孕穗期杂交中稻对淹涝胁迫的响应

doi:10.16819/j.1001-7216.2023.230304

摘要/Abstract

摘要：

【目的】 为分析水稻孕穗期对淹涝胁迫的响应，减少淹涝胁迫导致的水稻产量损失。【方法】 于2017和2018年在中稻丰两优香1号孕穗期设计了不同淹涝时间(3 d、6 d和9 d)和淹涝深度[淹1/4(水深达试验开始时水稻株高的25%)、淹1/2、3/4和淹4/4]交互试验。【结果】 水稻孕穗期受淹涝胁迫后，株高显著增高，其主要是第4节间显著增长，单穗干质量显著降低。水稻剑叶叶绿素a含量、叶绿素b含量、可溶性蛋白含量、净光合速率、气孔导度、蒸腾速率随着淹涝持续时间和淹涝深度的增加不断下降；胞间CO₂浓度随着淹涝深度的增加不断下降，但在全淹涝条件下浓度出现回升。水稻剑叶中的丙二醛(MDA)含量和过氧化物酶(POD)活性随着淹涝深度的增加而增高，剑叶可溶性糖含量随着淹涝深度的增加先上升后下降。淹涝3 d时，水稻剑叶超氧化物歧化酶(SOD)活性、根系MDA含量和SOD活性随淹涝深度的增加而升高；随着淹涝持续时间的延长，水稻剑叶SOD活性、根系MDA含量和SOD活性随淹涝深度的增加先升高后降低。根系POD活性在淹涝3 d和6 d时随淹涝深度的增加而升高，在淹涝9 d时随淹涝深度的增加先升高后降低。随着淹涝深度的加深，水稻有效穗数、穗实粒数、结实率、千粒重和实际产量均降低。水稻淹涝持续时间超过6 d或淹涝深度超过植株1/2时，水稻减产均超过25%，水稻实际产量与淹涝天数和淹涝深度拟合呈二元一次关系，且淹涝深度的影响大于淹涝持续时间。【结论】 叶绿素含量、可溶性蛋白含量、净光合速率、气孔导度和蒸腾速率与水稻实际产量的相关性最高，灰色关联系数均大于0.6，可作为水稻孕穗期受淹涝后的指标，确定其受淹及减产程度。

关键词: 杂交中稻, 淹涝, 形态指标, 生理指标, 产量

Abstract:

【Objective】 This study aims to investigate the response of rice and identify key indicators to flooding stress during the booting stage, with the goal of mitigating yield losses caused by such stress. 【Method】 Experiments involving different flooding durations (3 days, 6 days, and 9 days) and depths (1/4 flooding, 25% of the rice plant height before the experiment, 1/2 flooding, 3/4 flooding, and full flooding) were conducted during the booting stage of middle-season rice (Fengliangyouxiang 1) in 2017 and 2018. 【Results】 Following flooding stress during the booting stage, there was a significant increase in rice plant height, primarily attributed to a significant increase in the 4th internode, while the dry weight of individual panicle decreased notably. Chlorophyll a (Chl a) content, chlorophyll b (Chl b) content, soluble protein content, net photosynthetic rate (P_n), stomatal conductance (G_s), and transpiration rate (T_r) of rice leaves decreased with increasing flooding duration and depth. The concentration of inter-cellular CO₂ (C_i) declined with greater flooding depth, yet exhibited a rebound under fully flooding conditions. Malondialdehyde (MDA) content and peroxidase (POD) activity in rice leaves increased with deeper flooding. The content of soluble sugars in leaves increased, then decreased, with greater flooding depth. At 3 days of flooding, superoxide dismutase (SOD) activity in rice leaves, root MDA content, and SOD activity increased with deeper flooding. With prolonged flooding, SOD activity in rice leaves, root MDA content, and SOD activity initially increased, then decreased, with increasing flooding depth. Root POD activity increased with increasing flooding depth at 3 days and 6 days, and increased, then decreased, at 9 days. The effective panicle number, filled grain number per panicle, seed setting rate, thousand grain weight, and actual rice yield all decreased with greater flooding depth. When the duration of flooding exceeded 6 days or the depth of flooding exceeded half the plant height, rice yield reduction exceeded 25%. Actual rice yield exhibited a highly significant binary correlation with both days of flooding and depth of flooding, with depth having a greater impact than duration. 【Conclusion】 Chlorophyll b (Chl b), chlorophyll a (Chl a), soluble protein, P_n, G_s, and T_r display the strongest correlation with actual rice yield, with gray correlation coefficients all exceeding 0.6. These can serve as indicators for evaluating flooding severity and predicting yield reduction during booting.

