Grain-filling Characteristics and Its Relationship with Grain Yield Formation of japonica Rice Nanjing 9108 Under Combined Salinity-drought Stress

doi:10.16819/j.1001-7216.2025.240515

Chinese Journal OF Rice Science ›› 2025, Vol. 39 ›› Issue (3): 373-386.DOI: 10.16819/j.1001-7216.2025.240515

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Grain-filling Characteristics and Its Relationship with Grain Yield Formation of japonica Rice Nanjing 9108 Under Combined Salinity-drought Stress

WEI Huanhe¹, WANG Lulu¹^,², MA Weiyi¹, ZHANG Xiang¹, ZUO Boyuan¹, GENG Xiaoyu¹, ZHU Wang¹^,², ZHU Jizou¹, MENG Tianyao¹^,², CHEN Yinglong¹, GAO Pinglei¹, XU Ke¹, DAI Qigen¹^,^*()

¹Jiangsu Key Laboratory of Crop Genetics and Physiology / Jiangsu Key Laboratory of Crop Cultivation and Physiology / Co-Innovation Center for Modern Production Technology of Grain Crops / Research Institute of Rice Industrial Engineering Technology / Key Laboratory of Saline-Alkali Soil Reclamation and Utilization in Coastal Areas, Ministry of Agriculture and Rural Affairs of China / East China Branch of National Center of Technology Innovation for Saline-Alkali Tolerant Rice, Yangzhou University, Yangzhou 225009, China
²Joint International Research Laboratory of Agri-culture and Agro-product Safety, Ministry of Education / Institute of Agricultural Science and Technological Development, Yangzhou University, Yangzhou 225009, China

Received:2024-05-29 Revised:2024-10-15 Online:2025-05-10 Published:2025-05-21
Contact: *email: qgdai@yzu.edu.cn

盐−旱复合胁迫下粳稻品种南粳9108籽粒灌浆特性及其与产量形成的关系

韦还和¹, 汪璐璐¹^,², 马唯一¹, 张翔¹, 左博源¹, 耿孝宇¹, 朱旺¹^,², 朱济邹¹, 孟天瑶¹^,², 陈英龙¹, 高平磊¹, 许轲¹, 戴其根¹^,^*()

¹江苏省作物遗传生理重点实验室/江苏省作物栽培生理重点实验室/江苏省粮食作物现代产业技术协同创新中心/扬州大学水稻产业工程技术研究院/农业农村部盐碱土改良与利用(滨海盐碱地)重点实验室/国家耐盐碱水稻技术创新中心华东中心, 江苏扬州 225009
²扬州大学教育部农业与农产品安全国际合作联合实验室/扬州大学农业科技发展研究院, 江苏扬州 225009

通讯作者: *email: qgdai@yzu.edu.cn
基金资助:
国家自然科学基金资助项目(32472222);国家自然科学基金资助项目(32001466);江苏省重点研发计划资助项目(BE2023355);江苏省重点研发计划资助项目(1020);江苏省高等学校基础科学（自然科学）研究重大项目(23KJA210004);国家重点研发计划资助项目(2022YFE0113400);江苏省碳达峰碳中和科技创新专项资金资助项目(BE2022304);江苏省碳达峰碳中和科技创新专项资金资助项目(BE2022305);扬州大学‘青蓝工程'项目;江苏省高校优势学科建设工程资助项目(PAPD)

Abstract

Abstract:

【Objective】This study aims to investigate the effects of combined salinity-drought stress on rice grain-filling characteristics and its relationship with yield formation.【Method】The study was conducted with conventional japonica rice cultivar Nanjing 9108, which is widely cultivated in the coastal saline soil of Jiangsu Province, China. The experimental design consisted of factorial combinations of salinity and drought stresses: salinity stress induded non-salinity (NS, salt concentration 0%) and high-salinity stress (HS, salt concentration 0.3%); while drought regimes comprised well-watered (WW), moderate drought (MD), and severe drought (SD) treatments from jointing stage to heading stage.【Result】Compared with non-stress, 1) the yield reduction under salinity stress, drought stress, and combined salinity-drought stress was 36.8%, 13.7%−30.0%, and 60.3%−107.1%, respectively (two-year average). 2) the number of panicles per unit area, grains per panicle, seed setting rate, and the thousand-grain weight were all lower under single salinity stress, single drought stress, and combined salinity-drought stress, and the reduction was greater under combined salinity-drought stress; the panicle length, and grain weight and seed setting rate of superior and inferior grains showed a similar trend. 3) single salinity stress, single drought stress, and combined salinity-drought stress significantly reduced dry matter weight at jointing and heading and the accumulation of dry matter weight from heading to maturity, as well as leaf photosynthetic rate and SPAD value at 20 and 40 days after heading; the reductions under combined salinity-drought stress were greater than those under single salinity and single drought stress. 4) the maximum grain-filling rate and the mean grain-filling rate of superior and inferior grains were lower under single salinity stress, single drought stress, and combined salinity-drought stress, while the time to reach the maximum grain-filling rate and the effective grain-filling period of superior grains showed an increasing trend, and the differences in the time to reach the maximum grain-filling rate and the effective grain-filling period of inferior grains were smaller. The number of days in the early, middle, and late periods were greater under single salinity stress, single drought stress, and combined salinity-drought stress, while lower for the mean grain-filling rate and grain-filling amount. 5) single salinity stress, single drought stress, and combined salinity-drought stress treatments reduced the activities of key starch synthesis enzymes in grains, especially for AGPase and GBSS enzymes.【Conclusion】This study demonstrats that the grain-filling rate and key starch synthesis enzyme activities of superior and inferior grains decreased significantly and the grain-filling characteristics of superior and inferior grains were deteriorated under single salinity stress, single drought stress, and especially combined salinity-drought stress. The inhibitory effects of combined salinity-drought stress on the grain-filling rate, grain-filling amount of superior and inferior grains, and grain yield were greater than single salinity and drought stress.

