水稻分蘖期干旱锻炼对幼穗分化期高温下穗发育和产量形成的影响

doi:10.16819/j.1001-7216.2024.231110

摘要/Abstract

摘要：

【目的】明确分蘖期干旱锻炼对幼穗分化期高温下水稻穗发育和产量形成的影响，并探究其生理机理。【方法】以两优培九(热敏感型)和汕优63(耐热型)为材料，在盆栽条件下设置全生育期淹灌和分蘖期干旱锻炼、幼穗分化期适温和高温共四种水分和温度处理组合，研究干旱锻炼对高温胁迫下水稻颖花分化与退化、花粉活力、颖花育性、颖花大小、产量及穗生理特性的影响。【结果】与淹灌相比，在适温下干旱锻炼对两个品种产量无显著影响，但高温下干旱锻炼后两优培九和汕优63的产量分别显著提高了54.0%和20.1%，这主要是因为结实率和千粒重显著提高。与淹灌相比，幼穗分化期高温下，分蘖期干旱锻炼显著增强两优培九颖花过氧化物酶和超氧化物歧化酶活性，降低了丙二醛含量，其花粉活力和颖花育性分别显著提高了26.0%和39.0%。与适温相比，全生育期淹灌下，高温显著降低了两个品种的颖花大小和总颖花分化数。与淹灌相比，分蘖期干旱锻炼显著提高了幼穗分化期高温下两品种穗非结构性碳水化合物含量和颖花细胞分裂素(反式玉米素+反式玉米素核苷)含量，两个品种颖花长、颖花宽和总颖花分化数平均显著增加了3.2%、4.4%和15.0%。【结论】分蘖期干旱锻炼可通过增加抗氧化酶活性、非结构性碳水化合物和细胞分裂素含量来促进颖花发育和提高颖花育性，进而缓解幼穗分化期高温对水稻产量的不利影响，研究结果可为高温热害频发地区的水稻生产实践提供理论指导。

关键词: 干旱锻炼, 幼穗分化期高温, 颖花育性, 颖花发育, 产量

Abstract:

【Objective】 The aim of this study was to investigate the impact of drought priming during the tillering stage on rice panicle development and yield formation under high temperatures during the panicle initiation stage, along with the underlying physiological mechanisms.【Method】 Two rice varieties, Liangyoupeijiu (heat-sensitive) and Shanyou 63 (heat-tolerant), were chosen for the study. A pot experiment was conducted with two water treatments (flooding irrigation and drought priming during tillering stage) and two temperature treatments (normal temperature and high temperature during panicle initiation stage). The effects of drought priming on spikelet differentiation and degradation, pollen viability, spikelet fertility, spikelet size, grain yield, and underlying physiological characteristics were evaluated.【Result】 Under normal temperature treatment, there was no significant difference in grain yield between flooding irrigation and drought priming for both varieties. However, under high temperature treatment, drought priming led to a significant increase in seed setting rate and 1000-grain weight, resulting in a 54.0% increase in grain yield for Liangyoupeijiu and a 20.1% increase for Shanyou 63 compared to flooding irrigation. Drought priming also resulted in the significant increases in the activities of peroxidase and superoxide dismutase and significant decrease in malondialdehyde content of spikelet, and significant increase in the pollen viability by 26.0%, and in spikelet fertility by 39.0% in Liangyoupeijiu under high temperature treatment. Under flooding irrigation, high temperature treatment resulted in significant reductions in spikelet size and differentiated spikelet number for both varieties compared to normal temperature treatment. However, under high temperature treatment, drought priming led to significant increases in the contents of panicle non-structural carbohydrates and spikelet cytokinins (trans-zeatin and trans-zeatin-riboside), as well as increases in spikelet length, width, and differentiated spikelet number across both varieties.【Conclusion】 Drought priming during the tillering stage effectively mitigated the negative effects of high temperature stress on spikelet development and fertility by enhancing antioxidant enzyme activities, cytokinin content in spikelets, and non-structural carbohydrate levels in panicles. This resulted in improved rice yield formation under high temperature stress conditions, offering insights for rice production in regions prone to high temperature stress.