Key words: hybrid midde-season rice, flooding, morphological index, physiological index, yield

邹宇傲, 吴启侠, 周乾顺, 朱建强, 晏军. 孕穗期杂交中稻对淹涝胁迫的响应[J]. 中国水稻科学, 2023, 37(6): 642-656.

ZOU Yuao, WU Qixia, ZHOU Qianshun, ZHU Jianqiang, YAN Jun. Response of Middle-season Hybrid Rice to Flooding Stress at the Booting Stage[J]. Chinese Journal OF Rice Science, 2023, 37(6): 642-656.

图/表 16

参考文献 40

[1]	Ivanic M, Martin W. Implications of higher global food prices for poverty in low-income countries 1[J]. Agricultural economics, 2008, 39: 405-416.
[2]	张福锁, 王激清, 张卫峰, 崔振岭, 马文奇, 陈新平, 江荣风. 中国主要粮食作物肥料利用率现状与提高途径[J]. 土壤学报, 2008(5): 915-924.
	Zhang F S, Wang J Q, Zhang W F, Cui Z L, Ma W Q, Chen X P, Jiang R F. Nutrient use efficiencies of major cereal crops in China and measures for improvement[J]. Acta Pedologica Sinica, 2008(5): 915-924. (in Chinese with English abstract)
[3]	Shao G, Wang M, Yu S, Liu N, Xiao M, Yuan M. Potential of controlled irrigation and drainage for reducing nitrogen emission from rice paddies in Southern China[J]. Journal of Chemistry, 2015, 913470.
[4]	Kotera A, Nawata E. Role of plant height in the submergence tolerance of rice: A simulation analysis using an empirical model[J]. Agricultural Water Management, 2007, 89(1-2): 49-58.
[5]	李阳生, 李绍清. 淹涝胁迫对水稻生育后期的生理特性和产量性状的影响[J]. 武汉植物学研究, 2000(2): 117-122.
	Li Y S, Li S Q. Effect of submergence on physiologic indexes and yield component at reproductive stage in rice[J]. Plant Science Journal, 2000(2): 117-122. (in Chinese with English abstract)
[6]	李开江, 石鹤付, 史健, 顾长善. 分蘖期淹水对水稻生长发育和产量的影响[J]. 安徽农学通报, 2007, 90(20): 64-65.
	Li K J, Shi H F, Shi J, Gu Z S. Effects of flooding at tillering stage on growth and yield of rice[J]. Anhui Agricultural Science Bulletin, 2007, 90(20): 64-65. (in Chinese with English abstract)
[7]	姜丽霞, 闫敏慧, 翟墨, 闫平, 韩俊杰, 何锋, 王铭, 于艳梅. 关键生育期淹涝胁迫对黑龙江省水稻的影响[J]. 灾害学, 2020, 35(4): 128-134.
	Jiang L X, Yan M H, Zhai M, Yan P, Han J J, He F, Wang M, Yu Y M. Effects of waterlogging stress on japonica rice during critical growth period in Heilongjiang province[J]. Journal of Catastrophology, 2020, 35(4): 128-134. (in Chinese with English abstract)
[8]	姜丽霞, 于艳梅, 刘泽恩, 王萍, 孙丽莉, 闫平, 赵慧颖. 淹涝胁迫对寒地水稻生长和产量的影响研究[J]. 海洋气象学报, 2020, 40(2): 140-148.
	Jiang L X, Yu Y M, Liu Z E, Wang P, Sun L P, Yan P, Zhao H Y. Impact of waterlogging stress on japonica rice growth and vield in cold region[J]. Journal of Marine Meteorology, 2020, 40(2): 140-148. (in Chinese with English abstract)
[9]	吴凤燕, 王煌, 朱瑞, 翟丽妮, 胡铁松. 拔节-孕穗期淹涝胁迫对水稻生长的后效影响[J]. 农业工程学报, 2021, 37(24): 85-93.
	Wu F Y, Wang H, Zhu R, Zhai L N, Hu T S. Post-effects of waterlogging on the rice growth at the jointing-booting stage[J]. Transactions of the Chinese Society of Agricultural Engineering, 2021, 37(24): 85-93. (in Chinese with English abstract)
[10]	成添, 胡继超, 李映雪, 谢晓金, 李永秀. 淹涝胁迫对水稻植株叶片光合性能的影响[J]. 气象与环境科学, 2019, 42(1): 26-33.
	Cheng T, Hu J C, Li Y X, Xie X J, Li Y X,. Effects of flooding stress on leave's photosynthetic capability of paddy rice[J]. Meteorological and Environmental Sciences, 2019, 42(1): 26-33. (in Chinese with English abstract)