Key words: rice, combined salinity-drought stress, grain-filling characteristics, grain yield formation

摘要：

【目的】明确盐−旱复合胁迫对水稻籽粒灌浆特性的影响及其与产量形成的关系。【方法】以江苏滨海盐碱地主栽的粳稻品种南粳9108为试材，设置盐胁迫与干旱胁迫组合处理，盐胁迫设置无盐(Non-salinity, NS, 盐浓度0%)和高盐(High-salinity stress, HS, 盐浓度0.3%)处理；干旱胁迫于拔节期至抽穗期设置浅水层灌溉(Well-watered, WW)、中度干旱(Moderate drought, MD)和重度干旱(Severe drought, SD)。【结果】与无胁迫相比，1）单一盐胁迫、单一干旱胁迫和盐−旱复合胁迫后产量分别下降36.8%、13.7%~30.0%和60.3%~107.1%(两年平均值)。2）单一盐、单一干旱和盐−旱复合胁迫下单位面积穗数、每穗粒数、结实率和千粒重均较低，且以盐−旱复合胁迫下的降幅更大；水稻穗长、穗部强、弱势粒的结实率和千粒重亦呈类似趋势。3）单一盐、单一干旱和盐−旱复合胁迫显著降低了抽穗期和成熟期干物质量及抽穗期至成熟期干物质积累量、抽穗后20 d和40 d叶片光合速率和SPAD值，盐−旱复合胁迫下的降幅高于单一盐和单一干旱胁迫。4）单一盐、单一干旱和盐−旱复合胁迫下强、弱势粒的最大灌浆速率和平均灌浆速率均较低，强势粒达最大灌浆速率的时间和有效灌浆天数则呈增加趋势，弱势粒达最大灌浆速率的时间和有效灌浆天数差异则较小。单一盐、单一干旱和盐−旱复合胁迫下灌浆前、中、后期的天数较高，灌浆速率和灌浆量则较低。5）单一盐、单一干旱和盐−旱复合胁迫处理降低了籽粒中淀粉合成关键酶活性，尤其是AGPase和GBSS酶活性。【结论】本研究表明，单一盐、单一干旱胁迫和盐−旱复合胁迫下水稻强、弱势粒灌浆速率和淀粉合成关键酶活性显著下降，弱化强、弱势粒充实性状，且以盐−旱复合胁迫下降幅更大。盐−旱复合胁迫对籽粒强、弱势粒灌浆速率和灌浆量以及产量的抑制效应高于单一盐和单一干旱胁迫。

关键词: 水稻, 盐−旱复合胁迫, 籽粒灌浆特性, 产量形成

WEI Huanhe, WANG Lulu, MA Weiyi, ZHANG Xiang, ZUO Boyuan, GENG Xiaoyu, ZHU Wang, ZHU Jizou, MENG Tianyao, CHEN Yinglong, GAO Pinglei, XU Ke, DAI Qigen. Grain-filling Characteristics and Its Relationship with Grain Yield Formation of japonica Rice Nanjing 9108 Under Combined Salinity-drought Stress[J]. Chinese Journal OF Rice Science, 2025, 39(3): 373-386.

韦还和, 汪璐璐, 马唯一, 张翔, 左博源, 耿孝宇, 朱旺, 朱济邹, 孟天瑶, 陈英龙, 高平磊, 许轲, 戴其根. 盐−旱复合胁迫下粳稻品种南粳9108籽粒灌浆特性及其与产量形成的关系[J]. 中国水稻科学, 2025, 39(3): 373-386.

Figures/Tables 11

Table 1. Grain yield and yield components of rice under salinity and drought stresses

年份 Year	盐处理 Salinity treatment	干旱处理 Drought treatment	单位面积穗数 NP (×10⁴ hm^-2)	每穗粒数 SPP	结实率 SSR (%)	千粒重 1000-GW (g)	实产 Actual yield (t hm^-2)
2021	无盐 NS	浅水层灌溉 WW	304 a	165 a	89.3 a	26.7 a	11.6 a
		中度干旱 MD	291 b	152 b	88.7 ab	26.0 b	10.3 b
		重度干旱 SD	273 c	145 c	87.2 b	25.6 c	8.9 c
	高盐 HS	浅水层灌溉 WW	271 c	140 d	84.3 c	25.4 c	8.5 d
		中度干旱 MD	256 d	133 e	82.8 d	24.8 d	7.4 e
		重度干旱 SD	238 e	117 f	79.7 e	24.0 e	5.7 f
2022	无盐 NS	浅水层灌溉 WW	307 a	162 a	89.7 a	26.8 a	11.7 a
		中度干旱 MD	287 b	154 b	88.9 ab	26.5 b	10.2 b
		重度干旱 SD	271 c	143 c	86.7 b	25.7 c	9.1 c
	高盐 HS	浅水层灌溉 WW	267 cd	137 d	84.0 c	25.6 c	8.6 d
		中度干旱 MD	253 d	134 e	82.3 d	24.8 d	7.2 e
		重度干旱 SD	241 e	119 f	78.9 e	24.2 e	5.6 f
方差分析 Analysis of variance
年份 Year(Y)			ns	ns	ns	ns	ns
盐处理 Salinity treatment(S)			**	**	**	**	**
干旱处理 Drought treatment(D)			**	**	**	**	**
年份×盐处理Y×S			ns	ns	ns	ns	ns
年份×干旱处理 Y×D			ns	ns	ns	ns	ns
盐处理×干旱处理 S×D			ns	ns	ns	ns	ns
年份×盐处理×干旱处理 Y×S×D			ns	ns	ns	ns	ns

Table 2. Panicle traits of rice under salinity and drought stresses

年份 Year	盐处理 Salinity treatment	干旱处理 Drought treatment	穗长 PL (cm)	强势粒 SG			弱势粒 IG
年份 Year	盐处理 Salinity treatment	干旱处理 Drought treatment	穗长 PL (cm)	籽粒数 NG	结实率 SSR (%)	千粒重 1000-GW (g)	籽粒数 NG	结实率 SSR (%)	千粒重 1000-GW (g)
2021	无盐 NS	浅水层灌溉 WW	13.9 a	19.3 a	92.6 a	29.0 a	35.0 a	87.5 a	24.6 a
		中度干旱 MD	12.7 b	17.6 b	91.7 ab	28.5 ab	31.8 b	86.6 ab	23.8 ab
		重度干旱 SD	12.1 bc	16.9 b	90.5 b	27.9 b	29.6 bc	84.7 b	23.7 ab
	高盐 HS	浅水层灌溉 WW	11.1 c	15.7 c	87.5 c	27.6 bc	28.3 c	81.7 c	23.2 b
		中度干旱 MD	10.7 cd	14.8 d	86.5 c	27.2 bc	26.5 d	79.6 cd	22.8 bc
		重度干旱 SD	10.2 d	13.6 e	84.1 d	25.8 c	22.9 e	76.2 d	22.2 c
2022	无盐 NS	浅水层灌溉 WW	14.1 a	18.5 a	92.2 a	29.1 a	34.5 a	88.2 a	24.8 a
		中度干旱 MD	13.0 b	17.2 b	91.5 ab	28.9 ab	32.2 ab	86.9 ab	24.7 a
		重度干旱 SD	12.3 bc	17.0 b	90.3 b	27.9 b	29.6 b	83.7 b	24.0 ab
	高盐 HS	浅水层灌溉 WW	11.7 c	16.6 bc	87.3 c	27.9 b	28.2 bc	81.2 bc	23.2 b
		中度干旱 MD	11.2 cd	16.5 bc	86.4 cd	27.2 bc	27.3 c	78.4 c	23.1 b
		重度干旱 SD	10.4 d	15.0 c	83.1 d	26.4 c	24.2 d	74.4 d	22.2 c
方差分析 Analysis of variance
年份 Year(Y)			**	ns	ns	ns	ns	ns	ns
盐处理 Salinity treatment(S)			**	**	**	**	**	**	**
干旱处理 Drought treatment(D)			**	**	**	**	**	**	*
年份×盐处理 Y×S			ns	*	ns	ns	ns	ns	ns
年份×干旱处理 Y×D			ns	ns	ns	ns	ns	ns	ns
盐处理×干旱处理 S×D			*	ns	ns	ns	ns	ns	ns
年份×盐处理×干旱处理 Y×S×D			ns	ns	ns	ns	ns	ns	ns

