不同种植方式下温度升高对水稻产量及同化物转运的影响

doi:10.3864/j.issn.0578-1752.2021.07.017

摘要/Abstract

摘要： 【目的】气候变暖对水稻生产系统的影响备受关注,研究不同种植方式下,水稻产量及其形成对气候变化的响应规律,为水稻种植区划、栽培措施和品种调整提供依据。【方法】 2017—2018年以南粳9108和南粳46为供试品种,模拟机插秧移栽和机械化直播2种种植方式,以常温（NT）为对照,于始穗期进行中度升温（平均增加2℃,MT）和极端高温胁迫（平均增加5℃,HT）,研究不同种植方式下温度升高对不同水稻品种的产量及其构成、同化物转运、光合生产特性的影响。【结果】在中度升温和极端高温胁迫下,南粳9108和南粳46产量降幅均为移栽<直播,长生育期品种南粳46产量降幅较小。穗干物重增长速率表现为NT>MT>HT,水稻茎叶向穗的干物质转运量、转运率均随着温度升高而递减,且南粳9108下降趋势大于南粳46。穗后21 d至成熟期,剑叶SPAD值总体随着温度的升高而增加,差异达极显著水平;剑叶净光合速率穗后14—21 d均以极端高温胁迫处理下最小,而到穗后35 d以极端高温胁迫处理下最大。剑叶气孔导度、蒸腾速率均呈NT>MT>HT趋势,生育后期差异更显著。通径分析表明,产量各构成因子对产量的影响程度为结实率>千粒重>穗数>每穗粒数,温度处理对产量各构成因子的影响都表现为负效应,且以结实率影响最大（-0.819）。相关分析表明,不同种植方式下受中度升温、极端高温胁迫后,成熟期干物质总重量、茎叶干物质转运量与产量构成因子（穗数除外）,一、二次枝梗籽粒结实率都呈极显著正相关。【结论】始穗期2—5℃升温均显著降低粳稻结实率,从而导致水稻产量降低。从光合物质特性究其原因是由于温度升高降低了干物质向穗的转运率和穗干物质积累速率,从而导致生育后期水稻剑叶SPAD值增加,延长叶片持绿时间,抑制“源”向“库”转移。从气候变暖应对措施来看,选择采用移栽种植方式和长生育期品种易于表现出对极端高温胁迫逆境较好的抗性。

关键词: 高温胁迫, 种植方式, 产量, 水稻, 同化物转运

Abstract:

【Objective】The effect of global warming on rice production has attracted much attention, leading to changes on planting area, cultivated practices and rice varieties. Thus, it is necessary to clarify the responses of rice yield and its formation to warming under different planting patterns.【Method】During 2017 to 2018, we selected two varieties (Nanjing 9108 and Nanjing 46), two planting patterns (simulating machine transplanting and mechanized direct sowing), and three temperature treatments, including normal temperature (NT) as control, moderate temperature (average increase 2℃, MT) and extreme high temperature stress (average increase 5℃, HT), to study the effects of elevated temperature on rice yield, yield composition, assimilation transport and the characteristics of photosynthesis production.【Result】Under MT and HT treatments, the decreasing yields of Nanjing 9108 and Nanjing 46 under transplanting were less than that under direct sowing, while the yield of the long-growth variety Nanjing 46 had smaller decline. The growth rate of the spike dry material showed the trend of NT>MT>HT. The amount and rate of dry matter translocation from rice stem and leaves to spike decreased with the increase of temperature, and the decreasing effects on Nanjing 9108 was greater than that for Nanjing 46. Moreover, after 21 days of heading to maturity, the SPAD of flag leaf increased with elevated temperatures significantly. And the lowest net photosynthetic rate of flag leaf was found under HT at 14-21 days after heading period, while the highest was found at 35 days after heading period. In addition, The stomatal conductance and transpiration rate of flag leaf showed the increasing trend of NT>MT>HT, and the difference was more significant in the later growth stage. The path analysis indicated that the importance of yield components on yield was in the order of filled grain percentage>1000-grain weight>number of panicles>spikelet number per panicle, and all the elevated temperature treatments had negative impacts on yield components, with the greatest effect (-0.819) on filled grain percentage. The relevant analysis showed that the total weight of dry matter at maturity, the amount of translocation were significantly positively correlated to the yield composition factor (except the number of spikes) and the filled grain percentage of primary and secondary branches under MT and HT. 【Conclusion】Increasing 2℃ to 5℃ at the initial heading stage significantly reduced the filled grain percentage of rice, which led to the decrease of rice yield. From the perspective of photosynthetic characteristics, temperature rising reduced the rate of dry matter transported to spikes and declined dry matter accumulation in spikes, increasing SPAD of rice flag leaves in late growth period and thus prolonging the green holding time, which could inhibit the translocation from source to sink. Overall, we suggest that long-growth period varieties with proper plant patterns could have better resistance to elevated temperatures, which could be adaptive to the global warming.

