作物杂志, 2026, 42(3): 21-29 doi: 10.16035/j.issn.1001-7283.2026.03.004

第二十八届中国科协年会学术论文专栏(主要粮食作物产能品质提升与高质量发展路径)

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

郭威,1,2,3, 孙冬1,2, 闫治霖1,2, 丁蓓1,2, 杜跃辰1,2, 吴慧扬1,2, 徐强1,2,3, 窦志,1,2,3, 高辉1,2,3

1 江苏省粮食作物现代产业技术协同创新中心225009江苏扬州

2 江苏省优质粳稻产业工程研究中心225009江苏扬州

3 扬州大学水稻产业工程技术研究院225009江苏扬州

Influence of Semi-Deep Water Irrigation during Grain-Filling Period on Rice Yield, Dry Matter Transportation and Grain-Filling

Guo Wei,1,2,3, Sun Dong1,2, Yan Zhilin1,2, Ding Bei1,2, Du Yuechen1,2, Wu Huiyang1,2, Xu Qiang1,2,3, Dou Zhi,1,2,3, Gao Hui1,2,3

1 Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops, Yangzhou 225009, Jiangsu, China

2 Jiangsu High-Quality Japonica Rice Industrial Engineering Research Center, Yangzhou 225009, Jiangsu, China

3 Research Institute of Rice Industrial Engineering Technology, Yangzhou 225009, Jiangsu, China

通讯作者: 窦志,主要从事水稻高产优质栽培和稻渔综合种养研究,E-mail:douzhi@yzu.edu.cn

收稿日期: 2025-12-12   修回日期: 2026-03-25   网络出版日期: 2026-04-10

基金资助: 天长市横向课题(20230705000010)
国家重点研发计划(2024YFD2300504)

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:1034177399@qq.com

摘要

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: 水稻; 半深水灌溉; 产量; 干物质转运; 籽粒灌浆

Abstract

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: Rice; Semi-deep water irrigation; Yield; Dry matter transportation; Grain filling

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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. 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. 作物杂志, 2026, 42(3): 21-29 doi:10.16035/j.issn.1001-7283.2026.03.004

Guo Wei, Sun Dong, Yan Zhilin, Ding Bei, Du Yuechen, Wu Huiyang, Xu Qiang, Dou Zhi, Gao Hui. Influence of Semi-Deep Water Irrigation during Grain-Filling Period on Rice Yield, Dry Matter Transportation and Grain-Filling. Crops, 2026, 42(3): 21-29 doi:10.16035/j.issn.1001-7283.2026.03.004

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。

水浆管理是调控水稻产量形成的重要农艺措施。高产栽培研究[8]表明,“寸水缓苗、浅水分蘖、够苗搁田、孕穗抽穗有水、干湿交替灌浆”的水分管理方式,有助于促进秧苗返青与分蘖、控制无效分蘖、提高土壤氧化还原电位、增强根系与茎秆发育,促进灌浆期间同化物向籽粒转运,这种“浅、湿、露、晒”相结合的水浆管理模式,能有效提高水稻生理活性、协调各产量因素,从而实现水稻高产[9-10]。而发生涝害或持续淹灌会抑制水稻分蘖发生[11]、阻碍根系发育[12]、减少干物质积累与转运[13]、增加倒伏风险[14],最终导致产量与品质下降。

灌浆结实期是水稻产量形成的关键阶段,该时期干物质转运与籽粒灌浆水平直接决定水稻产量的高低,且部分稻渔共作模式在灌浆结实期仍实施半深水灌溉,例如湖北和浙江地区的稻鱼共作模式[5,7]。然而,结实期半深水灌溉是否会对水稻产量构成显著影响,干物质转运及籽粒灌浆对半深水灌溉又作何响应,目前还鲜有报道。

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  Water management methods in the whole growth period of each treatment

生育时期Growth periodCKWD20WD40
缓苗期Seedling recovering0~2 cm水层0~2 cm水层0~2 cm水层
分蘖期Tillering2~4 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

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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:

$\begin{array}{c}\text{W}\text{=}\text{A}{\left(\text{1+}\text{B}{\text{e}}^{\text{-}\text{Kt}}\right)}^{\text{-}\frac{\text{1}}{\text{N}}}\end{array}$

式中,W为各期生长量,即千粒重,g;t为齐穗后天数,d;ABKN为方程参数,并用决定系数(r2)(Wt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:

R0=K/N
Tmax=(lnB-lnN)/K
GRmean=AK/[2(N+2)]
$\begin{array}{c}\text{G}\text{R}\text{m}\text{a}\text{x}\text{=}\text{A}{\text{(}\text{N}\text{+1}\text{)}}^{\text{-}\frac{\text{1}}{\text{N}}}\mathit{ }\end{array}$
D=[2(N+2)]/K

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  Effects of different irrigation depths on rice yield and harvest index during grain filling stage

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
CK316.30a168.86a91.91a26.44a12.25a12.19a58.71a
WD20311.60a159.92ab88.60b25.54b11.02b10.98b56.93ab
WD40309.13a153.73b83.22c25.18c9.41c9.31c52.86b
徽两优898
HLY898
CK234.64a242.98a88.73a25.80a12.08a12.39a60.33a
WD20232.22a234.78ab84.19b24.42b10.42b10.63b58.16ab
WD40234.96a223.82b80.07c23.87c9.12c9.05c53.58b
方差分析
Analysis of variance
V2453.09**2723.53**90.91**46.41**8.63*28.82**23.28**
ID2.1682.01**178.20**39.41**186.48**33.34**104.70**
V×ID1.962.991.211.711.090.052.27

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.