[11]	晏军, 吴启侠, 朱建强. 中稻灌浆期对淹水胁迫的响应及排水指标研究[J]. 灌溉排水学报, 2017, 36(5): 59-65.
	Yan J, Wu Q X, Zhu J Q. Response of mid-season rice to waterlogging at filing stage and its consideration in design of drainage standard[J]. Journal of Irrigation and Drainage, 2017, 36(5): 59-65. (in Chinese with English abstract)
[12]	晏军, 吴启侠, 朱建强, 徐笑笑, 张露萍. 拔节期杂交中稻对淹水胁迫的响应及指示性指标探讨[J]. 中国稻米, 2017, 23(1): 17-25.
	Yan J, Wu Q X, Zhu J Q, Xu X X, Zhang L P. Response of hybrid mid-season rice to flooding and discussion of indicative index at the jointing stage[J]. China Rice, 2017, 23(1): 17-25. (in Chinese with English abstract)
[13]	彭斯敏, 耿延琢, 程勤学, 王毅. 丰两优香1号在湖北省种植表现及其高产栽培技术[J]. 杂交水稻, 2009, 24(5): 52-53.
	Peng S M, Geng Y Z, Cheng Q X, Wang Y. Performance and high-yielding cultural techniques of fengliangyou xiang 1 in Hubei Province[J]. Hybrid Rice, 2009, 24(5): 52-53. (in Chinese with English abstract)
[14]	潘澜, 薛立. 植物淹水胁迫的生理学机制研究进展[J]. 生态学杂志, 2012, 31(10): 2662-2672.
	Pan L, Xue L. Plant physiological mechanisms in adapting to waterlogging stress: A review[J]. Chinese Journal of Ecology, 2012, 31(10): 2662-2672. (in Chinese with English abstract)
[15]	谭淑端, 朱明勇, 张克荣, 党海山, 张全发. 植物对水淹胁迫的响应与适应[J]. 生态学杂志, 2009, 28(9): 1871-1877.
	Tan S D, Zhu M Y, Zhang K R, Dang H S, Zhang Q F. Response and adaptation of plants to submergence stress[J]. Chinese Journal of Ecology, 2009, 28(9): 1871-1877. (in Chinese with English abstract)
[16]	Karalija E, Selović A. The effect of hydro and proline seed priming on growth, proline and sugar content, and antioxidant activity of maize under cadmium stress[J]. Environmental Science and Pollution Research, 2018, 25(33): 33370-33380.
[17]	李一路, 郭照辉, 肖蓉, 程伟, 张敏, 胡丹, 王玉双, 单世平, 魏小武. 外源柠檬酸杆菌对水稻镉吸收和亚细胞分布及生理特性的影响[J]. 湖南农业大学学报: 自然科学版, 2023, 49(1): 35-42.
	Li Y L, Guo Z H, Xiao R, Cheng W, Zhang M, Hu D, Wang Y S, Dan S P, Wei X W. Effects of exogenous addition of Citrobacter sp. on Cd uptake and the subcellular distribution of Cd and physiological characteristics in rice[J]. Journal of Hunan Agricultural University(Natural Sciences), 2023, 49(1): 35-42. (in Chinese with English abstract).
[18]	Zeng L S, Liao M, Chen C L, Huang C Y. Effects of lead contamination on soil enzymatic activities, microbial biomass, and rice physiological indices in soil-lead-rice (Oryza sativa L.) system[J]. Ecotoxicology and Environmental Safety, 2007, 67(1): 67-74.
[19]	Sone C, Ito O, Sakagami J I. Characterizing submergence survival strategy in rice via chlorophyll fluorescence[J]. Journal of Agronomy and Crop Science, 2012, 198(2): 152-160.
[20]	向镜, 陈惠哲, 张玉屏, 张义凯, 朱德峰. 淹涝条件下水温对水稻幼苗形态和生理的影响[J]. 中国水稻科学, 2016, 30(5): 525-531.
	Xiang J, Chen H Z, Zhang Y P, Zhang Y K, Zhu D F. Morphological and physiological responses of rice seedlings to water temperature under complete submergence[J]. Chinese Journal of Rice Science, 2016, 30(5): 525-531. (in Chinese with English abstract)
[21]	甄博, 周新国, 陆红飞, 李会贞. 高温与涝交互胁迫对水稻孕穗期生理指标的影响[J]. 灌溉排水学报, 2019, 38(3): 1-7.
