根际氧环境对水稻幼苗根系微形态结构的影响及其生理机制

doi:10.16819/j.1001-7216.2022.211103

摘要/Abstract

摘要：

【目的】研究根际氧环境对水稻幼苗根系微形态建成的影响及其生理机制，以期为水稻幼苗理想根型的构建提供理论依据。【方法】以春优84和秀水09为材料，用国际水稻所营养液配方进行水培试验。秧苗移栽一周后用在线溶氧仪(氮气、氧气调节)设定低氧(0~1.0 mg/L)、中氧(2.5~3.5 mg/L)和高氧(>6.0 mg/L，饱和溶解氧处理，在水稻生长过程中用充气泵连续向水体中充入空气)和常规水培(CK，不进行氧调节，水稻移栽一周后水中氧含量约为0.3~2.5 mg/L) 4个氧处理，研究水稻幼苗生物量、根系微形态结构、根系分泌有机酸、呼吸强度等指标。【结果】1）各处理水稻幼苗根系直径为d₁(0~0.50 mm)、d₂(0.51~1.00 mm)和d₃(1.01~1.50 mm)的细根占总根的比例均超过90%，对水稻苗期根系形态构建尤为重要。中氧处理后，春优84直径为d₁、d₂和秀水09直径为d₁、d₃的根的根长和根表面积均明显高于对照。2）低氧处理减少地上部和地下部生物量，抑制细根(d₁)生长，促进粗根(d₅, 2.01~2.50 mm)的生长，降低呼吸强度，增加有机酸分泌量。中氧和高氧处理后水稻植株的生物量均有增加，但生理反应不尽相同。中氧处理后水稻根系呼吸强度上升，而高氧处理后根系呼吸强度降低。各氧处理均减少水稻幼苗根系生长素(IAA)含量，中氧和高氧处理与对照间差异达显著水平。高氧处理显著增加根系一氧化氮(NO)含量，其他处理间差异不显著。3）相关分析结果表明，两供试品种总根和细根(d₂)的各形态指标(根长、表面积和体积)均与根系有机酸分泌量、呼吸强度、NO含量正相关，与生长素(IAA)含量负相关。【结论】适当增氧(中氧)处理增加水稻幼苗根系呼吸强度和有机酸分泌量，根系生理活性增强;增加细根(直径为0~1.50 mm)的根长、吸收面积的占比，优化其根系形态结构。因此，生产上可以通过栽培措施改变根际氧环境调控分蘖期水稻根系微形态结构，增强根系功能。

关键词: 水稻, 根际氧环境, 根系生理, 根系微形态结构

Abstract:

【Objective】This study was conducted to research the effect of rhizosphere oxygen environment on the root micromorphological structure of rice seedlings and its physiological mechanism, so as to lay a theoretical basis for the construction of ideal root type of rice.【Method】Two varieties, Chunyou 84 and Xiushui 09, were cultured in nutrient solution with different oxygen content (OC) levels (low oxygen, OC 0－1.0 mg/L; medium oxygen, OC 2.5－3.5 mg/L; high oxygen, OC >6.0 mg/L, continuous bubbling with atmospheric air, and normal conditions as CK, conventional hydroponics without oxygen regulation, the oxygen content in water approximately 0.3－2.5 mg/L) controlled by on-line dissolved oxygen meters. After different oxygen treatments, the biomass, root micromorphological structure, and some root physiological indexes (organic acid content, respiratory intensity, and so on) of rice seedlings were measured and analyzed.【Result】1) The proportions of fine roots (d₁, 0－0.50 mm; d₂, 0.51－1.00 mm; d₃, 1.01－1.50 mm) were more than 90%, which were particularly important for the construction of root morphology for rice seedlings. At medium oxygen level, the root length and root surface area of Chunyou 84 with diameter d₁ and d₂, and Xiushui 09 with diameter d₁ and d₃ were significantly higher than those of the control. 2) Low oxygen treatment reduced the biomass of shoot and root, inhibited the growth of fine roots (d₁), promoted the growth of coarse roots (d₅, 2.01－2.50 mm), reduced the root respiration rate and increased the secretion of organic acids. Medium and high oxygen treatments both increased the biomass of rice plants, but had different effects on root physiology. Medium oxygen treatment increased root respiration rate, while high oxygen treatment decreased. Different oxygen treatments reduced the content of root auxin (IAA). Especially, there was obvious difference between medium and high oxygen treatments with the control. The content of nitric oxide (NO) increased significantly under the high oxygen treatment, but there was no significant difference among other treatments. 3) The results of correlation analysis showed that the morphological indexes (root length, surface area, and volume) of the total and fine roots (d₂) of the two varieties were positively correlated with root organic acid secretion, respiratory intensity, and nitric oxide content, on the contrary, they were negatively correlated with auxin content.【Conclusion】Moderate increase of dissolved oxygen concentration (medium oxygen treatment) could improve the root respiration rate and organic acid secretion of rice seedlings, enhancing the root function and increasing the proportion of the length and absorption area of fine root (d₁－d₃), optimizing the root morphological structure. Therefore, appropriate cultivation measures could change the rhizosphere oxygen environment, regulate the micro morphological structure of rice roots at tillering stage and enhance the function of roots in field.