Table 3. Dry matter weight and harvest index of rice under salinity and drought stresses

年份 Year	盐处理 Salinity treatment	干旱处理 Drought treatment	干物质量 Dry matter weight (t/hm²)		抽穗期至成熟期干物重积累量 Dry weight accumulation from heading to maturity (t/hm²)	收获指数 Harvest index
年份 Year	盐处理 Salinity treatment	干旱处理 Drought treatment	抽穗期 Heading	成熟期 Maturity		收获指数 Harvest index
2021	无盐 NS	浅水层灌溉 WW	12.2 a	20.4 a	8.1 a	0.493 d
		中度干旱 MD	11.1 b	17.6 b	6.5 b	0.505 cd
		重度干旱 SD	9.5 c	14.9 c	5.4 cd	0.515 c
	高盐 HS	浅水层灌溉 WW	8.8 d	14.5 cd	5.7 c	0.508 cd
		中度干旱 MD	7.5 e	12.2 d	4.7 d	0.525 b
		重度干旱 SD	5.7 f	9.1 e	3.4 e	0.541 a
2022	无盐 NS	浅水层灌溉 WW	12.2 a	20.6 a	8.4 a	0.492 e
		中度干旱 MD	10.8 b	17.4 b	6.6 b	0.508 d
		重度干旱 SD	9.4 c	15.2 c	5.9 c	0.517 cd
	高盐 HS	浅水层灌溉 WW	9.0 cd	14.7 cd	5.7 c	0.506 d
		中度干旱 MD	7.4 d	11.9 d	4.6 d	0.522 b
		重度干旱 SD	5.7 e	9.0 e	3.3 e	0.539 a
方差分析 Analysis of variance
年份 Year(Y)			ns	ns	ns	ns
盐处理 Salinity treatment(S)			**	**	**	**
干旱处理 Drought treatment(D)			**	**	**	**
年份×盐处理 Y×S			ns	ns	ns	ns
年份×干旱处理 Y×D			ns	ns	ns	ns
盐处理×干旱处理 S×D			ns	ns	ns	ns
年份×盐处理×干旱处理 Y×S×D			ns	ns	ns	ns

Table 4. Leaf photosynthetic rate and SPAD values of rice under salinity and drought stresses

年份 Year	盐处理 Salinity treatment	干旱处理 Drought treatment	叶片光合速率 Leaf photosynthetic rate (µmol m^-2 s^-1)		叶片SPAD值 Leaf SPAD value
年份 Year	盐处理 Salinity treatment	干旱处理 Drought treatment	抽穗后20 d 20 DAH	抽穗后40 d 40 DAH	抽穗后20 d 20 DAH	抽穗后40 d 40 DAH
2021	无盐 NS	浅水层灌溉 WW	20.8 a	13.2 a	42.6 a	34.0 a
		中度干旱 MD	20.3 ab	12.2 b	41.3 b	31.0 b
		重度干旱 SD	18.6 c	10.6 d	39.0 c	26.6 d
	高盐 HS	浅水层灌溉 WW	19.4 b	11.2 c	38.6 c	28.6 c
		中度干旱 MD	18.2 cd	9.9 e	35.7 d	24.8 e
		重度干旱 SD	17.7 d	9.4 f	33.1 e	21.6 f
2022	无盐 NS	浅水层灌溉 WW	22.3 a	12.6 a	43.4 a	34.1 a
		中度干旱 MD	21.7 ab	12.1 b	42.1 b	31.3 b
		重度干旱 SD	20.9 b	10.5 d	39.0 cd	26.7 c
	高盐 HS	浅水层灌溉 WW	20.4 bc	11.3 c	40.0 c	30.2 bc
		中度干旱 MD	19.8 c	9.5 e	36.4 d	26.1 c
		重度干旱 SD	19.1 d	9.2 f	32.1 e	21.0 d
方差分析 Analysis of variance
年份 Year(Y)			**	ns	ns	ns
盐处理 Salinity treatment(S)			**	**	**	**
干旱处理 Drought treatment(D)			**	**	**	**
年份×盐处理 Y×S			ns	ns	ns	ns
年份×干旱处理 Y×D			ns	ns	ns	ns
盐处理×干旱处理 S×D			ns	ns	ns	ns
年份×盐处理×干旱处理Y×S×D			ns	ns	ns	ns

Fig. 1. Dynamics in grain weight of rice under salinity and drought stresses NS, Non-salinity; HS, High-salinity. WW, Well-watered conditions; MD, Moderate drought; SD, Severe drought. SG, Superior grains; IG, Inferior grains.

Table 5. Stimulation equations of grain-filling process of rice under salinity and drought stresses