Key words: drought priming, high temperature during panicle initiation stage, spikelet fertility, spikelet development, grain yield

赵艺婷, 谢可冉, 高逖, 崔克辉. 水稻分蘖期干旱锻炼对幼穗分化期高温下穗发育和产量形成的影响[J]. 中国水稻科学, 2024, 38(3): 277-289.

ZHAO Yiting, XIE Keran, GAO Ti, CUI Kehui. Effects of Drought Priming During Tillering Stage on Panicle Development and Yield Formation Under High Temperature During Panicle Initiation Stage in Rice[J]. Chinese Journal OF Rice Science, 2024, 38(3): 277-289.

图/表 10

图1 干旱锻炼处理期间的土壤水势数据为均值 ± 标准误 (n=3)。LYPJ：两优培九；SY63：汕优63。W1：淹灌；W2：干旱锻炼。下同。

Fig. 1. Soil water potential during drought priming treatment Data are shown as mean ± SE (n=3). LYPJ, Liangyoupeijiu; SY63, Shanyou 63. W1, Flooding irrigation; W2, Drought priming. The same as below.

图2 高温处理期间温室的平均温度和湿度数值为高温处理期间各时间点的均值 ± 标准误 (n=4)。T1：适温处理；T2：高温处理。

Fig. 2. Average daytime temperature and relative humidity in the greenhouse during high temperature treatment The data at each point-in-time are the average values over the duration of the high temperature treatment ± SE (n=4). T1, Normal temperature; T2, High temperature.

表1 干旱锻炼对幼穗分化期高温下水稻产量的影响

Table 1. Effects of drought priming on rice yield under high temperature during panicle initiation stage

品种 Variety	处理 Treatment	产量 Yield(g/plant)	单株穗数 Panicles per plant	每穗粒数 Grains per panicle	结实率 Seed setting rate(%)	千粒重 1000-grain weight(g)
两优培九	W1T1	24.6±0.1 a	11.4±0.6 a	133.7±2.8 a	69.0±1.7 a	23.6±0.1 a
Liangyoupeijiu	W2T1	24.4±1.0 a	11.9±0.5 a	133.2±2.0 a	64.7±2.1 a	23.9±0.3 a
	W1T2	15.5±1.1 b	11.0±0.2 a	131.6±5.6 a	48.5±3.1 b	22.3±0.2 b
	W2T2	23.9±1.6 a	12.0±0.7 a	131.9±11.6 a	66.2±1.8 a	23.3±0.3 a
汕优63	W1T1	35.5±0.4 a*	12.3±0.4 a	133.9±5.8 ab	84.6±3.5 a*	25.6±0.1 a*
Shanyou 63	W2T1	34.6±1.9 a*	13.3±0.2 a	142.5±8.3 a	73.4±1.5 b*	25.2±0.1 a*
	W1T2	25.9±1.2 b*	12.3±0.4 a	119.7±3.1 b	78.7±3.5 ab*	22.6±0.1 c
	W2T2	31.2±1.2 a*	13.4±0.6 a	128.1±2.3 ab	78.4±1.4 ab*	23.5±0.2 b
ANOVA
品种 Variety(V)		++	ns	ns	++	++
温度 Temperature(T)		++	ns	ns	ns	++
水分 Water content(W)		+	++	ns	ns	+
V×T		ns	ns	ns	ns	++
V×W		ns	ns	ns	++	ns
T×W		++	ns	ns	++	+
V×T×W		ns	ns	ns	ns	ns

图3 干旱锻炼对幼穗分化期高温下水稻花粉活力和颖花育性的影响数据为均值 ± 标准误 (n=3)。不同小写字母表示同一品种在不同水分和温度处理组合下同一性状在P < 0.05水平下差异显著(LSD法)。*表示相同水分与温度处理组合下两个品种间同一性状在P < 0.05水平下差异显著。V表示品种，T表示温度，W表示水分。+和++分别表示在P < 0.05和P < 0.01水平下差异显著，ns表示在P < 0.05水平下差异不显著。W1T1表示淹灌和适温，W2T1表示干旱锻炼和适温，W1T2表示淹灌和高温，W2T2表示干旱锻炼和高温。