Key words: high temperature stress, planting modes, yield, rice, assimilate translocation

张明静,韩笑,胡雪,臧倩,许轲,蒋敏,庄恒扬,黄丽芬. 不同种植方式下温度升高对水稻产量及同化物转运的影响[J]. 中国农业科学, 2021, 54(7): 1537-1552.

ZHANG MingJing,HAN Xiao,HU Xue,ZANG Qian,XU Ke,JIANG Min,ZHUANG HengYang,HUANG LiFen. Effects of Elevated Temperature on Rice Yield and Assimilate Translocation Under Different Planting Patterns[J]. Scientia Agricultura Sinica, 2021, 54(7): 1537-1552.

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https://www.chinaagrisci.com/CN/Y2021/V54/I7/1537

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温度升高对水稻产量及其构成因素的影响"

品种 Variety	种植方式 Planting modes	温度处理 Temperature treatment	穗数 No. of panicles per hole	每穗粒数 Spikelet number per panicle	千粒重 1000-grain weight (g)	结实率 Filled grain percentage (%)	实际产量 Actual yield per hole (g)	收获指数 Harvest index
南粳9108 NJ9108	移栽 TP	常温 NT	10.89±0.29Aa	118.24±0.61Aa	25.85±0.16Aa	88.26±0.07Aa	24.50±0.03Aa	0.52±0.01Aa
		中度升温 MT	10.89±0.11Aa	116.10±0.64Aa	24.66±0.13Ab	47.51±0.32Bb	19.38±0.12Bb	0.45±0.02Ab
		极端高温 HT	10.56±0.29Aa	109.41±0.17Bb	25.01±0.26Aab	7.32±0.17Cc	11.01±0.03Cc	0.27±0.01Bc
	直播 DS	常温 NT	12.78±0.11Aa	122.99±0.23Aa	25.63±0.06Aa	87.32±0.71Aa	28.65±0.09Aa	0.56±0.01Aa
		中度升温 MT	12.33±0.19Aa	115.81±1.12ABb	24.95±0.08Ab	52.45±0.68Bb	22.32±0.14Bb	0.47±0.01Ab
		极端高温 HT	12.11±0.29Aa	111.47±0.67Bb	25.25±0.15Aab	5.04±0.11Cc	9.33±0.09Cc	0.22±0.02Bc
南粳46 NJ46	移栽 TP	常温 NT	11.56±0.11Aa	123.54±1.21Aa	26.89±0.02Aa	87.32±0.45Aa	32.11±0.18Aa	0.53±0.00Aa
		中度升温 MT	11.67±0.33Aa	120.41±0.45Aa	26.36±0.54Aa	76.64±0.63Bb	27.26±0.03Bb	0.48±0.00Bb
		极端高温 HT	11.22±0.22Aa	117.80±1.40Aa	25.59±0.20Aa	56.92±0.54Cc	20.79±0.11Cc	0.38±0.00Cc
	直播 DS	常温 NT	13.33±0.19Aa	121.55±0.36Aa	26.27±0.08Aa	88.39±0.62Aa	33.89±0.16Aa	0.51±0.01Aa
		中度升温 MT	11.33±0.38Bb	126.99±1.72Aa	25.82±0.06ABb	65.47±0.19Bb	25.28±0.11Bb	0.48±0.01ABa
		极端高温 HT	11.22±0.29Bb	125.50±1.16Aa	25.36±0.02Bc	55.02±1.35Cc	19.58±0.16Cc	0.41±0.01Bb
南粳9108 NJ9108	种植方式 Planting modes		74.54**	16.69*	0.66	61.91**	64.20**	0.17
	温度处理 Temperature treatments		2.33	122.07**	19.73**	19670.00**	1831.00**	338.68**
	种植方式×温度处理 Planting modes×Temperature treatments		0.51	7.49*	1.74	28.23**	62.60**	8.08*
南粳46 NJ46	种植方式 Planting modes		4.68	18.81**	5.64	42.78**	2.04	0.29
	温度处理 Temperature treatments		11.07**	1.58	10.86*	940.85**	514.37**	232.72**
	种植方式×温度处理 Planting modes×Temperature treatments		8.68**	10.49*	0.38	36.24**	12.35**	8.21*

