强势粒活跃灌浆期较CK和WD20明显延长,WD20处理下弱势粒活跃灌浆期明显长于CK处理,WD40处理下弱势粒活跃灌浆期又明显长于WD20处理。徽两优898强势粒与弱势粒活跃灌浆期均表现为CK<WD20<WD40,且弱势粒活跃灌浆期在不同处理间差异大于强势粒。
2.4 不同灌溉深度对水稻剑叶光合参数的影响
本试验测定了抽穗后15和35 d水稻剑叶光合参数和SPAD值(表6)。方差分析结果表明,品种对D35的Ci和D15的Tr具有显著影响,而灌溉深度对2个时期的Pn、D15的Gs和2个时期的Tr均具有显著影响,品种和灌溉深度均对D15和D35的剑叶SPAD值具有极显著影响,两因素互作除了对D15的SPAD值有显著影响外,对其他不同时期剑叶光合参数和SPAD值均无显著影响。
表6 结实期不同灌溉深度对水稻抽穗后15和35 d剑叶光合参数和SPAD值的影响
Table 6
| 品种 Variety | 处理 Treatment | Pn [μmol/(m2·s)] | Gs [mmol/(m2·s)] | Ci (μmol/mol) | Tr [mmol/(m2·s)] | SPAD值SPAD value | |||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| D15 | D35 | D15 | D35 | D15 | D35 | D15 | D35 | D15 | D35 | ||||||
| 南粳5718 NG5718 | CK | 27.76a | 14.90a | 0.44a | 0.29a | 373.00a | 294.67a | 8.57a | 3.27b | 44.77a | 36.50a | ||||
| WD20 | 24.60ab | 13.10a | 0.39ab | 0.31a | 359.00b | 298.00b | 8.39ab | 3.95a | 49.50a | 38.67a | |||||
| WD40 | 22.59b | 19.73b | 0.33b | 0.35a | 351.33b | 324.67b | 7.53b | 4.43a | 47.17a | 43.83b | |||||
| 徽两优898 HLY898 | CK | 25.60a | 11.14a | 0.38a | 0.32a | 379.00a | 342.00a | 9.47a | 3.63a | 43.03a | 26.20a | ||||
| WD20 | 25.53a | 10.60a | 0.36a | 0.29a | 374.67a | 330.33a | 9.19ab | 3.84a | 41.70a | 31.77ab | |||||
| WD40 | 23.47b | 13.76b | 0.32b | 0.36a | 371.00a | 338.33a | 8.21b | 3.27b | 41.80a | 35.07b | |||||
| 方差分析 Analysis of variance | V | 0.52 | 6.95* | 2.89 | 0.06 | 0.55 | 8.55* | 6.80* | 1.12 | 37.34** | 99.70** | ||||
| ID | 8.44* | 5.51* | 8.70* | 0.09 | 0.57 | 0.42 | 4.82* | 5.90* | 12.22** | 11.92** | |||||
| V×ID | 3.17 | 1.78 | 1.71 | 0.22 | 0.13 | 0.40 | 0.07 | 3.51 | 7.28* | 0.64 | |||||
南粳5718与徽两优898的光合参数在不同灌溉处理下表现出明显的响应差异。在D15时期,南粳5718剑叶的Pn在WD20和WD40处理下较CK处理分别降低了11.4%和18.6%,徽两优898的Pn在CK与WD20处理之间无显著差异,但该品种WD40下的Pn较CK处理降低了8.6%;剑叶Gs和Ci在不同水稻品种中均呈现出CK>WD20>WD40的趋势,但在WD20与WD40处理之间无显著差异;南粳5718剑叶Tr在WD20和WD40处理下较CK分别下降了2.1%和12.1%,而徽两优898的Tr在WD20和WD40处理下较CK分别下降了2.9%和16.9%。D15时期2个品种剑叶SPAD值在不同灌溉处理之间无显著差异,随着灌浆进行,2个品种剑叶SPAD值在D35时期均明显小于D15,在D35时期,2个品种剑叶SPAD值均表现为WD40>WD20>CK,且WD40处理与CK之间的差异达到了显著水平。图2是抽穗后35 d植株形态照片,可知WD40处理下水稻植株整体最绿,其次是WD20处理,最后是CK处理。
图2
图2
抽穗后35 d不同灌溉深度下水稻植株形态