Fig. 3. Effects of drought priming on rice pollen viability and spikelet fertility under high temperature during panicle initiation stage Data are shown as mean ± SE (n=3). Different lowercase letters indicate significant difference for a given trait among the four water and temperature treatment combinations for the same variety at P<0.05 (LSD test). * indicates the significant difference for a given trait between two varieties in the same combination of water and temperature treatment at P<0.05. V indicates variety, T indicates temperature, and W indicates water. + and ++ indicate significant difference at P <0.05 and P <0.01, respectively, while ns indicates no significant difference at P <0.05. W1T1 indicates flooded irrigation and normal temperature, W2T1 indicates drought priming and normal temperature, W1T2 indicates flooded irrigation and high temperature, and W2T2 indicates drought priming and high temperature.

表2 干旱锻炼对幼穗分化期高温下颖花大小和籽粒大小的影响

Table 2. Effects of drought priming on rice spikelet size and grain size under high temperature during panicle initiation stage

品种 Variety	处理 Treatment	抽穗期Heading stage		成熟期Mature stage
品种 Variety	处理 Treatment	颖花长 Spikelet length(mm)	颖花宽 Spikelet width(mm)	籽粒长 Grain length(mm)	籽粒宽 Grain width(mm)
两优培九	W1T1	9.07±0.02 a*	3.00±0.03 a	8.85±0.05 a*	2.62±0.05 b
Liangyoupeijiu	W2T1	8.97±0.00 a*	2.98±0.04 a	8.82±0.02 ab*	2.69±0.02 a
	W1T2	8.77±0.07 b*	2.82±0.02 b	8.72±0.03 b*	2.54±0.03 c
	W2T2	9.00±0.13 a*	2.96±0.07 a	8.80±0.08 ab*	2.63±0.02 b
汕优63	W1T1	8.40±0.11 a	3.28±0.04 a*	8.44±0.06 a	2.96±0.00 a*
Shanyou 63	W2T1	8.37±0.02 a	3.22±0.01 a*	8.48±0.04 a	2.94±0.02 a*
	W1T2	7.98±0.01 b	2.97±0.02 c*	8.15±0.07 c	2.78±0.01 c*
	W2T2	8.28±0.03 a	3.09±0.01 b*	8.32±0.09 b	2.87±0.01 b*
ANOVA
品种 Variety(V)		++	++	++	++
温度 Temperature(T)		+	++	+	+
水分 Water content(W)		ns	ns	ns	++
V×T		ns	ns	ns	ns
V×W		ns	ns	ns	+
T×W		+	+	ns	++
V×T×W		ns	ns	ns	+

表3 干旱锻炼对幼穗分化期高温下每穗枝梗数的影响

Table 3. Effects of drought priming on the number of rachis branches per panicle under high temperature during panicle initiation stage

品种 Variety	处理 Treatment	总枝梗分化数 NBD (No./panicle)	一次枝梗分化数 NPBD (No./panicle)	二次枝梗分化数 NSBD (No./panicle)	一次枝梗退化数 NPBG (No./panicle)	二次枝梗退化数 NSBG (No./panicle)
两优培九	W1T1	73.7±2.3 a	12.8±0.4 a	60.9±1.9 a	0.0±0.0 a	22.3±2.3 ab
Liangyoupeijiu	W2T1	68.0±1.6 a	12.3±0.2 a	55.7±1.5 a	0.0±0.0 a	18.9±1.6 ab
	W1T2	66.8±1.5 a	12.1±0.1 a	54.7±1.4 a	0.0±0.0 a	23.8±1.7 a
	W2T2	68.0±1.9 a	12.2±0.1 a	55.8±1.8 a	0.0±0.0 a	18.6±1.1 b
汕优63	W1T1	67.3±0.7 a	14.0±0.3 a	53.3±1.0 a*	0.0±0.0 a	17.7±1.2 a
Shanyou 63	W2T1	67.0±2.1 a	13.2±0.7 a	53.8±1.5 a	0.0±0.0 a	17.2±3.6 a
	W1T2	64.4±0.2 a	13.4±0.1 a*	51.0±0.3 a	0.0±0.0 a	22.4±1.3 a
	W2T2	65.8±1.0 a	13.1±0.1 a*	52.7±1.1 a	0.0±0.0 a	17.9±0.9 a
ANOVA
品种 Variety(V)		ns	ns	+	ns	ns
温度 Temperature(T)		+	ns	+	ns	ns
水分 Water content(W)		ns	ns	ns	ns	ns
V×T		ns	ns	ns	ns	ns
V×W		ns	ns	ns	ns	ns
T×W		ns	ns	ns	ns	ns
V×T×W		ns	ns	ns	ns	ns