Key words: rice, rhizosphere oxygen environment, root physiology, root micro morphological structure

肖德顺, 徐春梅, 王丹英, 章秀福, 陈松, 褚光, 刘元辉. 根际氧环境对水稻幼苗根系微形态结构的影响及其生理机制[J]. 中国水稻科学, 2022, 36(4): 399-409.

XIAO Deshun, XU Chunmei, WANG Danying, ZHANG Xiufu, CHEN Song, CHU Guang, LIU Yuanhui. Effect of Rhizosphere Oxygen Environment on the Root Micromorphological Structure of Rice Seedlings and Its Physiological Mechanism[J]. Chinese Journal OF Rice Science, 2022, 36(4): 399-409.

图/表 8

参考文献 47

[1]	李香玲, 冯跃华. 水稻根系生长特性及其与地上部分关系的研究进展[J]. 中国农学通报, 2015, 31(6): 1-6.
	Li X L, Feng Y H. Research advance on relation of aerial part and root traits of rice[J]. Chinese Agricultural Science Bulletin, 2015, 31(6): 1-6. (in Chinese with English abstract)
[2]	石庆华, 黄英金, 李木英, 徐益群, 谭雪明, 张佩莲. 水稻根系性状与地上部的相关及根系性状的遗传研究[J]. 中国农业科学, 1997, 30(4): 62-68.
	Shi Q H, Huang Y J, Li M Y, Xu Y Q, Tan X M, Zhang P L. Studies on the heredity of root characteristics and correlation between the characteristics of roots and upperground parts in rice[J]. Scientia Agricultura Sinica, 1997, 30(4): 62-68. (in Chinese with English abstract)
[3]	张福锁, 曹一平. 根际动态过程与植物营养[J]. 土壤学报, 1992, 29(3): 239-250.
	Zhang F S, Cao Y P. Rhizosphere dynamics and plant nutrition[J]. Acta Pedologica Sinica, 1992, 29(3): 239-250. (in Chinese with English abstract)
[4]	Kiba T, Krapp A. Plant nitrogen acquisition under low availability: Regulation of uptake and root architecture[J]. Plant and Cell Physiology, 2016, 57(4): 707-714.
[5]	赵霞, 杜朝云, 徐春梅, 杨华伟, 吕泽林, 章秀福. 水稻对低氧环境的适应及其机制研究进展[J]. 作物杂志, 2015, 31(3): 5-12.
	Zhao X, Du Z Y, Xu C M, Yang H W, Lü Z L, Zhang X F. Review of acclimating mechanism to hypoxia condition in rice[J]. Crops, 2015, 31(3): 5-12. (in Chinese with English abstract)
[6]	Colmer T D, Cox M C H, Voesenek L A C J. Root aeration in rice (Oryza sativa): Evaluation of oxygen, carbon dioxide, and ethylene as possible regulators of root acclimatizations[J]. New Phytologist, 2006, 170(4): 767-777.
[7]	胡继杰, 钟楚, 胡志华, 张均华, 曹小闯, 刘守坎, 金千瑜, 朱练峰. 溶解氧浓度对水稻分蘖期根系生长及氮素利用特性的影响[J]. 中国农业科学, 2021, 54(7): 1525-1536.
	Hu J J, Zhong C, Hu Z H, Zhang J H, Cao X C, Liu S K, Jin Q Y, Zhu L F. Effects of dissolved oxygen concentration on root growth at tillering stage and nitrogen utilization characteristics of rice[J]. Scientia Agricultura Sinica, 2021, 54(7): 1525-1536. (in Chinese with English abstract)
[8]	胡志华, 朱练峰, 林育炯, 胡继杰, 张均华, 金千瑜. 根际氧浓度对水稻产量及其氮素利用的影响[J]. 中国水稻科学, 2015, 29(4): 382-389.
	Hu Z H, Zhu L F, Lin Y J, Hu J J, Zhang J H, Jin Q Y. Effects of rhizosphere oxygen concentration rice grain yield and nitrogen utilization[J]. Chinese Journal of Rice Science, 2015, 29(4): 382-389. (in Chinese with English abstract)
[9]	徐春梅, 王丹英, 陈松, 陈丽萍, 章秀福. 增氧对水稻根系生长与氮代谢的影响[J]. 中国水稻科学, 2012, 26(3): 320-324.