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温度处理Temperature treatment	日期 Date (M-D)	2:00-5:00	5:00-8:00	8:00-11:00	11:00-14:00	14:00-17:00	17:00-20:00	20:00-23:00	23:00-2:00	平均温度Average temperature
常温 NT	08-23—08-31	27.0	27.0	28.0	31.0	33.0	31.0	28.0	27.0	29.0
	09-01—09-05	26.0	26.0	27.0	30.0	32.0	30.0	27.0	26.0	28.0
	09-06—09-10	24.0	24.0	25.0	28.0	30.0	28.0	25.0	24.0	27.0
	09-11—09-16	23.0	23.0	24.0	28.0	30.0	27.0	25.0	24.0	25.5
	09-17—09-20	23.0	23.0	24.0	28.0	29.0	26.0	25.0	24.0	25.3
	09-21—09-25	23.0	23.0	24.0	26.0	29.0	25.0	24.0	23.0	24.6
中度升温 MT	08-23—08-31	29.0	29.0	30.0	33.0	35.0	33.0	30.0	29.0	31.0
	09-01—09-05	28.0	28.0	29.0	32.0	34.0	32.0	29.0	28.0	30.0
	09-06—09-10	26.0	26.0	27.0	30.0	32.0	30.0	27.0	26.0	29.0
	09-11—09-16	25.0	25.0	26.0	30.0	32.0	29.0	27.0	26.0	27.5
	09-17—09-20	25.0	25.0	26.0	30.0	31.0	28.0	27.0	26.0	27.3
	09-21—09-25	25.0	25.0	26.0	28.0	31.0	27.0	26.0	25.0	26.6
极端高温 HT	08-23—08-31	32.0	32.0	33.0	36.0	38.0	36.0	33.0	32.0	34.0
	09-01—09-05	31.0	31.0	32.0	35.0	37.0	35.0	32.0	31.0	33.0
	09-06—09-10	29.0	29.0	30.0	33.0	35.0	33.0	30.0	29.0	32.0
	09-11—09-16	28.0	28.0	29.0	33.0	35.0	32.0	30.0	29.0	30.5
	09-17—09-20	28.0	28.0	29.0	32.0	34.0	32.0	30.0	29.0	30.3
	09-21—09-25	28.0	28.0	29.0	31.0	34.0	30.0	29.0	28.0	29.6