	Zhen B, Zhou X G, Lu H F, Li H Z. The effects of alternate hot wave and waterlogging on physiological traits of rice at booting stage[J]. Journal of Irrigation and Drainage, 2019, 38(3): 1-7. (in Chinese with English abstract)
[22]	向丽霞, 胡立盼, 孟森, 邹志荣. 叶面喷施亚精胺对高温胁迫下番茄叶绿素合成代谢的影响[J]. 西北植物学报, 2020, 40(5): 846-851.
	Xiang L X, Hu L P, Meng S, Zou Z R. Effects of foliar-spraying spermidine on chlorophyll synthesis metabolism of tomato seedlings under heat[J]. Acta Botanica Boreali-Occidentalia Sinica, 2020, 40(5): 846-851. (in Chinese with English abstract)
[23]	吴思佳, 李仁英, 谢晓金, 张婍, 陈佳林, 徐向华, 胡宗荟, 卢炳浩, 张娜. 抽穗期高温对水稻叶片光合特性、叶绿素荧光特性和产量构成因素的影响[J]. 南方农业学报, 2021, 52(1): 20-27.
	Wu S J, Li R Y, Xie X J, Zhang Q, Chen J L, Xu X H, Hu Z H, Lu B H, Zhang N. Effects of high temperature on characteristics of photosynthesis and chlorophyll fluorescence and yield components of rice at heading stage[J]. Journal of Southern Agriculture, 2021, 52(1): 20-27. (in Chinese with English abstract)
[24]	Flexas J, Bota J, Loreto F, Cornic G, Sharkey T D. Diffusive and metabolic limitations to photosynthesis under drought and salinity in C3 plants[J]. Plant Biology, 2004, 6(3): 269-279.
[25]	Grassi G, Magnani F. Stomatal, mesophyll conductance and biochemical limitations to photosynthesis as affected by drought and leaf ontogeny in ash and oak trees[J]. Plant, Cell & Environment, 2005, 28(7): 834-849.
[26]	Kumar A, Nayak A K, Hanjagi P S, Kumari K, Vijayakumar S, Mohanty S, Tripathi R, Panneerselvam P. Submergence stress in rice: Adaptive mechanisms, coping strategies and future research needs[J]. Environmental and Experimental Botany, 2021, 186: 104448.
[27]	徐涛, 才硕, 时红, 时元智, 谢亨旺, 刘方平, 梁举. 拔节期淹水胁迫对水稻叶片酶活性及产量的影响[J]. 中国农村水利水电, 2020, 457(11): 89-93.
	Xu T, Cai S, Shi H, Shi Y Z, Xie H W, Liu F P, Liang J. The effects of different flooding stresses on enzymatic activity of the blade and the yield of rice at elongation stage[J]. China Rural Water and Hydropower, 2020, 457(11): 89-93. (in Chinese with English abstract).
[28]	李学昊, 罗强, 秦婷婷, 袁帅, 程伦国, 刘路广. 暴雨后不同滞蓄水深对虾稻田中水稻拔节期生长及产量的影响[J]. 中国农村水利水电, 2021, 467(9): 124-127.
	Li X H, Luo Q, Qin T T, Yuan S, Cheng L G, Liu L G. Effects of different detention water depths on rice growth and yield in shrimp rice fields after a heavy rain[J]. China Rural Water and Hydropower, 2021, 467(9): 124-127. (in Chinese with English abstract)
[29]	彭克勤, 夏石头, 李阳生. 涝害对早中稻生理特性及产量的影响[J]. 湖南农业大学学报: 自然科学版, 2001(3): 173-176.
	Peng K Q, Xia S T, Li Y S. Effects of complete submergence on some physiological and yield characteristics of early and middle-season rice[J]. Journal of Hunan Agricultural University: Natural Sciences, 2001(3): 173-176. (in Chinese with English abstract)
[30]	王矿, 王友贞, 汤广民. 水稻在拔节孕穗期对淹水胁迫的响应规律[J]. 中国农村水利水电, 2016(9): 81-87.
	Wang K, Wang Y Z, Tang G M. Experimental study of response rice jointing-booting stage to inundation condition[J]. China Rural Water and Hydropower, 2016(9): 81-87. (in Chinese with English abstract)
[31]	姬静华, 霍治国, 唐力生, 杜尧东, 胡飞. 早稻灌浆期淹水对剑叶理化特性及产量和品质的影响[J]. 中国水稻科学, 2016, 30(2): 181-192.