	Xu C M, Wang D Y, Chen S, Chen L P, Zhang X F. Effect of aeration on root growth and nitrogen metabolism in rice[J]. Chinese Journal of Rice Science, 2012, 26(3): 320-324. (in Chinese with English abstract)
[10]	徐春梅, 陈丽萍, 王丹英, 陈松, 章秀福, 石庆华. 低氧胁迫对水稻幼苗根系功能和氮代谢相关酶活性的影响[J]. 中国农业科学, 2016, 49(8): 1625-1634.
	Xu C M, Chen L P, Wang D Y, Zhang X F, Shi Q H. Effects of low oxygen stress on the root function and enzyme activities related to nitrogen metabolism in roots of rice seedlings[J]. Scientia Agricultura Sinica, 2016, 49(8): 1625-1634. (in Chinese with English abstract)
[11]	茆智. 水稻节水灌溉及其对环境的影响[J]. 中国工程科学, 2002, 4(7): 8-16.
	Mao Z. Water saving irrigation for rice and its effect on environment[J]. Engineering Science, 2002, 4(7): 8-16. (in Chinese with English abstract)
[12]	赵锋, 王丹英, 徐春梅, 张卫建, 李凤博, 毛海军, 章秀福. 根际增氧模式的水稻形态、生理及产量响应特征[J]. 作物学报, 2010, 36(2): 303-312.
	Zhao F, Wang D Y, Xu C M, Zhang W J, Li F B, Mao H J, Zhang X F. Response of morphological, physiological and yield characteristics of rice (Oryza sativa L.) to different oxygen-increasing patterns in rhizosphere[J]. Acta Agronomica Sinica, 2010, 36(2): 303-312. (in Chinese with English abstract)
[13]	朱练峰, 刘学, 禹盛苗, 欧阳由男, 金千瑜. 增氧灌溉对水稻生理特性和后期衰老的影响[J]. 中国水稻科学, 2010, 24(3): 257-263.
	Zhu L F, Liu X, Yu S M, Ouyang Y N, Jin Q Y. Effects of aerated irrigation on physiological characteristics and senescence at late growth stage of rice[J]. Chinese Journal of Rice Science, 2010, 24(3): 257-263. (in Chinese with English abstract)
[14]	Thakup A K, Mandal K G, Mohanty R K, Ambast S K. Rice root growth, photosynthesis, yield and water productivity improvements through modifying cultivation practices and water management[J]. Agricultural Water Management, 2018, 206: 67-77.
[15]	Drescher G L, Silva L S D, Sarfaraz Q, Roberts T L, Nicoloso F T, Schwalbert R, Marques A C R. Available nitrogen in paddy soils depth: Influence on rice root morphology and plant nutrition[J]. Journal of Soil Science and Plant Nutrition, 2020, 20(3): 1029-1041.
[16]	Nicola C, Christina A, Eva B. Auxin-regulated lateral root organogenesis[J]. Cold Spring Harbor Perspectives in Biology, 2021, 13(7): a039941.
[17]	Xu C, Chen L, Chen S, Chu G, Wang D, Zhang X. Effects of rhizosphere oxygen concentration on root physiological characteristics and anatomical structure at the tillering stage of rice[J]. Annals of Applied Biology, 2020, 177(1): 61-73.
[18]	Xu C M, Wang D Y, Chen S, Chen L P, Zhang X F. Effects of aeration on root physiology and nitrogen metabolism in rice[J]. Rice Science, 2013, 20(2): 148-253.
[19]	吴龙龙, 田仓, 张露, 黄晶, 朱练峰, 张均华, 曹小闯, 金千瑜. 稻田水氮氧环境因子对水稻生长发育、光合作用和氮利用的调控研究进展[J]. 应用生态学报, 2021, 32(4): 1498-1508.
	Wu L L, Tian C, Zhang L, Huang J, Zhu L F, Zhang J H, Cao X C, Jin Q Y. Research advance in the roles of water-nitrogen-oxygen factors in mediating rice growth, photosynthesis and nitrogen utilization in paddy soils[J]. Chinese Journal of Applied Ecology, 2021, 32(4): 1498-1508. (in Chinese with English abstract)
