结实期低温胁迫对水稻强、弱势粒淀粉形成与积累的影响

doi:10.16819/j.1001-7216.2022.211105

摘要/Abstract

摘要：

【目的】研究结实期低温胁迫对水稻强、弱势粒淀粉组分含量、积累速率及关键酶活性的影响，明确淀粉合成关键酶活性变化对淀粉积累速率的调控效应，探究强、弱势粒淀粉形成积累差异对水稻产量的影响。【方法】以耐冷型品种东农428和冷敏型品种松粳10为试验材料，设置1个常温处理(白天温度28℃，14h/夜间温度22℃，10 h，7 d)和4个低温处理(17℃，低温处理时间分别为1、3、5、7 d)，分析了结实期低温胁迫对强、弱势粒淀粉组分积累、合成关键酶活性、水稻产量及构成因素的影响，并探讨了灌浆期各阶段淀粉积累差异与酶活性变化的关系。【结果】与对照相比，结实期低温胁迫降低了两个品种水稻强、弱势粒中腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)、可溶性淀粉合成酶(SSS)、淀粉分支酶(SBE)的峰值活性以及支链淀粉和总淀粉含量，提高了齐穗后28~38 d 低温处理3、5、7 d的颗粒结合型淀粉合成酶(GBSS)活性和直链淀粉含量。与对照相比，各低温处理酶活性最高和淀粉积累最快的时间均有不同程度的推迟，低温处理7 d的影响最大，强、弱势粒低温处理7 d的支链淀粉和总淀粉含量分别在齐穗后13、18 d降幅最大，直链淀粉含量在28 d增幅最大。相关分析表明，强、弱势粒直链淀粉含量及强势粒支链淀粉、总淀粉含量与其最大积累速率呈极显著正相关，弱势粒支链淀粉、总淀粉含量还受到其最大积累速率出现时间的影响。AGPase、GBSS、SSS、SBE活性变化与淀粉积累速率和积累时间早晚密切相关，对淀粉及淀粉组分含量的变化有着明显的影响。同时，结实期低温胁迫显著降低了水稻的千粒重、结实率和产量，且随低温处理天数的增加降幅逐渐增大。结实期低温胁迫对弱势粒中淀粉合成关键酶活性变化影响大于强势粒，弱势粒淀粉合成积累减慢，含量降低，导致水稻千粒重显著下降，产量降低。【结论】就品种而言，耐冷型东农428具有较高的淀粉合成关键酶活性，淀粉及其组分含量较高，在低温胁迫下产量能维持在较高的水平。因此，强、弱势粒淀粉合成关键酶活性对淀粉合成起着非常关键的调控作用，淀粉组分和含量及其变化对产量有着十分重要的影响。

关键词: 水稻, 结实期, 低温, 淀粉合成

Abstract:

【Objective】Our purposes are to illuminate the effects of low temperature stress during grain filling on the contents, accumulation rates and key enzyme activities of starch components in superior and inferior grains in rice, to clarify the regulatory effect of changes in the activities of key enzymes in starch synthesis on the rate of starch accumulation and to explore the effect of difference in granule starch formation and accumulation in superior and inferior grains on rice yield formation. 【Method】This experiment was carried out with the chill-tolerant variety Dongnong 428 and chill-sensitive variety Songjing 10 as the test materials at room temperature (daytime temperature 28℃, 14 h/night temperature 22℃, 10 h; 7 days), and low temperature (17℃, four treatments). The low temperature treatments lasted 1, 3, 5, and 7 days(abbreviated to D₁, D₃, D₅ and D₇), respectively. The difference in the accumulation of starch components of superior and inferior grains and the changes in the activities of key synthesizing enzymes under low temperature stress during grain-filling period were analyzed. 【Result】Compared with the control, low temperature stress during grain filling period reduced the peak activities of adenosine diphosphoglucose pyrophosphorylase (AGPase), soluble starch synthase (SSS), starch branching enzyme (SBE) and the contents of amylopectin and total starch in the superior and inferior grains of the two rice varieties, improved granule-bound starch synthase (GBSS) activity and amylose content in D₃, D₅, and D₇ treatments 28－38 days after heading. Compared with the control, the highest enzyme activity and the fastest accumulation of starch in low temperature treatments occurred at a later time, and the D₇ treatment had the greatest impact. The contents of amylopectin and total starch of the superior and inferior grains in D₇ decreased most on the 13th and 18th days after heading, and the amylose content increased most on the 28th day after heading. Correlation analysis showed that the amylose content of superior and inferior grains and the content of superior amylopectin and total starch are extremely significantly positively correlated with their maximum accumulation rates. The accumulation of inferior grain amylopectin and total starch is also related to the time to reach the maximum accumulation rate. The changes in the activities of AGPase, GBSS, SSS, and SBE are closely related to the accumulation rate and time of starch accumulation, and have a significant impact on the changes in starch and starch component contents. At the same time, low temperature stress during the grain filling stage significantly reduced the thousand-grain weight, seed setting rate and yield of rice, and the decrease gradually increased with the increasing low temperature treatment days. Low temperature stress during the grain filling stage has a greater impact on the changes in the key enzyme activities of starch synthesis in the inferior grains than the superior ones. The starch synthesis and accumulation of inferior grains slowed down and the content decreased, resulting in a significant decrease in thousand-grain weight and yield. 【Conclusion】In terms of varieties, the cold-tolerant Dongnong 428 has relatively high key enzyme activities of starch synthesis, and the contents of starch and its components is relatively high, so that the rice yield can be maintained at a relatively high level under low temperature stress. Therefore, the key enzyme activities of starch synthesis of granules in superior and inferior grains play a very critical role in the regulation of starch synthesis, and the changes and levels of starch components have a very important impact on yield.