Fig.2
Rice plant morphology under different irrigation depths 35 days after heading
3 讨论
水浆管理是调控水稻生长发育的重要农艺措施。水稻高产栽培研究[8-9,17]表明,分蘖期和孕穗期浅湿交替为主、结实期干湿交替的灌溉方式既能满足水稻生理和生态需水,也能促进水稻高产优质形成。近年来,在水稻价格持续不振的背景下,稻渔综合种养因其经济效益好、生态环境友好而得到大面积推广[18],稻虾、稻鱼和稻蟹等稻渔共作模式均需在水稻生长期间建立较深的水层来为水产动物创造适宜的生长环境,所以半深水灌溉是稻渔共作农事管理区别于水稻单作生产的一个重要特征[2],因此,必须要重视半深水灌溉之于水稻生长的效应,而灌浆结实期是水稻产量形成至关重要的阶段,该阶段光合物质生产及其转运利用直接关系到水稻籽粒灌浆水平[15,19]。本研究结果表明,结实期20和40 cm灌溉深度均降低了水稻产量,且灌水越深,产量降幅越大,结实率和千粒重下降是深水灌溉下水稻减产的主要原因,而WD40处理下每穗颖花数相较CK也出现了一定程度下降。南粳5718千粒重下降幅度小于徽两优898。
水稻籽粒灌浆的同化物主要来源于2个部分,其一是抽穗前茎鞘积累的非结构性碳水化合物(non-structural carbohydrates,NSC),灌浆启动后NSC被分解成可溶性糖,然后被动员、运输到籽粒中,这部分约占籽粒灌浆来源物质的30%[20];本研究发现,相较常规灌溉方式,结实期半深水灌溉下水稻成熟期穗干重、茎鞘物质输出量和转运率明显下降,且茎鞘物质输出率和转运率在WD40和WD20之间的差异明显大于WD20和CK之间,而成熟期茎鞘干重显著高于CK,这说明半深水灌溉抑制了茎鞘和叶片干物质向籽粒的转运及利用,且灌溉深度达40 cm时加重了这一效应。其二是抽穗后功能叶片光合作用生产的蔗糖运输到籽粒中,在一系列酶的作用下合成淀粉和蛋白质,从而完成粒重的积累[21]。本研究发现,抽穗后35 d剑叶的SPAD值和Pn较抽穗后15 d明显下降,表明随着灌浆进行,水稻叶片逐渐衰老,叶片光合能力显著下降,抽穗后15 d CK和WD20的SPAD值和Pn无显著差异,而WD40的Pn和Gs相较CK和WD20均明显下降,而在抽穗后35 d两个品种在WD40下的Pn均明显高于CK和WD20,同时SPAD值表现出WD40>WD20>CK的趋势,且成熟期半深水灌溉处理下叶片干重较CK处理高,但半深水灌溉下水稻籽粒灌浆和茎鞘转运却明显差于常规灌溉处理,再结合图2中半深水灌溉下水稻植株明显较常规灌溉下更深绿的表型,说明半深水灌溉下水稻存在明显的“滞绿”现象,即虽然水稻叶片衰老过慢,但其虽有较高的光合速率但生产的光合物质却无法被充分转运利用。Zang等[22]研究发现,叶片“滞绿”的水稻籽粒灌浆时淀粉合成相关酶活性明显低于叶片正常衰老的水稻,导致粒重下降,这与本研究半深水灌溉下的“叶绿粒轻”的现象相似,推测库活性不足可能是半深水灌溉下水稻籽粒灌浆变差的原因。同时期同一灌溉处理下南粳5718的SPAD值和Pn始终高于徽两优898,表明南粳5718叶片的光合能力强于徽两优898且衰老速度更慢,这可能是南粳5718在半深水灌溉下粒重下降幅度相对较小的原因。
水稻颖花根据其着生位置可分为强势粒和弱势粒,位于上部和一次枝梗上的强势粒相比位于基部和二次枝梗上的弱势粒开花更早,强势粒通常结实率高、灌浆快且籽粒重,而弱势粒结实率低、灌浆慢且籽粒轻[23]。本研究对强势粒和弱势粒灌浆过程分析发现,半深水灌溉相较常规灌溉并未明显改变水稻强势粒的灌浆启动势,但明显降低了强势粒的GRmax和GRmean,而弱势粒的灌浆启动势、GRmean和GRmax在WD20和WD40处理下均明显下降,虽然半深水灌溉下水稻籽粒活跃灌浆期有所延长,但不足以弥补灌浆速率下降的损失,这导致粒重显著下降,水稻在40 cm灌溉下相较20 cm灌溉下籽粒灌浆速率进一步下降,且弱势粒GRmean和粒重相较强势粒在半深水灌溉下表现出更大的降幅,其他研究[26-
半深水既是稻渔共作主动采取的灌溉深度,也可能是水稻生产被动遭遇的场景,2024年8-10月我国南方地区台风暴雨频发,而南方水稻正大量处于灌浆结实期[29],持续强降雨可能会引发稻田被动处于半深水环境。本研究明确了半深水灌溉对水稻籽粒灌浆和产量具有负面影响,且灌水越深,负效应越大,因此稻渔综合种养应尽可能规避于水稻灌浆结实期进行半深水灌溉,如确有半深水养殖水产动物的需要,也应尽可能降低灌溉深度和选择对半深水环境相对钝感的水稻品种。
4 结论
灌浆结实期半深水灌溉降低了水稻结实率、千粒重和产量,叶片及植株滞绿、光合物质转运利用低、籽粒灌浆不良是结实期半深水灌溉下千粒重、结实率和产量低的重要原因,灌溉深度增加会进一步加剧半深水灌溉对水稻籽粒灌浆和产量的负面影响。南粳5718较徽两优898对半深水灌溉钝感,弱势粒灌浆较强势粒对半深水灌溉更加敏感。