	Ji J H, Huo Z G, Tang L S, Du Y D, Hu F. Grain yield and quality and physiological and biochemical characteristics of glag leaf in early rice as affected by submergence at filling stage[J]. Chinese Journal of Rice Science, 2016, 30(2): 181-192. (in Chinese with English abstract)
[32]	徐涛, 才硕, 时红, 时元智, 刘方平, 谢亨旺, 梁举. 分蘖期淹水胁迫对早稻生长发育及产量构成的影响[J]. 节水灌溉, 2020, 301(9): 16-20.
	Xu T, Cai S, Shi H, Shi Y Z, Liu F P, Xie H W, Liang J. Effects of waterlogging stress on growth and yield of early rice at tillering stage[J]. Water Saving Irrigation, 2020, 301(9): 16-20. (in Chinese with English abstract).
[33]	Fu J, Jian Y, Wang X, Li L, Ciais P, Zscheischler J, Wang Y, Tang Y, Müller C, Webber H. Extreme rainfall reduces one-twelfth of China’s rice yield over the last two decades[J]. Nature Food, 2023: 1-11.
[34]	逯涛, 曾庆涛, 张文, 王文博, 王政洋, 杨芮, 孙玉岩. 主成分分析及灰色关联度分析综合评价棉花产量与品质[J]. 新疆农业科学, 2023, 60(5): 1099-1109.
	Lu T, Ceng Q T, Zhang W, Wang W B, Wang Z Y, Yang R, Sun Y Y. Comprehensive evaluation of cotton yield and quality by principal component analysis and grey correlation analysis[J]. Xinjiang Agricultural Sciences, 2023, 60(5): 1099-1109. (in Chinese with English abstract).
[35]	袁伟玲, 曹凑贵, 程建平. 水稻产量及构成因素的灰色关联度分析[J]. 湖北农业科学, 2005(2): 24-25.
	Yuan W L, Cao C G, Cheng J P. Gray correlation analysis of rice yield and its compositions[J]. Hubei Agricultural Sciences, 2005(2): 24-25. (in Chinese with English abstract)
[36]	曹燕燕, 张宏套, 郭春强, 葛昌斌, 廖平安, 黄杰, 乔冀良, 齐双丽, 李雷雷. 不同冬小麦品种拔节期低温生理生化反应及其灰色关联度分析[J]. 山东农业科学, 2021, 53(8): 37-42.
	Cao Y Y, Zhang H T, Guo C Q, Ge C B, Liao P A, Huang J, Qiao J L, Qi S L, Li L L. Physiological and biochemical reactions of different winter wheat varieties at jointing stage under low temperature and their grey correlation analysis[J]. Shandong Agricultural Sciences, 2021, 53(8): 37-42. (in Chinese with English abstract)
[37]	胡江龙, 郭林涛, 王友华, 周治国. 棉花渍害恢复的生理指示指标探讨[J]. 中国农业科学, 2013, 46(21): 4446-4453.
	Hu J L, Guo L T, Wang Y H, Zhou Z G. Physiological indicator of cotton plant in recovery from waterlogging damage[J]. Scientia Agricultura Sinica, 2013, 46(21): 4446-4453. (in Chinese with English abstract)
[38]	Ennahli S, Earl H J. Physiological limitations to photosynthetic carbon assimilation in cotton under water stress[J]. Crop Science, 2005, 45(6): 2374-2382.
[39]	冯方剑, 宋敏, 陈全家, 姚正培, 李杨阳, 刘艳, 王兴安, 曲延英. 棉花苗期抗旱相关指标的主成分分析及综合评价[J]. 新疆农业大学学报, 2011, 34(3): 211-217.
	Feng F J, Song M, Chen Q J, Yao Z P, Li Y Y, Liu Y, Wang X A, Qu Y Y. Analysis and comprehensive evaluation on principal component of relative indices of drought resistance at the seedling stage of cotton[J]. Journal of Xinjiang Agricultural University, 2011, 34(3): 211-217. (in Chinese with English abstract)
[40]	周广生, 梅方竹, 周竹青, 朱旭彤. 小麦不同品种耐湿性生理指标综合评价及其预测[J]. 中国农业科学, 2003(11): 1378-1382.
	Zhou G S, Mei F Z, Zhou Z Q, Zhu X T. Comprehensive evaluation and forecast on physiological indices of waterlogging resistance of different wheat varieties[J]. Scientia Agricultura Sinica, 2003(11): 1378-1382. (in Chinese with English abstract)