[20]	徐国伟, 陆大克, 刘聪杰, 王贺正, 陈明灿, 李友军. 干湿交替灌溉和施氮量对水稻内源激素及氮素利用的影响[J]. 农业工程学报, 2018, 34(7): 137-146.
	Xu G W, Lu D K, Liu C J, Wang H Z, Chen M C, Li Y J. Effect of alternate wetting and drying irrigation and nitrogen coupling on endogenous hormones, nitrogen utilization[J]. Transactions of the Chinese Society of Agricultural Engineering, 2018, 34(7): 137-146. (in Chinese with English abstract)
[21]	何俊瑜, 王阳阳, 任艳芳, 周国强, 杨良静. 镉胁迫对不同水稻品种幼苗根系形态和生理特性的影响[J]. 生态环境学报, 2009, 18(5): 1863-1868.
	He J Y, Wang Y Y, Ren Y F, Zhou G Q, Yang L J. Effect of cadmium on root morphology and physiological characteristics of rice seedlings[J]. Ecology and Environmental Sciences, 2009, 18(5): 1863-1868.
[22]	郜红建, 常江, 张自立, 丁士明, 魏俊岭. 研究植物根系分泌物的方法[J]. 植物生理学通讯, 2003, 39(1): 56-60.
	Gao H J, Chang J, Zhang Z L, Ding S M, Wei J L. Methods for investigating root exudates of plants[J]. Plant Physiology Communications, 2003, 39(1): 56-60. (in Chinese with English abstract)
[23]	Kelm M, Dahmann R, Wink D, Feelisch M. The nitric oxide superoxide assay: Insights into the biological chemistry of the NO/O₂^― interaction[J]. Journal of Biological Chemistry, 1997, 272(15): 9922-9932.
[24]	杨建昌. 水稻根系形态生理与产量、品质形成及养分吸收利用的关系[J]. 中国农业科学, 2011, 44(1): 36-46.
	Yang J C. Relationships of rice root morphology and physiology with the formation of grain yield and quality and the nutrient absorption and utilization[J]. Scientia Agricultura Sinica, 2011, 44(1): 36-46. (in Chinese with English abstract)
[25]	Yang J C, Zhang H, Zhang J H. Root morphology and physiology in relation to the yield formation of rice[J]. Journal of Integrative Agriculture, 2012, 11(6): 920-926.
[26]	章怡兰, 林雪, 吴仪, 李梦佳, 张晟婕, 路梅, 饶玉春, 王跃星. 水稻根系遗传育种研究进展[J]. 植物学报, 2020, 55(3): 382-393.
	Zhang Y L, Lin X, Wu Y, Li M J, Zhang S J, Lu M, Rao Y C, Wang Y X. Research progress on genetics and breeding of rice roots[J]. Chinese Bulletin of Botany, 2020, 55(3): 382-393. (in Chinese with English abstract)
[27]	丁仕林, 刘朝雷, 钱前. 水稻根系遗传研究进展[J]. 中国稻米, 2019, 25(5): 24-29.
	Ding S L, Liu Z L, Qian Q. Research progress on genetic of rice root[J]. China Rice, 2019, 25(5): 24-29. (in Chinese with English abstract)
[28]	王昱, 王玮, 范杰英, 姜晓莉. 水稻根系分布同产量之间关系的研究进展[J]. 农业与技术, 2017, 37(5): 43-45.
	Wang Y, Wang W, Fan J Y, Jiang X L. Research progress on the relationship between rice root distribution and yield[J]. Agriculture and Technology, 2017, 37(5): 43-45. (in Chinese with English abstract)
[29]	王丹英, 韩勃, 章秀福, 邵国胜, 徐春梅, 符冠富. 水稻根际含氧量对根系生长的影响[J]. 作物学报, 2008, 34(5): 803-808.
	Wang D Y, Han B, Zhang X F, Shao G S, Xu C M, Fu G F. Influence of rhizosphere oxygen concentration on rice root growth[J]. Acta Agronomica Sinica, 2008, 34(5): 803-808. (in Chinese with English abstract)