参考文献
稻虾共作模式研究进展
DOI:10.13304/j.nykjdb.2020.1114
[本文引用: 1]
水稻(Oryzasativa)-克氏原螯虾(Procambarusclarkii,以下称小龙虾)共作模式作为稻渔综合种养的重要模式,近些年在我国尤其是长江中下游地区发展迅速。综述了近10年来稻虾共作模式相关理论研究进展和产业发展情况。在理论研究方面,从稻田土壤、水质、生物多样性和温室气体排放等方面阐述了稻虾共作对稻田生态的影响;在技术研究方面,稻虾共作模式的推广有利于减少化肥、农药投入,可在稳定水稻播种面积的基础上达到虾稻同步增产、品质同步提升的效果,但稻虾产业的发展壮大还需进行产业结构优化和升级。与此同时,稻虾共作模式的发展还存在入侵风险、种质退化、“重虾轻稻”等问题,制约了稻虾共作模式的可持续发展。为推动稻虾共作模式研究和广泛应用,提出3点建议:建立稻田信息库和农情分析系统,对稻虾共作推广区域进行等级划分;推进产学研一体化,强化基础理论和集成技术模式研究;加强政策支持和技术服务,保障经营主体经济效益。
Deepwater rice: A model plant to study stem elongation
DOI:10.1104/pp.118.4.1105 PMID:9847084 [本文引用: 1]
Agronomic performance of high- yielding rice variety grown under alternate wetting and drying irrigation
DOI:10.1016/j.fcr.2011.09.018 URL [本文引用: 2]
水稻超高产栽培研究与探讨
DOI:10.3969/j.issn.1006-8082.2012.01.001
[本文引用: 1]
扼要回顾了国内外水稻超高产栽培的研究进展,总结了扬州大学农学院关于水稻超高产栽培研究得到的水稻超高产形成规律、栽培途径以及关键技术与精确定量栽培技术的因地制宜集成,并对水稻超高产栽培研究进行了展望与探讨。
Mechanisms associated with tiller suppression under stagnant flooding in rice
DOI:10.1111/jac.2019.205.issue-2 URL [本文引用: 1]
Flooding stress: the effects of planting pattern and water regime on root morphology, physiology and grain yield of rice
DOI:10.1111/jac.2010.196.issue-5 URL [本文引用: 1]
水稻不同灌溉方式下的高产生理特性
以3个高产杂交水稻组合中优6号、两优培九、Ⅱ优7954为材料,研究了水稻好气灌溉条件下的生理特性。与传统淹水灌溉相比较,好气灌溉条件下水稻整个生育期田间水位降低,土壤氧化还原电位提高;分蘖早发,前期分蘖和大分蘖比例提高,分蘖成穗数提高7.1%~18.3%,增产8.6%~10.8%。同时,好气灌溉在单丛叶面积没有显著下降的情况下,提高群体各层次的透光率。与传统方式比较,水稻好气灌溉处理在孕穗期和开花期根系单茎伤流量增加,花后叶片光合速率、气孔导度和比叶重提高。
Effects of semi-deep water irrigation on hybrid indica rice lodging resistance
DOI:10.3389/fpls.2022.1038129
URL
[本文引用: 1]
Recently, rice-aquatic animal integrated farming (RAAIF) has grown rapidly in China due to its favorable benefits and the lower application of pesticides and fertilizers. However, rice lodging occurs frequently under RAAIF, which restricts rice yield. We assumed that semi-deep water irrigation may cause weaker rice-lodging resistance since it is the most significant environmental factor for RAAIF that distinguishes it from rice monoculture. To investigate the response