年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	方程参数 Equation parameter				方程拟合 Simulated equation
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	A	B	K	N	方程拟合 Simulated equation
2021	强势粒	无盐 NS	浅水层灌溉 WW	24.38	6107.27	0.5664	3.0308	Y=24.38 $(1+6107.27 e -0.5664 X) - 1 3.0308$ R²=0.981
	SG		中度干旱 MD	23.79	8607.28	0.5637	3.2093	Y=23.79 $(1+8607.28 e -0.5637 X) - 1 3.2093$ R²=0.971
			重度干旱 SD	23.24	9336.26	0.4808	3.4410	Y=23.24 $(1+9336.26 e -0.4808 X) - 1 3.4410$ R²=0.972
		高盐 HS	浅水层灌溉 WW	23.03	7951.21	0.4742	3.0184	Y=23.03 $(1+7951.21 e -0.4742 X) - 1 3.0184$ R²=0.978
			中度干旱 MD	23.00	3160.03	0.3391	2.8003	Y=23.00 $(1+3160.03 e -0.3391 X) - 1 2.8003$ R²=0.989
			重度干旱 SD	21.80	4246.39	0.2936	3.1752	Y=21.80 $(1+4246.39 e -0.2936 X) - 1 3.1752$ R²=0.973
	弱势粒	无盐 NS	浅水层灌溉 WW	20.26	11274.68	0.2949	2.9366	Y=20.26 $(1+11274.68 e -0.2949 X) - 1 2.9366$ R²=0.992
	IG		中度干旱 MD	19.68	9977.78	0.2934	2.8548	Y=19.68 $(1+9977.78 e -0.2934 X) - 1 2.8548$ R²=0.992
			重度干旱 SD	19.63	10484.08	0.2878	2.8548	Y=19.63 $(1+10484.08 e -0.2878 X) - 1 2.8548$ R²=0.990
		高盐 HS	浅水层灌溉 WW	19.29	11237.63	0.2908	2.8821	Y=19.29 $(1+11237.63 e -0.2908 X) - 1 2.8821$ R²=0.992
			中度干旱 MD	18.92	10053.42	0.2898	2.8003	Y=18.92 $(1+10053.42 e -0.2898 X) - 1 2.8003$ R²=0.992
			重度干旱 SD	18.47	9707.16	0.2903	2.7730	Y=18.47 $(1+9707.16 e -0.2903 X) - 1 2.7730$ R²=0.993
2022	强势粒	无盐 NS	浅水层灌溉 WW	24.27	6107.27	0.5634	3.0593	Y=24.27 $(1+6107.27 e -0.5634 X) - 1 3.0593$ R²=0.981
	SG		中度干旱 MD	23.86	8179.72	0.5446	3.1547	Y=23.86 $(1+8179.72 e -0.5446 X) - 1 3.1547$ R²=0.973
			重度干旱 SD	23.34	7318.14	0.4809	3.1274	Y=23.34 $(1+7318.14 e -0.4809 X) - 1 3.1274$ R²=0.973
		高盐 HS	浅水层灌溉 WW	23.04	7951.20	0.4680	3.1274	Y=23.04 $(1+7951.20 e -0.4680 X) - 1 3.1274$ R²=0.970
			中度干旱 MD	23.02	2586.37	0.3154	2.8003	Y=23.02 $(1+2586.37 e -0.3154 X) - 1 2.8003$ R²=0.986
			重度干旱 SD	22.24	4265.16	0.2946	3.2638	Y=22.24 $(1+4265.16 e -0.2946 X) - 1 3.2638$ R²=0.972
	弱势粒	无盐 NS	浅水层灌溉 WW	20.59	12001.81	0.2960	2.9364	Y=20.59 $(1+12001.81 e -0.2960 X) - 1 2.9364$ R²=0.997
	IG		中度干旱 MD	20.62	12553.42	0.2910	2.9093	Y=20.62 $(1+12553.42 e -0.2910 X) - 1 2.9093$ R²=0.994
			重度干旱 SD	20.08	12553.42	0.2906	3.0184	Y=20.08 $(1+12553.42 e -0.2906 X) - 1 3.0184$ R²=0.992
		高盐 HS	浅水层灌溉 WW	19.23	11237.63	0.2908	2.8821	Y=19.23 $(1+11237.63 e -0.2908 X) - 1 2.8821$ R²=0.991
			中度干旱 MD	19.16	9957.77	0.2873	2.8412	Y=19.16 $(1+9957.77 e -0.2873 X) - 1 2.8412$ R²=0.992
			重度干旱 SD	18.60	10053.42	0.2902	2.8003	Y=18.60 $(1+10053.42 e -0.2902 X) - 1 2.8003$ R²=0.993

Table 5. Stimulation equations of grain-filling process of rice under salinity and drought stresses

年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	方程参数 Equation parameter				方程拟合 Simulated equation
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	A	B	K	N	方程拟合 Simulated equation
2021	强势粒	无盐 NS	浅水层灌溉 WW	24.38	6107.27	0.5664	3.0308	Y=24.38 $(1+6107.27 e -0.5664 X) - 1 3.0308$ R²=0.981
	SG		中度干旱 MD	23.79	8607.28	0.5637	3.2093	Y=23.79 $(1+8607.28 e -0.5637 X) - 1 3.2093$ R²=0.971
			重度干旱 SD	23.24	9336.26	0.4808	3.4410	Y=23.24 $(1+9336.26 e -0.4808 X) - 1 3.4410$ R²=0.972
		高盐 HS	浅水层灌溉 WW	23.03	7951.21	0.4742	3.0184	Y=23.03 $(1+7951.21 e -0.4742 X) - 1 3.0184$ R²=0.978
			中度干旱 MD	23.00	3160.03	0.3391	2.8003	Y=23.00 $(1+3160.03 e -0.3391 X) - 1 2.8003$ R²=0.989
			重度干旱 SD	21.80	4246.39	0.2936	3.1752	Y=21.80 $(1+4246.39 e -0.2936 X) - 1 3.1752$ R²=0.973
	弱势粒	无盐 NS	浅水层灌溉 WW	20.26	11274.68	0.2949	2.9366	Y=20.26 $(1+11274.68 e -0.2949 X) - 1 2.9366$ R²=0.992
	IG		中度干旱 MD	19.68	9977.78	0.2934	2.8548	Y=19.68 $(1+9977.78 e -0.2934 X) - 1 2.8548$ R²=0.992
			重度干旱 SD	19.63	10484.08	0.2878	2.8548	Y=19.63 $(1+10484.08 e -0.2878 X) - 1 2.8548$ R²=0.990
		高盐 HS	浅水层灌溉 WW	19.29	11237.63	0.2908	2.8821	Y=19.29 $(1+11237.63 e -0.2908 X) - 1 2.8821$ R²=0.992
			中度干旱 MD	18.92	10053.42	0.2898	2.8003	Y=18.92 $(1+10053.42 e -0.2898 X) - 1 2.8003$ R²=0.992
			重度干旱 SD	18.47	9707.16	0.2903	2.7730	Y=18.47 $(1+9707.16 e -0.2903 X) - 1 2.7730$ R²=0.993
2022	强势粒	无盐 NS	浅水层灌溉 WW	24.27	6107.27	0.5634	3.0593	Y=24.27 $(1+6107.27 e -0.5634 X) - 1 3.0593$ R²=0.981
	SG		中度干旱 MD	23.86	8179.72	0.5446	3.1547	Y=23.86 $(1+8179.72 e -0.5446 X) - 1 3.1547$ R²=0.973
			重度干旱 SD	23.34	7318.14	0.4809	3.1274	Y=23.34 $(1+7318.14 e -0.4809 X) - 1 3.1274$ R²=0.973
		高盐 HS	浅水层灌溉 WW	23.04	7951.20	0.4680	3.1274	Y=23.04 $(1+7951.20 e -0.4680 X) - 1 3.1274$ R²=0.970
			中度干旱 MD	23.02	2586.37	0.3154	2.8003	Y=23.02 $(1+2586.37 e -0.3154 X) - 1 2.8003$ R²=0.986
			重度干旱 SD	22.24	4265.16	0.2946	3.2638	Y=22.24 $(1+4265.16 e -0.2946 X) - 1 3.2638$ R²=0.972
	弱势粒	无盐 NS	浅水层灌溉 WW	20.59	12001.81	0.2960	2.9364	Y=20.59 $(1+12001.81 e -0.2960 X) - 1 2.9364$ R²=0.997
	IG		中度干旱 MD	20.62	12553.42	0.2910	2.9093	Y=20.62 $(1+12553.42 e -0.2910 X) - 1 2.9093$ R²=0.994
			重度干旱 SD	20.08	12553.42	0.2906	3.0184	Y=20.08 $(1+12553.42 e -0.2906 X) - 1 3.0184$ R²=0.992
		高盐 HS	浅水层灌溉 WW	19.23	11237.63	0.2908	2.8821	Y=19.23 $(1+11237.63 e -0.2908 X) - 1 2.8821$ R²=0.991
			中度干旱 MD	19.16	9957.77	0.2873	2.8412	Y=19.16 $(1+9957.77 e -0.2873 X) - 1 2.8412$ R²=0.992
			重度干旱 SD	18.60	10053.42	0.2902	2.8003	Y=18.60 $(1+10053.42 e -0.2902 X) - 1 2.8003$ R²=0.993

Fig. 2. Dynamics of grain-filling rate of rice under salinity and drought stresses SG, Superior grains; IG, Inferior grains. NS, Non-salinity; HS, High-salinity. WW, Well-watered conditions; MD, Moderate drought; SD, Severe drought.