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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  Effects of different irrigation depths on dry matter accumulation and translocation of rice during grain filling stage

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
CK3.27a1.17a0.56a5.00a2.56b0.49b
WD203.28a1.07a0.58a4.93a2.65b0.55ab
WD403.24a1.05a0.57a4.86a2.81a0.60a
徽两优898
HLY898
CK3.88a0.93a0.81a5.62a3.03b0.59b
WD203.92a0.94a0.79a5.65a3.10b0.64b
WD403.86a0.93a0.83a5.60a3.35a0.74a
方差分析
Analysis of variance
V87.38**35.97**161.88**16.32*62.42**3.68**
ID0.830.820.450.4724.91**2.64**
V×ID0.120.771.360.380.053.71
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
CK4.34a7.39a3.78a0.71a21.80a21.81a
WD204.08b7.29a3.49ab0.62a19.04b17.84a
WD403.82c7.22a3.25b0.42b13.15c13.10b
徽两优898
HLY898
CK5.82a9.49a5.01a0.85a21.91a17.01a
WD205.39b9.27a4.60b0.78a19.81b16.89a
WD404.52c8.44b3.69c0.53b13.66c14.29b
方差分析
Analysis of variance
V162.47**266.42**74.39**30.98**1.932.29
ID73.18**28.87**60.01**172.96**217.86**58.08**
V×ID12.34*15.40*10.61**0.520.316.67*

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.

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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  Richards equation parameter estimates for grain filling process

品种
Variety
处理
Treatment
粒位
Grain position
ABKN生长量
W(g)
标准差
S
决定系数
r2
南粳5718
NG5718
CK强势粒28.7575.890.252.0528.740.570.9994
弱势粒23.79542.920.252.8623.740.560.9992
WD20强势粒26.4854.170.251.8026.471.330.9964
弱势粒21.532404.840.273.9421.450.440.9993
WD40强势粒24.7614.340.181.3324.690.960.9976
弱势粒19.915440.380.244.0319.311.450.9917
徽两优898
HLY898
CK强势粒29.46129.160.272.8729.450.90.9983
弱势粒25.429970.800.304.6725.361.360.9958
WD20强势粒27.5920.100.201.8227.541.160.9967
弱势粒24.603532.450.244.7624.221.230.9961
WD40强势粒26.4218.740.181.8026.331.060.9970
弱势粒22.657485.260.215.0120.920.030.9999

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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显示,2个品种籽粒灌浆的GRmeanGRmax均表现为CK>WD20>WD40。由图1可知,2个品种不同处理强、弱势粒的灌浆速率均表现出先上升后下降的趋势,强势粒的GRmaxGRmean均高于相同处理下的弱势粒,且强势粒达到GRmax的时间明显早于弱势粒,强势粒的活跃灌浆期均短于同处理下的弱势粒。

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  Effects of different irrigation depths on grain filling characteristic parameters of rice at grain filling stage

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0.124114.210.90251.805131.85
弱势粒0.088420.720.61891.237938.43
WD20强势粒0.136713.830.85751.715130.88
弱势粒0.067824.010.48430.968644.45
WD40强势粒0.136113.170.67211.344336.84
弱势粒0.058730.500.39030.780651.01
徽两优898
HLY898
CK强势粒0.095013.930.82551.651035.68
弱势粒0.065325.140.58071.161443.77
WD20强势粒0.107912.260.70841.416838.94
弱势粒0.059727.940.49210.984249.99
WD40强势粒0.098813.180.61791.235842.75
弱势粒0.042734.120.34600.691965.46

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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GRmaxGRmean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  Effects of different irrigation depths during the grain filling stage on photosynthetic parameters and SPAD values of flag leaves at 15 and 35 days after heading

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
D15D35D15D35D15D35D15D35D15D35
南粳5718
NG5718
CK27.76a14.90a0.44a0.29a373.00a294.67a8.57a3.27b44.77a36.50a
WD2024.60ab13.10a0.39ab0.31a359.00b298.00b8.39ab3.95a49.50a38.67a
WD4022.59b19.73b0.33b0.35a351.33b324.67b7.53b4.43a47.17a43.83b
徽两优898
HLY898
CK25.60a11.14a0.38a0.32a379.00a342.00a9.47a3.63a43.03a26.20a
WD2025.53a10.60a0.36a0.29a374.67a330.33a9.19ab3.84a41.70a31.77ab
WD4023.47b13.76b0.32b0.36a371.00a338.33a8.21b3.27b41.80a35.07b
方差分析
Analysis of variance
V0.526.95*2.890.060.558.55*6.80*1.1237.34**99.70**
ID8.44*5.51*8.70*0.090.570.424.82*5.90*12.22**11.92**
V×ID3.171.781.710.220.130.400.073.517.28*0.64

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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GsCi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PnGs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GRmaxGRmean,而弱势粒的灌浆启动势、GRmeanGRmax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[26-28]也发现,在面对非生物胁迫时弱势粒灌浆往往比强势粒表现差[24-25],笔者认为这可能与弱势粒灌浆启动晚、其快速灌浆阶段水稻根系活力已明显下降有关,因为水稻进入灌浆结实期后就不再长新根,而长期淹水灌溉下水稻根系缺氧造成大量有毒还原性物质积累、根系活力下降过快,这导致弱势粒灌浆的动能被进一步削弱。

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。

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