Key words: rice, grain filling stage, low temperature, starch synthesis

陈红阳, 贾琰, 赵宏伟, 瞿炤珺, 王新鹏, 段雨阳, 杨蕊, 白旭, 王常丞. 结实期低温胁迫对水稻强、弱势粒淀粉形成与积累的影响[J]. 中国水稻科学, 2022, 36(5): 487-504.

CHEN Hongyang, JIA Yan, ZHAO Hongwei, QU Zhaojun, WANG Xinpeng, DUAN Yuyang, YANG Rui, BAI Xu, WANG Changcheng. Effects of Low Temperature Stress During Grain Filling on Starch Formation and Accumulation of Superior and Inferior Grains in Rice[J]. Chinese Journal OF Rice Science, 2022, 36(5): 487-504.

图/表 13

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粒位 Grain position	品种 Variety	处理 Treatment	齐穗后天数Days after full heading stage
粒位 Grain position	品种 Variety	处理 Treatment	8~13	13~18	18~23		23~28	28~33	33~38
强势粒 Superior grains	东农428 Dongnong 428	D₀	0.81 a	0.61 a	0.48 e		0.24 e	0.17 a	0.07 b
		D₁	0.77 b	0.60 b	0.52 d		0.25 d	0.19 a	0.06 b
		D₃	0.51 c	0.47 d	0.67 c		0.83 c	0.03 c	0.10 a
		D₅	0.38 d	0.58 c	0.76 b		0.90 b	0.04 b	0.09 a
		D₇	0.21 e	0.57 c	0.93 a		1.05 a	0.06 b	0.10 a
	松粳10 Songjing 10	D₀	0.81 a	0.54 c	0.50 e		0.32 e	0.12 a	0.07 b
		D₁	0.76 b	0.53 c	0.52 d		0.40 d	0.11 a	0.09 a
		D₃	0.51 c	0.63 a	0.64 c		0.88 c	0.06 b	0.06 b
		D₅	0.20 e	0.56 b	0.74 b		1.28 b	0.07 b	0.10 a
		D₇	0.23 d	0.47 d	0.89 a		1.39 a	0.06 b	0.08 a
弱势粒 Inferior grains	东农428 Dongnong 428	D₀	0.40 a	0.79 a	0.60 d		0.44 e	0.19 b	0.20 a
		D₁	0.37 b	0.78 a	0.59 d		0.48 d	0.27 a	0.18 b
		D₃	0.34 c	0.54 b	0.78 b		0.83 c	0.11 c	0.21 a
		D₅	0.22 d	0.55 b	0.73 c		0.99 b	0.19 b	0.15 c
		D₇	0.15 e	0.44 c	0.85 a		1.10 a	0.19 b	0.19 a
	松粳10 Songjing 10	D₀	0.44 a	0.85 a	0.56 d		0.53 e	0.21 b	0.10 b
		D₁	0.38 b	0.85 a	0.56 d		0.57 d	0.23 a	0.07 c
		D₃	0.29 c	0.51 b	0.79 c		0.94 c	0.20 b	0.09 b
		D₅	0.18 d	0.41 d	0.84 a		1.25 b	0.18 c	0.12 a
		D₇	0.13 e	0.43 c	0.82 b		1.42 a	0.20 b	0.11 a
F值 F value	强势粒 Superior grains	处理Treatment(T)	166.11**	50.95**		142.15**	276.34**	109.63**	1.25
		品种Variety(V)	92.37**	51.36**		168.62**	223.58**	199.02**	1.06
		处理×品种T×V	15.38**	9.37**		11.72**	20.02**	17.05**	0.72
	弱势粒 Inferior grains	处理(T)	369.01**	167.58**		101.97**	525.66**	51.84**	66.18**
		品种(V)	121.30**	32.17*		96.38**	310.01**	34.79*	325.93**
		处理×品种T×V	31.10**	22.26**		24.67**	52.01**	9.13*	13.18**