pH	土壤速效养分含量Soil available nutrient content				每盆肥料用量 Application level per pot
pH	碱解氮 Alkaline nitrogen /(mg·kg⁻¹)	速效磷 Available nitrogen /(mg·kg⁻¹)	速效钾 Available nitrogen /(mg·kg⁻¹)		尿素 Urea/g	氯化钾 Potassium chloride/g	过磷酸钙 Superphosphate/g
7.6	69.4	28.7	118.7		2.5(70%基肥、30%分蘖肥) 2.5(70% as basal fertilizer, 30% for tillering)	1.95(基肥) 1.95(Basal fertilizer)	1.13(基肥) 1.13(Basal fertilizer)

pH	土壤速效养分含量Soil available nutrient content				每盆肥料用量 Application level per pot
pH	碱解氮 Alkaline nitrogen /(mg·kg⁻¹)	速效磷 Available nitrogen /(mg·kg⁻¹)	速效钾 Available nitrogen /(mg·kg⁻¹)		尿素 Urea/g	氯化钾 Potassium chloride/g	过磷酸钙 Superphosphate/g
7.6	69.4	28.7	118.7		2.5(70%基肥、30%分蘖肥) 2.5(70% as basal fertilizer, 30% for tillering)	1.95(基肥) 1.95(Basal fertilizer)	1.13(基肥) 1.13(Basal fertilizer)