变异来源 Source of variation	穗数 No. of panicles per hole	每穗粒数 Spikelet number per panicle	千粒重 1000-grain weight (g)	结实率 Filled grain percentage (%)	实际产量 Actual yield per hole (g)	收获指数 Harvest index	干物质转运量 Dry matter translocation amount (g·hole^-1)
年度Y	1.16ns	0.32ns	0.61ns	1.32ns	3.63ns	0.13ns	2.45ns
品种V	1.11ns	186.46**	93.22**	9296.25**	7157.96**	233.75**	474.33**
种植方式P	2.74ns	70.91**	10.30**	24.80**	63.37**	5.23**	13.18**
温度处理T	0.91ns	31.98**	31.71**	17056.86**	9731.15**	1336.83**	4303.98**
Y×V	0.97ns	0.32ns	0.02ns	0.83ns	0.01ns	0.71ns	0.01ns
Y×P	0.91ns	0.30ns	1.38ns	5.586ns	0.05ns	1.75ns	0.01ns
Y×T	1.00ns	2.67ns	2.21ns	2.94ns	2.28ns	3.26ns	0.35ns
V×P	0.33ns	7.20**	25.41**	67.39**	136.46**	0.71ns	56.03**
V×T	0.91ns	49.97**	1.36ns	3346.24**	270.61**	221.41**	84.07**
P×T	0.66ns	11.47**	0.68**	16.03**	200.42**	6.97**	26.29**
V×P×T	0.77ns	44.74**	2.61ns	134.66**	49.01**	31.15**	169.53**
Y×V×P	0.87ns	2.67ns	3.96ns	0.47*	0.59ns	1.93ns	1.07ns
Y×V×T	1.04ns	0.08ns	0.41ns	0.20ns	0.49ns	0.71ns	1.36ns
Y×P×T	1.06ns	0.30ns	3.30ns	0.28ns	2.26ns	0.71ns	4.76ns
Y×V×P×T	1.00ns	0.08ns	0.91ns	2.73ns	4.12ns	0.19ns	3.12ns

品种 Variety	种植方式 Planting modes	温度处理 Temperature treatments	一次枝梗千粒重 1000-grain weight of primary branch (g)	二次枝梗千粒重 1000-grain weight of secondary branch (g)	一次枝梗结实率 Filled grain percentage of primary branch (%)	二次枝梗结实率 Filled grain percentage of secondary branch (%)
南粳9108 NJ9108	移栽 TP	常温 NT	27.71±0.33Aa	25.52±0.09Aa	90.82±0.28Aa	84.88±0.09Aa
		中度升温MT	25.35±0.14ABb	24.03±0.216Bb	55.27±0.72Bb	36.30±0.43Bb
		极端高温HT	25.24±0.16Bb	23.96±0.05Bb	7.95±0.40Cc	7.13±0.12Cc
	直播 DS	常温 NT	26.47±0.50Aa	25.23±0.03Aa	89.48±0.44Aa	85.11±0.81Aa
		中度升温MT	26.53±0.22Aa	24.07±0.00Bb	60.64±1.12Bb	43.93±2.05Bb
		极端高温HT	25.57±0.17Aa	23.78±0.16Bb	5.52±0.07Cc	4.35±0.09Cc
南粳46 NJ46	移栽 TP	常温 NT	27.95±0.64Aa	26.62±0.39Aa	90.00±0.83Aa	83.17±0.18Aa
		中度升温MT	26.79±0.11Aa	26.34±0.56Aa	81.16±2.13Aa	72.60±0.52Bb
		极端高温HT	26.26±0.11Aa	25.23±0.23Aa	60.11±1.32Bb	53.47±0.52Cc
	直播 DS	常温 NT	28.68±0.02Aa	26.70±0.23Aa	90.44±0.43Aa	85.65±0.92Aa
		中度升温MT	27.50±0.01Bb	25.25±0.02Aab	73.69±1.27Bb	54.80±0.09Bb
		极端高温HT	27.06±0.17Bb	24.78±0.40Ab	63.97±0.84Cc	44.29±1.99Bc
南粳9108 NJ9108	种植方式 Planting modes		0.16	2.26	1.22	5.1
	温度处理 Temperature treatments		18.52**	97.08**	10160.00**	3748.00**
	种植方式×温度处理 Planting modes×Temperature treatments		9.39*	0.99	25.60**	17.02**
南粳46 NJ46	种植方式 Planting modes		11.05*	2.99	1.06	111.92**
	温度处理 Temperature treatments		18.99**	11.53**	252.87**	712.90**
	种植方式×温度处理 Planting modes×Temperature treatments		0.02	1.43	10.71**	57.97**