of rice stem lodging resistance to semi-deep water irrigation and its mechanism, three irrigation management modes, namely the typical high-yield irrigation model that is mainly based on swallow and wetting (CK), semi-deep water irrigation from the late tillering stage to the jointing stage (SDI1), and semi-deep water irrigation from the jointing stage to the middle grain-filling stage (SDI2), were conducted using three hybrid indica rice varieties: Shenliangyou136 (SLY136), Huiliangyousimiao (HLYSM), and Wanxiangyou982 (WXY982). Mechanics analysis indicated that the bending moment by the whole plant (WP) and the breaking strength (M) were both decreased by semi-deep water irrigation when compared with CK, while M presented a larger decreasing amplitude than WP, which induced the increased lodging index (LI) of rice, for all the tested varieties. SLY136 and HLYSM were affected more strongly by SDI1, whereas WXY982 was affected more strongly by SDI2. Significant weaker breaking force under two semi-deep water irrigation modes contributed to the decreased M relative to CK. Morphology results showed that semi-deep water irrigation reduced the thickness of mechanical tissues, sclerenchyma cells, and parenchyma cells; reduced the number of vascular bundles; and caused a looser arrangement, inducing the lower fullness of the rice basal internode. Decreased accumulation of lignin and cellulose was also linked to the weaker breaking force of the basal internode under semi-deep water irrigation, which was verified by correlation analysis. WXY982 had obvious lower structural carbohydrates content under semi-deep water irrigation than the other two varieties and thus showed worse breaking force and LI. In conclusion, the worse mechanical strength of the rice basal internode under semi-deep water irrigation was closely associated with weaker vascular bundle development and suppressed structural carbohydrate accumulation, and the decreasing degree of lodging resistance varied between rice varieties and semi-deep water irrigation periods.