[30]	赵霞, 徐春梅, 王丹英, 陈松, 陶龙兴, 章秀福. 根际溶氧量对分蘖期水稻生长特性及其氮素代谢的影响[J]. 中国农业科学, 2015, 48(18): 3733-3742.
	Zhao X, Xu C M, Wang D Y, Chen S, Tao L X, Zhang X F. Effect of rhizosphere oxygen on the growth characteristics of rice and its nitrogen metabolism at tillering stage[J]. Scientia Agricultura Sinica, 2015, 48(18): 3733-3742. (in Chinese with English abstract)
[31]	牛耀芳, 宗晓波, 都韶婷, 黄利东, 章永松. 大气CO₂浓度升高对植物根系形态的影响及其调控机理[J]. 植物营养与肥料学报, 2011, 17(1): 240-246.
	Niu Y F, Zong X B, Du S T, Huang L D, Zhang Y S. Effect of elevated CO₂ on morphology change of plant roots and its regulatory mechanism[J]. Journal of Plant Nutrition and Fertilizers, 2011, 17(1): 240-246. (in Chinese with English abstract)
[32]	孙虎威, 王文亮, 刘尚俊, 侯蒙蒙, 谢天宁, 梁志浩, 樊亚男, 张亚丽. 低氮胁迫下水稻根系的发生及生长素的响应[J]. 土壤学报, 2014, 51(5): 1096-1102.
	Sun H W, Wang W L, Liu S J, Hou M M, Xie T Y, Liang Z H, Fan Y N, Zhang Y L. Formation of rice root regulated by nitrogen deficiency[J]. Acta Pedologica Sinica, 2014, 51(5): 1096-1102. (in Chinese with English abstract)
[33]	宁倩, 吴金水, 李宝珍, 吴蔓莉. 水稻苗期生长特性对不同浓度IAA的响应[J]. 农业现代化研究, 2013, 34(2): 235-238.
	Ning Q, Wu J S, Li B Z, Wu M L. Response of Growth characteristics to different IAA concentrations in rice seedling[J]. Research of Agricultural Modernization, 2013, 34(2): 235-238. (in Chinese with English abstract)
[34]	李维芳, 王春蕾, 王妮, 邓雨正, 姚彦东, 魏丽娟, 廖伟彪. 一氧化氮影响植物不定根发生的研究进展[J]. 浙江农业学报, 2020, 32(4): 742-752.
	Li W F, Wang C L, Wang N, Deng Y Z, Yao Y D, Wei L J, Liao W B. Research progress on effect of nitric oxide on adventitious formation in plant[J]. Acta Agriculturae Zhejiangensis, 2020, 32(4): 742-752. (in Chinese with English abstract)
[35]	刘光亚, 张艳军, 孙学振, 董合忠. 一氧化氮对植物淹水伤害的缓解作用及其机制[J]. 分子植物育种, 2019, 17(22): 7579-7587.
	Liu G Y, Zhang Y J, Sun X Z, Dong H Z. Roles and underlying mechanisms of nitric oxide in alleviating waterlogging damage to plants[J]. Molecular Plant Breeding, 2019, 17(22): 7579-7587. (in Chinese with English abstract)
[36]	王金祥, 陈碧丽, 廖红, 严小龙. 生长素、乙烯和一氧化氮对拟南芥下胚轴插条形成不定根的调节[J]. 植物生理学通讯, 2009, 45(10): 986-990.
	Wang J X, Chen B L, Liao H, Yan X L. Regulation of auxin, ethylene and nitric oxide on adventitious rooting in Arabidopsis hypocotyl cuttings[J]. Plant Physiology Communications, 2009, 45(10): 986-990. (in Chinese with English abstract)
[37]	Correa-Aragunde N, Cejudo F J, Lamattina L. Nitric oxide is required for the auxin-induced activation of NADPH-dependent thioredoxin reductase and protein denitrosylation during root growth responses in arabidopsis[J]. Annals of Botany, 2015, 116(4): 695-702.
[38]	王金祥, 严小龙, 潘瑞炽. 不定根形成与植物激素的关系[J]. 植物生理学通讯, 2005, 41(2): 133-142.
	Wang J X, Yan X L, Pan R C. Relationship between adventitious root formation and plant hormones[J]. Plant Physiology Communications, 2005, 41(2): 133-142. (in Chinese with English abstract)