品种 Variety	种植方式 Planting modes	温度处理 Temperature treatments	始穗期 Initial heading stage (g·hole^-1)		成熟期 Mature stage (g·hole^-1)		转运量 Translocation amount (g·hole^-1)	转运率 Translocation rate (%)	贡献率 Contribution rate (%)
品种 Variety	种植方式 Planting modes	温度处理 Temperature treatments	茎叶 Stem and leaf	穗 Panicle	茎叶 Stem and leaf	穗 Panicle	转运量 Translocation amount (g·hole^-1)	转运率 Translocation rate (%)	贡献率 Contribution rate (%)
南粳9108 NJ9108	移栽 TP	常温 NT	40.75±0.93	2.48±0.06	30.73±0.80Bc	29.95±1.12Aa	10.02±0.46Aa	24.60	33.46
		中度升温MT			32.71±0.85Bb	21.32±0.55Bb	8.04±0.21Bb	19.73	37.71
		极端高温 HT			35.71±0.93Aa	12.11±1.31Cc	5.04±0.15Cc	12.37	41.63
	直播 DS	常温 NT	39.89±1.04	3.15±0.08	28.86±0.75Bc	31.52±0.82Aa	11.03±0.29Aa	27.65	35.00
		中度升温 MT			30.09±0.78Bb	25.55±0.56Bb	9.80±0.27Bb	24.56	38.34
		极端高温 HT			36.19±0.94Aa	9.39±0.34Cc	3.70±0.18Cc	9.28	39.40
南粳46 NJ46	移栽 TP	常温 NT	40.15±0.97	2.87±0.07	27.74±0.72Cc	35.32±0.92Aa	12.41±0.32Aa	30.91	35.13
		中度升温 MT			30.95±0.80Bb	29.99±1.08Bb	9.20±0.24Bb	22.92	30.69
		极端高温 HT			34.25±0.89Aa	22.87±0.59Cc	5.90±0.17Cc	14.69	25.80
	直播 DS	常温 NT	40.7±1.12	3.05±0.05	28.89±0.75Cc	35.28±1.32Aa	11.81±0.31Aa	29.02	33.48
		中度升温 MT			31.75±0.83Bb	29.81±0.78Bb	8.95±0.13Bb	21.99	30.02
		极端高温 HT			34.25±0.72Aa	21.58±0.57Cc	6.46±0.09Cc	15.86	29.91

相关性 Correlation	穗数 No. of panicles	每穗粒数 Spikelet number per panicle	千粒重 1000-grain weight	结实率 Filled grain percentage	一次枝梗籽粒结实率 Filled grain percentage of primary branch	二次枝梗籽粒结实率 Filled grain percentage of secondary branch	产量 Yield
干物质重量 Dry matter weight	0.309	0.700*	0.853**	0.932**	0.901**	0.956**	0.908**
转运量 Translocation amount	0.459	0.587*	0.619*	0.887**	0.879**	0.880**	0.936**
SPAD值 SPAD value	-0.164	0.352	0.146	-0.217	-0.183	-0.257	-0.153
剑叶净光合速率 Net photosynthetic rate of flag leaf	-0.194	-0.743**	-0.499	-0.689*	-0.729**	-0.628*	-0.676*
气孔导度 Stomatal conductance	0.452	0.502	0.427	0.742**	0.731**	0.740**	0.801**