Table 6. Grain-filling parameters of rice under salinity and drought stresses

年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	籽粒灌浆特征参数 Grain-filling parameters
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	最大灌浆速率 MGR_max (mg grain^-1 d^-1)	平均灌浆速率 MGR_mean (mg grain^-1 d^-1)	达最大灌浆速率的时间 T_max (d)	有效灌浆天数 EGP (d)
2021	强势粒	无盐	浅水层灌溉 WW	2.163	1.373	13.4	21.5
	SG	NS	中度干旱 MD	2.036	1.287	14.0	22.1
			重度干旱 SD	1.631	1.027	16.4	26.0
		高盐	浅水层灌溉 WW	1.714	1.088	16.6	26.3
		HS	中度干旱 MD	1.274	0.813	20.7	34.2
			重度干旱 SD	0.978	0.619	24.5	40.1
	弱势粒	无盐	浅水层灌溉 WW	0.952	0.605	28.0	43.5
	IG	NS	中度干旱 MD	0.934	0.595	27.8	43.4
			重度干旱 SD	0.914	0.582	28.5	44.4
		高盐	浅水层灌溉 WW	0.903	0.575	28.4	44.2
		HS	中度干旱 MD	0.896	0.571	28.2	44.1
			重度干旱 SD	0.880	0.562	28.1	43.9
2022	强势粒	无盐	浅水层灌溉 WW	2.131	1.351	13.5	21.6
	SG	NS	中度干旱 MD	1.992	1.261	14.4	22.8
			重度干旱 SD	1.729	1.095	16.1	25.7
		高盐	浅水层灌溉 WW	1.661	1.052	16.8	26.5
		HS	中度干旱 MD	1.186	0.756	21.6	36.2
			重度干旱 SD	0.985	0.622	24.4	39.9
	弱势粒	无盐	浅水层灌溉 WW	0.971	0.618	28.1	43.6
	IG	NS	中度干旱 MD	0.961	0.611	28.8	44.5
			重度干旱 SD	0.916	0.581	28.7	44.4
		高盐	浅水层灌溉 WW	0.900	0.573	28.4	44.2
		HS	中度干旱 MD	0.892	0.569	28.4	44.4
			重度干旱 SD	0.882	0.562	28.2	44.0

Table 7. Grain-filling characteristics of early, middle and late stages of rice under salinity and drought stresses

年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	前期 Early stage			中期 Middle stage
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	天数 Days (d)	平均灌浆速率 MGR_mean (mg grain^-1 d^-1)	灌浆量 GFA (mg)	天数 Days (d)	平均灌浆速率 MGR_mean (mg grain^-1 d^-1)	灌浆量 GFA (mg)
2021	强势粒 SG	无盐 NS	浅水层灌溉 WW	10.3	0.899	178.9	6.2	1.919	231.3
			中度干旱 MD	10.8	0.860	163.6	6.4	1.808	203.0
			重度干旱 SD	12.6	0.744	158.7	7.6	1.449	187.2
		高盐 HS	浅水层灌溉 WW	12.9	0.678	137.1	7.4	1.521	177.8
			中度干旱 MD	15.6	0.538	124.5	10.2	1.130	170.2
			重度干旱 SD	18.4	0.461	115.3	12.2	0.868	144.2
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	22.0	0.344	265.2	11.9	0.844	350.7
			中度干旱 MD	21.9	0.332	230.8	11.8	0.828	311.4
			重度干旱 SD	22.5	0.322	214.3	12.1	0.810	289.2
		高盐 HS	浅水层灌溉 WW	22.4	0.319	202.3	12.0	0.800	271.1
			中度干旱 MD	22.3	0.310	183.0	11.9	0.794	250.4
			重度干旱 SD	22.2	0.303	154.1	11.9	0.780	212.2
2022	强势粒 SG	无盐 NS	浅水层灌溉 WW	10.3	0.896	171.1	6.3	1.891	219.8
			中度干旱 MD	11.1	0.830	159.5	6.6	1.768	200.4
			重度干旱 SD	12.4	0.725	153.3	7.4	1.534	193.8
		高盐 HS	浅水层灌溉 WW	12.9	0.687	147.4	7.6	1.474	186.3
			中度干旱 MD	16.2	0.520	138.7	10.9	1.051	189.7
			重度干旱 SD	18.2	0.481	131.4	12.3	0.875	161.0
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	22.2	0.348	266.0	11.8	0.862	351.7
			中度干旱 MD	22.8	0.338	247.3	12.0	0.852	329.1
			重度干旱 SD	22.6	0.337	225.4	12.2	0.813	292.3
		高盐 HS	浅水层灌溉 WW	22.4	0.318	201.2	12.0	0.798	269.7
			中度干旱 MD	22.4	0.315	192.9	12.1	0.791	261.1
			重度干旱 SD	22.3	0.306	164.3	11.9	0.782	224.7
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	后期 Late stage
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	天数 Days (d)		平均灌浆速率 MGR_mean(mg grain^-1 d^-1)		灌浆量 GFA (mg)
2021	强势粒 SG	无盐 NS	浅水层灌溉 WW	5.0		0.581		55.8
			中度干旱 MD	4.9		0.550		47.9
			重度干旱 SD	5.7		0.445		42.9
		高盐 HS	浅水层灌溉 WW	5.9		0.460		43.0
			中度干旱 MD	8.4		0.339		42.4
			重度干旱 SD	9.5		0.264		34.1
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	9.6		0.255		85.6
			中度干旱 MD	9.7		0.249		76.9
			重度干旱 SD	9.9		0.244		71.4
		高盐 HS	浅水层灌溉 WW	9.8		0.241		66.7
			中度干旱 MD	9.9		0.239		62.3
			重度干旱 SD	9.9		0.234		53.0
2022	强势粒 SG	无盐 NS	浅水层灌溉 WW	5.0		0.573		52.8
			中度干旱 MD	5.1		0.537		47.5
			重度干旱 SD	5.8		0.466		46.1
		高盐 HS	浅水层灌溉 WW	6.0		0.448		44.4
			中度干旱 MD	9.1		0.316		47.2
			重度干旱 SD	9.4		0.267		37.7
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	9.6		0.260		85.9
			中度干旱 MD	9.8		0.257		80.7
			重度干旱 SD	9.7		0.246		70.6
		高盐 HS	浅水层灌溉 WW	9.8		0.240		66.3
			中度干旱 MD	9.9		0.238		64.6
			重度干旱 SD	9.9		0.235		55.9