不同水氮管理对水稻干物质积累和茎鞘物质运转及产量的影响
DOI:10.11869/j.issn.100-8551.2016.02.0347
[本文引用: 2]
为了探讨不同类型高产水稻水氮高效利用特性,以大穗型杂交粳稻甬优8号和穗粒兼顾型常规粳稻品种(系)苏10-100为材料,进行实地氮肥管理和全生育期轻干-湿交替灌溉技术联合运用,研究水稻干物质和茎鞘非结构性碳水化合物(NSC)积累与运转特性及其与籽粒产量形成的关系。结果表明,与常规水肥管理相比,实地氮肥管理和轻干-湿交替灌溉联合运用显著增加了幼穗分化期至成熟期的干物质积累量和抽穗期茎鞘中NSC含量,提高了茎鞘干物质和NSC运转率和对籽粒的贡献率,大穗型杂交粳稻甬优8号的运转率和贡献率明显大于常规粳稻苏10-100;水氮处理降低了穗数,但显著或极显著地提高了每穗粒数、结实率、充实度和千粒重,苏10-100和甬优8号分别增加了6.21%、2.53%,1.68%、13.63%,3.3%、8.1%和9.06%、10.35%,其中2个弱势籽粒千粒重的增幅分别达到了16.3%和15.9%,显著大于强势粒。因此,采用实地氮肥管理和轻干-湿交替灌溉水稻具有中后期单位面积干物质积累量大,物质运转率高,穗大且多,结实率高、充实度好的特点,有利于促进弱势籽粒灌浆充实,提高籽粒产量。本研究为水稻超高产栽培和不同类型水稻养分水份高效管理提供理论依据和实践指导。
Impact of alternate wetting and drying on rice physiology, grain production, and grain quality
DOI:10.1016/j.fcr.2017.01.016 URL [本文引用: 1]
孕穗期施硅对高温下扬花灌浆期水稻干物质转运及产量的影响
DOI:10.11869/j.issn.100-8551.2016.09.1833
[本文引用: 1]
为探明硅对高温逆境下水稻产量形成的调控效应,以圣稻19为试验材料,通过大田增温试验研究了孕穗期叶面施硅对高温下扬花灌浆期水稻干物质转运与产量的影响。结果表明,与不施硅相比,施硅显著提高了高温逆境下水稻叶片干物质输出量、输出率、转化率,对茎鞘干物质的输出转化无显著影响;减弱了自然温度下水稻叶片干物质的转化率。高温或自然温度下,施硅均显著提高了水稻库容量;高温逆境下,施硅显著增加了水稻结实率与产量,但显著降低了千粒重;自然温度下,显著提高了水稻结实率、千粒重与产量。施硅后叶片输出转化效率与结实率的增加是弥补高温下水稻产量损失的主要因素。本研究结果为完善水稻耐热栽培调控技术提供了有益参考。
Pre-anthesis non-structural carbohydrate reserve in the stem enhances the sink strength of inferior spikelets during grain filling of rice
DOI:10.1016/j.fcr.2011.05.015 URL [本文引用: 1]
The relationships among “STAY-GREEN” trait, post-anthesis assimilate remobili-zation, and grain yield in rice (Oryza sativa L.)
DOI:10.3390/ijms232213668
URL
[本文引用: 1]
The mobilization and translocation of carbohydrates and mineral nutrients from vegetative plant parts to grains are pivotal for grain filling, often involving a whole plant senescence process. Loss of greenness is a hallmark of leaf senescence. However, the relationship between crop yield and senescence has been controversial for many years. Here, in this study, the overexpression and RNA interference lines of gene of OsNYC3 (Non-Yellow Coloring 3), a chlorophyll catabolism gene, were investigated. Furthermore, exogenous phytohormones were applied, and a treatment of alternate wetting and moderate drying (AWMD) was introduced to regulate the processes of leaf senescence. The results indicated that the delayed senescence of the “STAY-GREEN” trait of rice is undesirable for the process of grain filling, and it would cause a lower ratio of grain filling and lower grain weight of inferior grains, because of unused assimilates in the stems and leaves. Through the overexpression of OsNYC3, application of exogenous chemicals of abscisic acid (ABA), and water management of AWMD, leaf photosynthesis was less influenced, a high ratio of carbohydrate assimilates was partitioned to grains other than leaves and stems as labeled by 13C, grain filling was improved, especially for inferior spikelets, and activities of starch-synthesizing enzymes were enhanced. However, application of ethephon not only accelerated leaf senescence, but also caused seed abortion and grain weight reduction. Thus, plant senescence needs to be finely adjusted in order to make a contribution to crop productivity.