[39]	陈佳宝, 闫道良, 郑炳松. 植物不定根诱导生成机制研究进展[J]. 北方园艺, 2021(6): 129-137.
	Chen J B, Yan D L, Zheng B S. Research progress of the mechanism on adventitious root induction in plant[J]. Northern Horticulture, 2021(6): 129-137. (in Chinese with English abstract)
[40]	许璐璐, 王涵, 高盼盼, 吴三桥, 张辰露. 环境胁迫对植物根系形态的影响[J]. 安徽农业科学, 2020, 48(14): 16-19.
	Xu L L, Wang H, Gao P P, Wu S Q, Zhang C L. Effects of environmental stress on plant root morphology[J]. Journal of Anhui Agricultural Sciences, 2020, 48(14): 16-19. (in Chinese with English abstract)
[41]	葛才林, 络剑峰, 刘冲, 殷朝珍, 王泽港, 马飞, 罗时石. 重金属对水稻呼吸速率及相关同功酶影响的研究[J]. 农业环境科学学报, 2005, 24(2): 222-226.
	Ge C L, Luo J F, Liu C, Yin C Z, Wang Z G, Ma F, Luo S S. Effects of heavy metals on respiration rate and expression of related isozymes in rice[J]. Journal of Agric-Environmental Sciences, 2005, 24(2): 222-226. (in Chinese with English abstract)
[42]	李志霞, 秦嗣军, 吕德国, 聂继云. 植物根系呼吸代谢及影响根系呼吸的环境因子研究进展[J]. 植物生理学报, 2011, 47(10): 957-966.
	Li Z X, Qin S J, Lü D G, Nie J T. Research progress in root respiratory metabolism of plant and the environ mental influencing factors[J]. Plant Physiology Journal, 2011, 47(10): 957-966. (in Chinese with English abstract)
[43]	师恺, 胡文海, 董德坤, Otieno O J, 宋兴舜, 夏晓剑, 周艳虹, 喻景权. 根系限制对番茄幼苗生长、根系呼吸、ATPase和PPase活性的影响[J]. 园艺学报, 2006, 33(4): 853-855.
	Shi K, Hu W H, Dong D K, Otieno O J, Song X S, Xia X J, Zhou Y H, Yu J Q. Effects of root restriction on plant growth, root respiration, activities of H⁺-ATPase and H⁺-PPase in tomato seedlings[J]. Acta Horticulturae Sinica, 2006, 33(4): 853-855. (in Chinese with English abstract)
[44]	陈静, 王朝云, 易永健, 汪洪鹰, 易镇邪, 周晚来. 短期淹水对水稻幼苗生长和根系氮代谢关键酶活性的影响[J]. 华北农学报, 2021, 36(2): 99-107.
	Chen J, Wang Z Y, Yi Y J, Wang H Y, Yi Z X, Zhou W L. Effects of short-term submergence on growth of rice seedlings and activities of key nitrogen metabolism enzymes in roots[J]. Acta Agriculturae Boreali-Sinica, 2021, 36(2): 99-107. (in Chinese with English abstract)
[45]	丁杰萍, 罗永清, 周欣, 岳祥飞, 连杰. 植物根系呼吸研究方法及影响因素研究进展[J]. 草业学报, 2015, 24(5): 206-216.
	Ding J P, Luo Y Q, Zhou X, Yue X F, Lian J. Review of methodology and factors influencing plant root respiration[J]. Acta Prataculturae Sinica, 2015, 24(5): 206-216. (in Chinese with English abstract)
[46]	Liang S, Wang Y H, Zhang H, Yun X Y, Wu Y. Response of root-exuded organic acids in irrigated rice to different water management practices[J]. Eurasian Soil Science, 2020, 53(11): 1572-1578.
[47]	Bi J G, Hou D P, Zhang X X, Tan J S, Bi Q Y, Zhang K K, Liu Y, Wang F M, Zhang A N, Chen L, Liu G L, Liu Z C, Yu X Q, Luo L J. A novel water-saving and drought-resistance rice variety promotes phosphorus absorption through root secreting organic acid compounds to stabilize yield under water-saving condition[J]. Journal of Cleaner Production, 2021, 315: 127992.