Table 7. Grain-filling characteristics of early, middle and late stages of rice under salinity and drought stresses

年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	前期 Early stage			中期 Middle stage
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	天数 Days (d)	平均灌浆速率 MGR_mean (mg grain^-1 d^-1)	灌浆量 GFA (mg)	天数 Days (d)	平均灌浆速率 MGR_mean (mg grain^-1 d^-1)	灌浆量 GFA (mg)
2021	强势粒 SG	无盐 NS	浅水层灌溉 WW	10.3	0.899	178.9	6.2	1.919	231.3
			中度干旱 MD	10.8	0.860	163.6	6.4	1.808	203.0
			重度干旱 SD	12.6	0.744	158.7	7.6	1.449	187.2
		高盐 HS	浅水层灌溉 WW	12.9	0.678	137.1	7.4	1.521	177.8
			中度干旱 MD	15.6	0.538	124.5	10.2	1.130	170.2
			重度干旱 SD	18.4	0.461	115.3	12.2	0.868	144.2
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	22.0	0.344	265.2	11.9	0.844	350.7
			中度干旱 MD	21.9	0.332	230.8	11.8	0.828	311.4
			重度干旱 SD	22.5	0.322	214.3	12.1	0.810	289.2
		高盐 HS	浅水层灌溉 WW	22.4	0.319	202.3	12.0	0.800	271.1
			中度干旱 MD	22.3	0.310	183.0	11.9	0.794	250.4
			重度干旱 SD	22.2	0.303	154.1	11.9	0.780	212.2
2022	强势粒 SG	无盐 NS	浅水层灌溉 WW	10.3	0.896	171.1	6.3	1.891	219.8
			中度干旱 MD	11.1	0.830	159.5	6.6	1.768	200.4
			重度干旱 SD	12.4	0.725	153.3	7.4	1.534	193.8
		高盐 HS	浅水层灌溉 WW	12.9	0.687	147.4	7.6	1.474	186.3
			中度干旱 MD	16.2	0.520	138.7	10.9	1.051	189.7
			重度干旱 SD	18.2	0.481	131.4	12.3	0.875	161.0
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	22.2	0.348	266.0	11.8	0.862	351.7
			中度干旱 MD	22.8	0.338	247.3	12.0	0.852	329.1
			重度干旱 SD	22.6	0.337	225.4	12.2	0.813	292.3
		高盐 HS	浅水层灌溉 WW	22.4	0.318	201.2	12.0	0.798	269.7
			中度干旱 MD	22.4	0.315	192.9	12.1	0.791	261.1
			重度干旱 SD	22.3	0.306	164.3	11.9	0.782	224.7
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	后期 Late stage
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	天数 Days (d)		平均灌浆速率 MGR_mean(mg grain^-1 d^-1)		灌浆量 GFA (mg)
2021	强势粒 SG	无盐 NS	浅水层灌溉 WW	5.0		0.581		55.8
			中度干旱 MD	4.9		0.550		47.9
			重度干旱 SD	5.7		0.445		42.9
		高盐 HS	浅水层灌溉 WW	5.9		0.460		43.0
			中度干旱 MD	8.4		0.339		42.4
			重度干旱 SD	9.5		0.264		34.1
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	9.6		0.255		85.6
			中度干旱 MD	9.7		0.249		76.9
			重度干旱 SD	9.9		0.244		71.4
		高盐 HS	浅水层灌溉 WW	9.8		0.241		66.7
			中度干旱 MD	9.9		0.239		62.3
			重度干旱 SD	9.9		0.234		53.0
2022	强势粒 SG	无盐 NS	浅水层灌溉 WW	5.0		0.573		52.8
			中度干旱 MD	5.1		0.537		47.5
			重度干旱 SD	5.8		0.466		46.1
		高盐 HS	浅水层灌溉 WW	6.0		0.448		44.4
			中度干旱 MD	9.1		0.316		47.2
			重度干旱 SD	9.4		0.267		37.7
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	9.6		0.260		85.9
			中度干旱 MD	9.8		0.257		80.7
			重度干旱 SD	9.7		0.246		70.6
		高盐 HS	浅水层灌溉 WW	9.8		0.240		66.3
			中度干旱 MD	9.9		0.238		64.6
			重度干旱 SD	9.9		0.235		55.9

Table 8. Key enzymes activities involved in starch synthesis of rice grains under salinity and drought stresses