Grain-filling problem in ‘super’ rice
DOI:10.1093/jxb/erp348 URL [本文引用: 1]
Effects of high temperature and shading on grain abscisic acid content and grain filling pattern in rice (Oryza sativa L.)
DOI:10.1080/1343943X.2018.1524264 URL [本文引用: 1]
Response of rice with overlapping growth stages to water stress by assimilates accumulation and transport and starch synthesis of superior and inferior grains
DOI:10.3390/ijms231911157
URL
[本文引用: 1]
Drought stress at jointing–booting directly affects plant growth and productivity in rice. Limited by natural factors, the jointing and booting stages of short-growth-period rice varieties are highly overlapped in high-latitude areas, which are more sensitive to water deficit. However, little is known about the dry matter translocation in rice and the strategies of starch synthesis and filling of superior and inferior grains under different drought stress was unclear. In this study, the rice plants were subjected to three degrees of drought stress (−10 kPa, −25 kPa, −40 kPa) for 15 days during the jointing–booting stage; we investigated dry matter accumulation and translocation, grain filling and enzyme activities to starch synthesis of superior and inferior grains in rice with overlapping growth stages from 2016 to 2017. The results showed that drought stress significantly reduced dry matter accumulation in the stems and leaves. Mild and moderate drought increased dry matter translocation efficiency. However, severe drought stress largely limited the dry matter accumulation and translocation. A large amount of dry matter remains in vegetative organs under severe drought stress. The high content in NSC in stem and sheath plays a key role in resisting drought stress. The drought stress at jointing–booting directly caused a change in the grain filling strategy. Under moderate and severe drought, the grain-filling active period of the superior grains was shortened to complete the necessary reproductive growth. The grain-filling active period of the inferior grains was significantly prolonged to avoid a decrease in grain yield. The significant decrease in the grain-filling rate of the superior and inferior grains caused a reduction in the thousand-grain weight. In particular, the influence of the grain-filling rate of inferior grains on the thousand-grain weight was more significant. Drought stress changed the starch synthesis strategies of the superior and inferior grains. Soluble starch synthase and starch branching enzyme activities of inferior grains increased significantly under drought stress. GBSS activity was not sensitive to drought stress. Therefore, amylose content was decreased and amylopectin synthesis was enhanced under drought stress, especially in inferior grains.
水稻氧营养的生理,生态机制及环境效应研究进展
DOI:10.3969/j.issn.1001-7216.2009.04.01
氧是水稻生命活动过程中重要的营养因子。水稻利用通气组织将大气中的氧从地上部分转运到根部,同时也能通过根系从生长介质中吸收一部分氧。良好的氧营养状况下水稻才能正常生长,不仅如此,氧营养还参与了水稻土壤环境系统的作用过程,具有重要的生态效应。从氧营养的水稻生物学响应和根际生态效应两个方面综述了近年来稻田氧营养相关研究状况,重点总结了目前国内外有关水稻需氧性,氧营养对水稻形态学、解剖学特征的影响,相应的生理、生态机制以及环境效应的研究成果,同时比较了水稻生产上所采用的氧营养调控途径,并对水稻氧营养研究作了展望。
Reprint of “Morphological and physiological traits of roots and their relationships with water productivity in water-saving and drought-resistant rice”
DOI:10.1016/j.fcr.2014.06.026 URL [本文引用: 1]
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