表4 干旱锻炼对幼穗分化期高温下每穗颖花数的影响

Table 4. Effects of drought priming on spikelets per panicle under high temperature during panicle initiation stage

品种 Variety	处理 Treatment	总颖花分化数 NSD (No./panicle)	一次颖花分化数 NPSD (No./panicle)	二次颖花分化数 NSSD (No./panicle)	一次颖花退化数 NPSG (No./panicle)	二次颖花退化数 NSSG (No./panicle)
两优培九	W1T1	189.4±5.5 a	69.6±2.9 a	119.9±4.1 a	0.0±0.0 a	7.4±0.9 b*
Liangyoupeijiu	W2T1	187.7±1.3 a	69.6±1.7 a	118.1±2.1 a	0.0±0.0 a	7.6±3.9 b
	W1T2	162.3±1.9 b	66.0±0.8 a	96.3±2.2 b*	0.1±0.1 a	16.4±3.5 a*
	W2T2	182.9±3.2 a	67.0±1.3 a	115.9±2.5 a	0.1±0.1 a	8.0±1.8 b
汕优63	W1T1	197.0±5.3 a	80.7±3.3 a	116.3±7.3 a	0.1±0.1 a	3.0±1.0 b
Shanyou 63	W2T1	193.2±0.8 a	80.6±3.6 a	112.7±3.8 a	0.0±0.0 a	3.4±0.7 b
	W1T2	160.3±3.7 b	74.0±2.0 a*	86.3±2.8 b	0.3±0.3 a	8.0±1.0 a
	W2T2	188.2±4.0 a	75.3±1.8 a*	112.9±5.1 a	0.0±0.0 a	4.7±0.9 ab
ANOVA
品种 Variety (V)		ns	ns	+	ns	ns
温度 Temperature (T)		++	ns	++	ns	+
水分 Water content (W)		++	ns	++	ns	ns
V×T		ns	ns	ns	ns	ns
V×W		ns	ns	ns	ns	ns
T×W		++	ns	++	ns	ns
V×T×W		ns	ns	ns	ns	ns

图4 干旱锻炼对幼穗分化期高温下穗部非结构性碳水化合物含量的影响数据为均值±标准误(n=3)。不同小写字母表示同一品种在不同水分和温度处理组合下同一性状在P < 0.05水平下差异显著(LSD法)。*表示相同水分与温度处理组合下两个品种间同一性状在P < 0.05水平下差异显著。V表示品种，T表示温度，W表示水分。+表示在P < 0.05水平下差异显著，ns表示在P < 0.05水平下差异不显著。W1T1表示淹灌和适温，W2T1表示干旱锻炼和适温，W1T2表示淹灌和高温，W2T2表示干旱锻炼和高温。

Fig. 4. Effects of drought priming on non-structural carbohydrate contents in panicle under high temperature during panicle initiation stage Data are shown as mean ± SE (n=3). Different lowercase letters indicate significant difference for a given trait among the four water and temperature treatment combinations for the same variety at P<0.05 (LSD test). * indicates significant difference for a given trait between two varieties in the same combination of water and temperature treatment at P<0.05. V indicates variety, T indicates temperature, and W indicates water. + indicates significant difference at P <0.05, and ns indicates no significant difference at P <0.05. W1T1 indicates flooded irrigation and normal temperature, W2T1 indicates drought priming and normal temperature, W1T2 indicates flooded irrigation and high temperature, and W2T2 indicates drought priming and high temperature.