粒位 Grain position	品种 Variety	处理 Treatment	齐穗后天数Days after full heading stage
粒位 Grain position	品种 Variety	处理 Treatment	8~13	13~18	18~23		23~28	28~33	33~38
强势粒 Superior grains	东农428 Dongnong 428	D₀	0.81 a	0.61 a	0.48 e		0.24 e	0.17 a	0.07 b
		D₁	0.77 b	0.60 b	0.52 d		0.25 d	0.19 a	0.06 b
		D₃	0.51 c	0.47 d	0.67 c		0.83 c	0.03 c	0.10 a
		D₅	0.38 d	0.58 c	0.76 b		0.90 b	0.04 b	0.09 a
		D₇	0.21 e	0.57 c	0.93 a		1.05 a	0.06 b	0.10 a
	松粳10 Songjing 10	D₀	0.81 a	0.54 c	0.50 e		0.32 e	0.12 a	0.07 b
		D₁	0.76 b	0.53 c	0.52 d		0.40 d	0.11 a	0.09 a
		D₃	0.51 c	0.63 a	0.64 c		0.88 c	0.06 b	0.06 b
		D₅	0.20 e	0.56 b	0.74 b		1.28 b	0.07 b	0.10 a
		D₇	0.23 d	0.47 d	0.89 a		1.39 a	0.06 b	0.08 a
弱势粒 Inferior grains	东农428 Dongnong 428	D₀	0.40 a	0.79 a	0.60 d		0.44 e	0.19 b	0.20 a
		D₁	0.37 b	0.78 a	0.59 d		0.48 d	0.27 a	0.18 b
		D₃	0.34 c	0.54 b	0.78 b		0.83 c	0.11 c	0.21 a
		D₅	0.22 d	0.55 b	0.73 c		0.99 b	0.19 b	0.15 c
		D₇	0.15 e	0.44 c	0.85 a		1.10 a	0.19 b	0.19 a
	松粳10 Songjing 10	D₀	0.44 a	0.85 a	0.56 d		0.53 e	0.21 b	0.10 b
		D₁	0.38 b	0.85 a	0.56 d		0.57 d	0.23 a	0.07 c
		D₃	0.29 c	0.51 b	0.79 c		0.94 c	0.20 b	0.09 b
		D₅	0.18 d	0.41 d	0.84 a		1.25 b	0.18 c	0.12 a
		D₇	0.13 e	0.43 c	0.82 b		1.42 a	0.20 b	0.11 a
F值 F value	强势粒 Superior grains	处理Treatment(T)	166.11**	50.95**		142.15**	276.34**	109.63**	1.25
		品种Variety(V)	92.37**	51.36**		168.62**	223.58**	199.02**	1.06
		处理×品种T×V	15.38**	9.37**		11.72**	20.02**	17.05**	0.72
	弱势粒 Inferior grains	处理(T)	369.01**	167.58**		101.97**	525.66**	51.84**	66.18**
		品种(V)	121.30**	32.17*		96.38**	310.01**	34.79*	325.93**
		处理×品种T×V	31.10**	22.26**		24.67**	52.01**	9.13*	13.18**