年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	AGPase酶活性 Activity of AGPase (mol min^-1 mg^-1)			GBSS酶活性 Activity of GBSS (mol min^-1 mg^-1)
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	15 DAH	30 DAH	45 DAH	15 DAH	30 DAH	45 DAH
2021	强势粒 SG	无盐 NS	浅水层灌溉 WW	16.3 a	44.6 a	13.9 a	6.9 a	17.2 a	5.6 a
			中度干旱 MD	14.9 b	40.7 b	13.1 b	6.6 ab	15.6 b	5.1 ab
			重度干旱 SD	13.2 c	39.0 bc	12.6 bc	6.2 b	13.5 cd	4.5 b
		高盐 HS	浅水层灌溉 WW	13.0 c	36.3 c	13.1 b	6.6 ab	13.9 c	4.2 c
			中度干旱 MD	11.5 d	34.2 cd	11.3 c	5.6 c	12.0 d	3.6 cd
			重度干旱 SD	10.2 e	31.4 d	10.2 d	4.1 d	10.1 e	2.4 d
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	12.1 a	32.3 a	10.6 a	4.3 a	12.7 a	4.4 a
			中度干旱 MD	11.2 b	30.2 b	10.0 b	4.1 ab	11.8 b	3.8 b
			重度干旱 SD	9.6 cd	28.9 c	9.4 c	3.7 bc	10.9 c	3.3 bc
		高盐 HS	浅水层灌溉 WW	10.0 c	29.0 c	10.0 b	4.1 ab	10.4 cd	3.2 bc
			中度干旱 MD	8.6 d	26.8 d	7.9 d	3.3 bc	9.1 d	2.1 c
			重度干旱 SD	7.9 e	22.6 e	6.9 e	2.5 c	7.9 e	1.3 d
2022	强势粒 SG	无盐 NS	浅水层灌溉 WW	15.3 a	48.4 a	14.5 a	5.9 a	17.4 a	6.0 a
			中度干旱 MD	14.2 b	45.2 b	14.1 ab	5.5 ab	16.2 b	5.5 b
			重度干旱 SD	12.3 cd	44.6 bc	13.2 c	4.7 b	13.9 c	4.9 c
		高盐 HS	浅水层灌溉 WW	12.7 c	43.4 c	13.7 b	4.5 b	14.2 c	4.8 c
			中度干旱 MD	11.2 d	41.3 d	11.6 d	3.5 c	12.8 d	3.6 d
			重度干旱 SD	10.3 e	38.1 e	9.3 e	2.4 d	11.2 e	2.4 e
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	11.7 a	35.7 a	11.1 a	4.5 a	13.1 a	4.1 a
			中度干旱 MD	10.7 b	32.5 b	10.8 ab	4.1 ab	11.9 b	3.8 ab
			重度干旱 SD	9.2 c	31.3 c	9.8 c	3.4 b	11.1 bc	3.5 b
		高盐 HS	浅水层灌溉 WW	9.3 c	29.0 d	10.5 b	3.3 b	10.6 cd	3.3 bc
			中度干旱 MD	8.6 cd	26.4 e	8.4 d	2.8 c	9.2 d	2.5 c
			重度干旱 SD	7.8 d	23.5 f	7.1 e	2.1 d	7.6 e	1.3 d
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	SSS酶活性 Activity of SSS(mol min^-1 mg^-1)			SBE酶活性 Activity of SBE(U g^-1)
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	15 DAH	30 DAH	45 DAH	15 DAH	30 DAH	45 DAH
2021	强势粒 SG	无盐 NS	浅水层灌溉 WW	3.4 a	6.5 a	2.2 a	4.5 a	7.1 a	3.1 a
			中度干旱 MD	3.3 a	5.9 b	2.0 ab	4.5 a	6.7 ab	2.8 ab
			重度干旱 SD	3.3 a	4.7 c	1.9 b	4.4 a	5.9 b	2.6 ab
		高盐 HS	浅水层灌溉 WW	3.2 ab	5.2 bc	1.8 b	4.3 a	5.8 b	2.5 ab
			中度干旱 MD	3.2 ab	4.5 c	1.7 bc	4.2 a	4.7 c	2.3 b
			重度干旱 SD	3.1 b	3.8 d	1.5 c	4.1 a	3.6 d	2.0 b
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	2.2 a	4.2 a	1.5 a	3.1 a	4.8 a	2.1 a
			中度干旱 MD	2.2 a	3.7 b	1.4 a	3.1 a	4.5 ab	2.0 a
			重度干旱 SD	2.1 a	3.1 cd	1.4 a	3.0 a	3.8 b	1.8 a
		高盐 HS	浅水层灌溉 WW	2.1 a	3.3 c	1.3 a	2.9 a	4.3 ab	1.7 ab
			中度干旱 MD	2.1 a	2.5 d	1.3 a	2.9 a	3.5 bc	1.7 ab
			重度干旱 SD	2.0 a	1.7 e	1.2 a	2.8 a	2.6 c	1.5 b
2022	强势粒 SG	无盐 NS	浅水层灌溉 WW	3.3 a	6.3 a	2.5 a	5.1 a	7.8 a	3.4 a
			中度干旱 MD	3.3 a	5.9 ab	2.3 a	5.1 a	7.5 ab	3.2 a
			重度干旱 SD	3.2 a	5.5 b	2.3 a	5.0 a	6.7 b	2.9 ab
		高盐 HS	浅水层灌溉 WW	3.1 a	5.2 bc	2.2 a	4.8 ab	6.5 b	2.8 ab
			中度干旱 MD	3.1 a	4.6 c	2.2 a	4.8 ab	5.3 c	2.7 ab
			重度干旱 SD	3.0 a	3.6 d	2.0 a	4.6 b	4.1 d	2.4 b
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	2.4 a	4.4 a	1.7 a	3.4 a	5.1 a	2.3 a
			中度干旱 MD	2.3 a	4.0 ab	1.5 a	3.4 a	4.8 ab	2.1 a
			重度干旱 SD	2.3 a	3.5 b	1.5 a	3.3 a	4.4 b	1.9 a
		高盐 HS	浅水层灌溉 WW	2.3 a	3.6 b	1.4 a	3.2 a	3.8 c	1.9 a
			中度干旱 MD	2.2 a	2.9 c	1.3 a	3.2 a	3.4 cd	1.7 ab
			重度干旱 SD	2.2 a	2.3 d	1.2 a	3.1 a	2.5 d	1.5 b

Table 8. Key enzymes activities involved in starch synthesis of rice grains under salinity and drought stresses