图5 干旱锻炼对幼穗分化期高温下颖花丙二醛含量和抗氧化酶活性的影响数据为均值 ± 标准误 (n=3)。不同小写字母表示同一品种在不同水分和温度处理组合下同一性状在P < 0.05水平下差异显著(LSD法)。*表示相同水分与温度处理组合下两个品种间同一性状在P < 0.05水平下差异显著。V表示品种，T表示温度，W表示水分。+和++分别表示在P < 0.05和P < 0.01水平下差异显著，ns表示在P < 0.05水平下差异不显著。W1T1表示淹灌和适温，W2T1表示干旱锻炼和适温，W1T2表示淹灌和高温，W2T2表示干旱锻炼和高温。

Fig. 5. Effects of drought priming on MDA content and antioxidant enzyme activities in spikelet under high temperature during panicle initiation stage Data are shown as mean ± SE (n=3). Different lowercase letters indicate significant difference for a given trait among the four water and temperature treatment combinations for the same variety at P<0.05 (LSD test). * indicates significant difference for a given trait between two varieties in the same combination of water and temperature treatment at P<0.05. V indicates variety, T indicates temperature, and W indicates water. + and ++ indicate significant difference at P <0.05 and P <0.01, respectively, while ns indicates no significant difference at P <0.05. W1T1 indicates flooded irrigation and normal temperature, W2T1 indicates drought priming and normal temperature, W1T2 indicates flooded irrigation and high temperature, and W2T2 indicates drought priming and high temperature.

表5 干旱锻炼对幼穗分化期高温下颖花细胞分裂素含量的影响

Table 5. Effects of drought priming on cytokinin contents in spikelet under high temperature during panicle initiation stage

品种 Variety	处理 Treatment	活性细胞分裂素总量 aCTKs(ng/g)	反式玉米素含量 tZ(ng/g)	反式玉米素核苷含量 tZR(ng/g)	异戊烯腺嘌呤含量 iP(ng/g)	异戊烯腺苷含量 iPA(ng/g)	异戊烯腺苷磷酸含量 iPMP(ng/g)
两优培九	W1T1	19.38±0.97 a	0.70±0.01 a	1.41±0.08 a	1.90±0.09 a	1.32±0.06 a*	14.05±1.11 a
Liangyoupeijiu	W2T1	19.54±1.34 a	0.69±0.02 a	1.55±0.09 a	1.74±0.07 a*	1.23±0.06 a*	14.33±1.16 a
	W1T2	15.05±0.53 c	0.11±0.00 c	0.69±0.02 b	2.01±0.06 a*	1.35±0.08 a	10.89±0.44 b
	W2T2	17.18±0.95 b	0.15±0.00 b	1.41±0.06 a	1.89±0.08 a	1.33±0.11 a	12.40±0.82 ab
汕优63	W1T1	19.43±0.73 a	0.64±0.06 a	1.71±0.14 a*	1.70±0.21 a	1.17±0.08 a	14.23±0.27 a
Shanyou 63	W2T1	19.56±0.97 a	0.65±0.07 a	1.69±0.11 a	1.58±0.07 a	1.09±0.08 a	14.54±0.77 a
	W1T2	18.59±0.65 a*	0.64±0.02 a*	1.36±0.03 b*	1.80±0.06 a	1.29±0.05 a	13.50±0.63 a*
	W2T2	19.29±1.12 a*	0.71±0.03 a*	1.65±0.07 a*	1.80±0.01 a	1.30±0.10 a	13.84±1.03 a
ANOVA
品种 Variety(V)		+	++	ns	+	ns	ns
温度 Temperature(T)		+	++	++	ns	ns	ns
水分 Water content(W)		ns	ns	++	ns	ns	ns
V×T		ns	++	ns	ns	ns	ns
V×W		ns	ns	++	ns	ns	ns
T×W		ns	ns	++	ns	ns	ns
V×T×W		ns	ns	ns	ns	ns	ns

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