粒位 Grain position	品种 Variety	处理 Treatment	齐穗后天数Days after full heading stage
粒位 Grain position	品种 Variety	处理 Treatment	8~13	13~18	18~23		23~28	28~33	33~38
强势粒SG	东农428 Dongnong 428	D₀	3.58 a	2.21 d	1.18 c		1.29 b	0.29 e	0.32 a
		D₁	3.47 b	2.31 c	1.16 d		1.36 a	0.33 d	0.28 b
		D₃	3.16 c	2.65 a	1.10 e		1.09 c	0.46 c	0.16 d
		D₅	3.14 c	2.37 b	1.46 a		0.70 e	0.65 a	0.25 c
		D₇	2.99 d	2.29 e	1.44 b		0.99 d	0.57 b	0.14 d
	松粳10 Songjing 10	D₀	3.69 a	2.14 b	1.33 d		0.95 d	0.38 e	0.49 a
		D₁	3.58 b	1.88 e	1.61 c		1.15 c	0.42 d	0.39 b
		D₃	3.15 c	2.10 c	1.70 b		1.18 c	0.46 c	0.34 c
		D₅	2.38 d	2.00 d	1.88 a		1.72 a	0.57 b	0.22 d
		D₇	1.60 e	2.19 a	1.89 a		1.51 b	0.89 a	0.32 c
弱势粒IG	东农428 Dongnong 428	D₀	1.46 a	2.36 a	1.75 d		1.76 c	1.07 b	0.41 e
		D₁	1.06 b	2.28 b	1.79 c		1.72 d	1.04 c	0.69 c
		D₃	0.85 c	2.26 b	1.80 c		1.71 d	1.10 a	0.43 d
		D₅	0.40 d	1.95 c	2.29 b		1.79 b	0.83 d	0.71 b
		D₇	0.40 d	1.64 d	2.51 a		1.84 a	0.84 d	0.95 a
	松粳10 Songjing 10	D₀	1.39 a	2.59 a	1.72 d		1.52 d	1.11 c	0.48 c
		D₁	1.14 b	2.24 b	1.94 c		1.52 d	1.08 c	0.71 a
		D₃	0.89 c	1.81 c	1.94 c		1.87 a	1.28 b	0.33 d
		D₅	0.74 d	1.48 d	2.14 b		1.74 b	1.39 a	0.46 c
		D₇	0.42 e	1.40 e	2.18 a		1.72 c	1.37 a	0.51 b
F值 F value	强势粒 Superior grains	处理Treatment(T)	205.62**	359.23**		296.56**	126.65**	119.98**	86.34*
		品种Variety(V)	392.38**	452.06**		296.76**	256.46**	206.26**	195.14**
		处理×品种T×V	35.18**	49.29**		43.06**	30.12**	19.35**	10.72*
	弱势粒 Inferior grains	处理(T)	134.45**	296.52**		104.46**	117.96**	96.88**	76.14**
		品种(V)	164.42**	203.80*		88.46**	102.44**	169.75**	103.42**
		处理×品种T×V	26.43**	33.45**		16.31**	12.11*	19.23**	10.78*

粒位 Grain position	品种 Variety	处理 Treatment	齐穗后天数Days after full heading stage
粒位 Grain position	品种 Variety	处理 Treatment	8~13	13~18	18~23		23~28	28~33	33~38
强势粒SG	东农428 Dongnong 428	D₀	3.58 a	2.21 d	1.18 c		1.29 b	0.29 e	0.32 a
		D₁	3.47 b	2.31 c	1.16 d		1.36 a	0.33 d	0.28 b
		D₃	3.16 c	2.65 a	1.10 e		1.09 c	0.46 c	0.16 d
		D₅	3.14 c	2.37 b	1.46 a		0.70 e	0.65 a	0.25 c
		D₇	2.99 d	2.29 e	1.44 b		0.99 d	0.57 b	0.14 d
	松粳10 Songjing 10	D₀	3.69 a	2.14 b	1.33 d		0.95 d	0.38 e	0.49 a
		D₁	3.58 b	1.88 e	1.61 c		1.15 c	0.42 d	0.39 b
		D₃	3.15 c	2.10 c	1.70 b		1.18 c	0.46 c	0.34 c
		D₅	2.38 d	2.00 d	1.88 a		1.72 a	0.57 b	0.22 d
		D₇	1.60 e	2.19 a	1.89 a		1.51 b	0.89 a	0.32 c
弱势粒IG	东农428 Dongnong 428	D₀	1.46 a	2.36 a	1.75 d		1.76 c	1.07 b	0.41 e
		D₁	1.06 b	2.28 b	1.79 c		1.72 d	1.04 c	0.69 c
		D₃	0.85 c	2.26 b	1.80 c		1.71 d	1.10 a	0.43 d
		D₅	0.40 d	1.95 c	2.29 b		1.79 b	0.83 d	0.71 b
		D₇	0.40 d	1.64 d	2.51 a		1.84 a	0.84 d	0.95 a
	松粳10 Songjing 10	D₀	1.39 a	2.59 a	1.72 d		1.52 d	1.11 c	0.48 c
		D₁	1.14 b	2.24 b	1.94 c		1.52 d	1.08 c	0.71 a
		D₃	0.89 c	1.81 c	1.94 c		1.87 a	1.28 b	0.33 d
		D₅	0.74 d	1.48 d	2.14 b		1.74 b	1.39 a	0.46 c
		D₇	0.42 e	1.40 e	2.18 a		1.72 c	1.37 a	0.51 b
F值 F value	强势粒 Superior grains	处理Treatment(T)	205.62**	359.23**		296.56**	126.65**	119.98**	86.34*
		品种Variety(V)	392.38**	452.06**		296.76**	256.46**	206.26**	195.14**
		处理×品种T×V	35.18**	49.29**		43.06**	30.12**	19.35**	10.72*
	弱势粒 Inferior grains	处理(T)	134.45**	296.52**		104.46**	117.96**	96.88**	76.14**
		品种(V)	164.42**	203.80*		88.46**	102.44**	169.75**	103.42**
		处理×品种T×V	26.43**	33.45**		16.31**	12.11*	19.23**	10.78*