指标 Indicator	年份 Year(Y)	淹涝天数 Flooding days(D)	淹涝深度 Flooding depth(H)	Y×D	Y×H	D×H	Y×D×H
株高增长量Plant height growth(PHG)	3.27	33.02**	40.25**	1.33	0.62	2.93*	0.27
第4节间长Length of the 4th internode(L4I)	0.38	3.81*	54.22**	0.06	0.08	NS	0.01
单穗干质量Dry weight per panicle(DWPP)	0.06	33.68**	92.72**	1.26	0.22	4.78**	0.36
剑叶MDA Flag leaf MDA(F-MDA)	1.52	NS	503.80**	1.20	1.15	NS	0.69
根系MDA Root MDA(R-MDA)	0.35	44.51**	123.88**	0.04	0.25	72.59**	0.32
叶绿素a含量Chl a content	0.03	103.11**	222.31**	0.03	0.04	9.12*	0.03
叶绿素b含量Chl b content	0.25	4.69*	35.93**	0.34	0.02	NS	0.01
可溶性糖含量Soluble sugar content(SS)	0.02	6.74**	136.62**	1.26	1.39	10.04**	0.58
可溶性蛋白含量Soluble protein content(SP)	1.58	30.44**	253.54**	1.69	2.07	NS	0.59
剑叶POD活性 Flag leaf POD activity(F-POD)	0.01	260.61**	751.44**	0.06	0.21	24.61**	0.06
根系POD活性 Root POD activity(R-POD)	0.09	170.49**	69.58**	69.58	0.57	43.77**	0.27
剑叶SOD活性 Flag leaf SOD activity(F-SOD)	0.71	286.95**	130.64**	0.63	0.57	166.97**	0.48
根系SOD活性 Root SOD activity(R-SOD)	0.46	121.99**	132.64**	0.07	1.84	126.59**	3.21**
净光合速率P_n	1.71	89.68**	464.66**	0.04	0.01	5.99**	0.06
气孔导度G_s	0.06	359.18**	1666.49**	1.68	0.06	21.44**	0.21
胞间CO₂浓度C_i	0.31	30.27**	81.00**	0.09	0.04	2.61*	0.01
蒸腾速率T_r	0.00	72.17**	115.21**	0.00	0.00	3.23**	0.00

指标 Indicator	年份 Year(Y)	淹涝天数 Flooding days(D)	淹涝深度 Flooding depth(H)	Y×D	Y×H	D×H	Y×D×H
株高增长量Plant height growth(PHG)	3.27	33.02**	40.25**	1.33	0.62	2.93*	0.27
第4节间长Length of the 4th internode(L4I)	0.38	3.81*	54.22**	0.06	0.08	NS	0.01
单穗干质量Dry weight per panicle(DWPP)	0.06	33.68**	92.72**	1.26	0.22	4.78**	0.36
剑叶MDA Flag leaf MDA(F-MDA)	1.52	NS	503.80**	1.20	1.15	NS	0.69
根系MDA Root MDA(R-MDA)	0.35	44.51**	123.88**	0.04	0.25	72.59**	0.32
叶绿素a含量Chl a content	0.03	103.11**	222.31**	0.03	0.04	9.12*	0.03
叶绿素b含量Chl b content	0.25	4.69*	35.93**	0.34	0.02	NS	0.01
可溶性糖含量Soluble sugar content(SS)	0.02	6.74**	136.62**	1.26	1.39	10.04**	0.58
可溶性蛋白含量Soluble protein content(SP)	1.58	30.44**	253.54**	1.69	2.07	NS	0.59
剑叶POD活性 Flag leaf POD activity(F-POD)	0.01	260.61**	751.44**	0.06	0.21	24.61**	0.06
根系POD活性 Root POD activity(R-POD)	0.09	170.49**	69.58**	69.58	0.57	43.77**	0.27
剑叶SOD活性 Flag leaf SOD activity(F-SOD)	0.71	286.95**	130.64**	0.63	0.57	166.97**	0.48
根系SOD活性 Root SOD activity(R-SOD)	0.46	121.99**	132.64**	0.07	1.84	126.59**	3.21**
净光合速率P_n	1.71	89.68**	464.66**	0.04	0.01	5.99**	0.06
气孔导度G_s	0.06	359.18**	1666.49**	1.68	0.06	21.44**	0.21
胞间CO₂浓度C_i	0.31	30.27**	81.00**	0.09	0.04	2.61*	0.01
蒸腾速率T_r	0.00	72.17**	115.21**	0.00	0.00	3.23**	0.00

年份 Year	回归关系式 Regression equation	标准化回归系数绝对值 Absolute value of standardized regression coefficient		相关系数 Correlation coefficients	F值 F-value
年份 Year	回归关系式 Regression equation	D	H	相关系数 Correlation coefficients	F值 F-value
2017	Y_PH=1.613D+0.121H−4.793	0.578	0.751	0.95	52.89**
2018	Y_PH=2.517D+0.156H−6.747	0.688	0.704	0.97	96.85**