年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	AGPase酶活性 Activity of AGPase (mol min^-1 mg^-1)			GBSS酶活性 Activity of GBSS (mol min^-1 mg^-1)
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	15 DAH	30 DAH	45 DAH	15 DAH	30 DAH	45 DAH
2021	强势粒 SG	无盐 NS	浅水层灌溉 WW	16.3 a	44.6 a	13.9 a	6.9 a	17.2 a	5.6 a
			中度干旱 MD	14.9 b	40.7 b	13.1 b	6.6 ab	15.6 b	5.1 ab
			重度干旱 SD	13.2 c	39.0 bc	12.6 bc	6.2 b	13.5 cd	4.5 b
		高盐 HS	浅水层灌溉 WW	13.0 c	36.3 c	13.1 b	6.6 ab	13.9 c	4.2 c
			中度干旱 MD	11.5 d	34.2 cd	11.3 c	5.6 c	12.0 d	3.6 cd
			重度干旱 SD	10.2 e	31.4 d	10.2 d	4.1 d	10.1 e	2.4 d
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	12.1 a	32.3 a	10.6 a	4.3 a	12.7 a	4.4 a
			中度干旱 MD	11.2 b	30.2 b	10.0 b	4.1 ab	11.8 b	3.8 b
			重度干旱 SD	9.6 cd	28.9 c	9.4 c	3.7 bc	10.9 c	3.3 bc
		高盐 HS	浅水层灌溉 WW	10.0 c	29.0 c	10.0 b	4.1 ab	10.4 cd	3.2 bc
			中度干旱 MD	8.6 d	26.8 d	7.9 d	3.3 bc	9.1 d	2.1 c
			重度干旱 SD	7.9 e	22.6 e	6.9 e	2.5 c	7.9 e	1.3 d
2022	强势粒 SG	无盐 NS	浅水层灌溉 WW	15.3 a	48.4 a	14.5 a	5.9 a	17.4 a	6.0 a
			中度干旱 MD	14.2 b	45.2 b	14.1 ab	5.5 ab	16.2 b	5.5 b
			重度干旱 SD	12.3 cd	44.6 bc	13.2 c	4.7 b	13.9 c	4.9 c
		高盐 HS	浅水层灌溉 WW	12.7 c	43.4 c	13.7 b	4.5 b	14.2 c	4.8 c
			中度干旱 MD	11.2 d	41.3 d	11.6 d	3.5 c	12.8 d	3.6 d
			重度干旱 SD	10.3 e	38.1 e	9.3 e	2.4 d	11.2 e	2.4 e
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	11.7 a	35.7 a	11.1 a	4.5 a	13.1 a	4.1 a
			中度干旱 MD	10.7 b	32.5 b	10.8 ab	4.1 ab	11.9 b	3.8 ab
			重度干旱 SD	9.2 c	31.3 c	9.8 c	3.4 b	11.1 bc	3.5 b
		高盐 HS	浅水层灌溉 WW	9.3 c	29.0 d	10.5 b	3.3 b	10.6 cd	3.3 bc
			中度干旱 MD	8.6 cd	26.4 e	8.4 d	2.8 c	9.2 d	2.5 c
			重度干旱 SD	7.8 d	23.5 f	7.1 e	2.1 d	7.6 e	1.3 d
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	SSS酶活性 Activity of SSS(mol min^-1 mg^-1)			SBE酶活性 Activity of SBE(U g^-1)
年份 Year	粒位 Grain position	盐处理 Salinity treatment	干旱处理 Drought treatment	15 DAH	30 DAH	45 DAH	15 DAH	30 DAH	45 DAH
2021	强势粒 SG	无盐 NS	浅水层灌溉 WW	3.4 a	6.5 a	2.2 a	4.5 a	7.1 a	3.1 a
			中度干旱 MD	3.3 a	5.9 b	2.0 ab	4.5 a	6.7 ab	2.8 ab
			重度干旱 SD	3.3 a	4.7 c	1.9 b	4.4 a	5.9 b	2.6 ab
		高盐 HS	浅水层灌溉 WW	3.2 ab	5.2 bc	1.8 b	4.3 a	5.8 b	2.5 ab
			中度干旱 MD	3.2 ab	4.5 c	1.7 bc	4.2 a	4.7 c	2.3 b
			重度干旱 SD	3.1 b	3.8 d	1.5 c	4.1 a	3.6 d	2.0 b
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	2.2 a	4.2 a	1.5 a	3.1 a	4.8 a	2.1 a
			中度干旱 MD	2.2 a	3.7 b	1.4 a	3.1 a	4.5 ab	2.0 a
			重度干旱 SD	2.1 a	3.1 cd	1.4 a	3.0 a	3.8 b	1.8 a
		高盐 HS	浅水层灌溉 WW	2.1 a	3.3 c	1.3 a	2.9 a	4.3 ab	1.7 ab
			中度干旱 MD	2.1 a	2.5 d	1.3 a	2.9 a	3.5 bc	1.7 ab
			重度干旱 SD	2.0 a	1.7 e	1.2 a	2.8 a	2.6 c	1.5 b
2022	强势粒 SG	无盐 NS	浅水层灌溉 WW	3.3 a	6.3 a	2.5 a	5.1 a	7.8 a	3.4 a
			中度干旱 MD	3.3 a	5.9 ab	2.3 a	5.1 a	7.5 ab	3.2 a
			重度干旱 SD	3.2 a	5.5 b	2.3 a	5.0 a	6.7 b	2.9 ab
		高盐 HS	浅水层灌溉 WW	3.1 a	5.2 bc	2.2 a	4.8 ab	6.5 b	2.8 ab
			中度干旱 MD	3.1 a	4.6 c	2.2 a	4.8 ab	5.3 c	2.7 ab
			重度干旱 SD	3.0 a	3.6 d	2.0 a	4.6 b	4.1 d	2.4 b
	弱势粒 IG	无盐 NS	浅水层灌溉 WW	2.4 a	4.4 a	1.7 a	3.4 a	5.1 a	2.3 a
			中度干旱 MD	2.3 a	4.0 ab	1.5 a	3.4 a	4.8 ab	2.1 a
			重度干旱 SD	2.3 a	3.5 b	1.5 a	3.3 a	4.4 b	1.9 a
		高盐 HS	浅水层灌溉 WW	2.3 a	3.6 b	1.4 a	3.2 a	3.8 c	1.9 a
			中度干旱 MD	2.2 a	2.9 c	1.3 a	3.2 a	3.4 cd	1.7 ab
			重度干旱 SD	2.2 a	2.3 d	1.2 a	3.1 a	2.5 d	1.5 b

Table 9. Correlation analysis between grain yield, biomass weight at maturity and grain-filling characteristics of rice under salinity and drought stresses

项目 Item			产量 Grain yield	成熟期干物质量 Dry matter weight at maturity
单位面积穗数 NP			0.99**	0.99**
每穗粒数 SPP			0.98**	0.98**
结实率 SSR			0.97**	0.96**
千粒重 1000-GW			0.98**	0.98**
抽穗期干物重 Dry matter weight at heading			0.99**	0.99**
抽穗期至成熟期干物重积累量 Dry matter accumulation from heading to maturity			0.98**	0.99**
收获指数 Harvest index			−0.95**	−0.96**
强势粒 SG		最大灌浆速率 MGR_max	0.98**	0.97**
		平均灌浆速率 MGR_mean	0.98**	0.98**
		达最大灌浆速率的时间T_max	−0.96**	−0.95**
		有效灌浆天数 EGP	−0.95**	−0.94**
弱势粒 IG		最大灌浆速率 MGR_max	0.93**	0.93**
		平均灌浆速率 MGR_mean	0.93**	0.92**
		达最大灌浆速率的时间T_max	−0.07	−0.09
		有效灌浆天数 EGP	−0.37	−0.39
强势粒 SG	前期 Early stage	天数Days	−0.97**	−0.96**
		平均灌浆速率 MGR_mean	0.97**	0.97**
		灌浆量 GFA	0.93**	0.93**
	中期 Middle stage	天数Days	−0.94**	−0.93**
		平均灌浆速率 MGR_mean	0.98**	0.97**
		灌浆量 GFA	0.94**	0.94**
	后期 Late stage	天数Days	−0.92**	−0.91**
		平均灌浆速率 MGR_mean	0.98**	0.97**
		灌浆量 GFA	0.90**	0.90**
弱势粒 IG	前期 Early stage	天数Days	−0.14	−0.16
		平均灌浆速率 MGR_mean	0.95**	0.94**
		灌浆量 GFA	0.98**	0.98**
	中期 Middle stage	天数Days	−0.18	−0.20
		平均灌浆速率 MGR_mean	0.93**	0.93**
		灌浆量 GFA	0.98**	0.98**
	后期 Late stage	天数Days	−0.86**	−0.87**
		平均灌浆速率 MGR_mean	0.94**	0.93**
		灌浆量 GFA	0.98**	0.98**

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[32]	Prathap V, Ali K, Singh A, Vishwakarma C, Krishnan V, Chinnusamy V, Tyagi A. Starch accumulation in rice grains subjected to drought during grain filling stage[J]. Plant Physiology and Biochemistry, 2019, 142: 440-451.

Grain-filling Characteristics and Its Relationship with Grain Yield Formation of japonica Rice Nanjing 9108 Under Combined Salinity-drought Stress

盐−旱复合胁迫下粳稻品种南粳9108籽粒灌浆特性及其与产量形成的关系

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