粒位 Grain position	品种 Variety	处理 Treatment	齐穗后天数Days after full heading stage
粒位 Grain position	品种 Variety	处理 Treatment	8~13	13~18	18~23		23~28	28~33	33~38
强势粒SG	东农428 Dongnong 428	D₀	4.39 a	2.83 d	1.66 d		1.53 d	0.46 d	0.39 a
		D₁	4.25 b	2.91 c	1.67 d		1.61 c	0.51 c	0.35 b
		D₃	3.70 c	3.12 a	1.76 c		1.92 b	0.49 c	0.25 d
		D₅	3.51 d	2.95 b	2.22 b		1.60 c	0.69 a	0.33 c
		D₇	3.20 e	2.86 e	2.37 a		2.05 a	0.62 b	0.24 d
	松梗10 Songjing 10	D₀	4.50 a	2.68 b	1.84 e		1.27 e	0.50 d	0.56 a
		D₁	4.35 b	2.42 d	2.14 d		1.56 d	0.53 c	0.52 b
		D₃	3.66 c	2.73 a	2.34 c		1.98 c	0.52 c	0.50 b
		D₅	2.59 d	2.56 c	2.63 b		2.99 a	0.64 b	0.32 c
		D₇	1.83 e	2.66 b	2.79 a		2.90 b	0.95 a	0.28 d
弱势粒IG	东农428 Dongnong 428	D₀	1.86 a	3.19 a	2.36 e		2.20 d	1.26 b	0.60 c
		D₁	1.42 b	3.05 b	2.39 d		2.20 d	1.31 a	0.87 b
		D₃	1.19 c	2.79 c	2.58 c		2.55 c	1.21 c	0.64 c
		D₅	0.63 d	2.50 d	3.03 b		2.78 b	1.02 d	0.86 b
		D₇	0.55 e	2.26 e	3.19 a		2.94 a	1.03 d	1.13 a
	松梗10 Songjing 10	D₀	1.83 a	3.44 a	2.29 d		2.05 d	1.33 d	0.58 e
		D₁	1.52 b	3.08 b	2.50 c		2.09 d	1.40 c	0.79 b
		D₃	1.17 c	2.31 c	2.73 b		2.81 c	1.48 b	0.41 d
		D₅	0.92 d	1.89 d	2.99 a		2.99 b	1.57 a	0.84 a
		D₇	0.55 e	1.82 d	3.00 a		3.14 a	1.56 a	0.63 c
F值 F value	强势粒 Superior grains	处理Treatment(T)	298.35**	202.65**		168.64**	245.40**	96.14**	79.64*
		品种Variety(V)	205.46**	462.62**		246.13**	301.14**	196.45**	106.54**
		处理×品种T×V	36.65**	29.35**		20.02**	44.54**	11.69**	9.47**
	弱势粒 Inferior grains	处理(T)	163.49**	341.48**		116.45**	99.84**	67.69**	56.74**
		品种(V)	196.35**	160.36*		156.23**	134.06**	106.99**	70.25**
		处理×品种T×V	22.64**	22.13**		19.47**	12.22**	9.15**	6.88**