品种 Variety	处理 Treatment	总根长 Total root length	不同直径根长 Length of roots with different diameters
品种 Variety	处理 Treatment	总根长 Total root length	d₁ (0~0.50 mm)	d₂ (0.51~1.00 mm)	d₃ (1.01~1.50 mm)	d₄ (1.51~2.00 mm)	d₅ (2.01~2.50 mm)
春优84 Chunyou 84	对照 Control	3570.30 a	2091.56 a	1121.01 b	335.62 b	13.66 a	8.44 a
	低氧 Low oxygen	3575.39 a	2077.17 a	1041.31 b	434.81 a	12.56 ab	9.54 a
	中氧 Medium oxygen	3816.75 a	2061.05 a	1403.56 a	344.33 b	7.57 b	0.24 b
	高氧 High oxygen	3758.83 a	2225.89 a	1152.14 b	368.87 b	11.35 ab	0.57 b
秀水09 Xiushui 09	对照 Control	2411.08 a	1284.53 a	998.64 b	112.95 a	7.91 a	7.05 a
	低氧 Low oxygen	1856.02 b	749.97 c	985.88 b	112.73 a	0.23 c	7.22 a
	中氧 Medium oxygen	2574.64 a	1343.62 a	1066.98 b	157.33 a	5.00 b	1.71 b
	高氧 High oxygen	2496.34 a	1150.37 b	1200.66 a	136.05 a	8.61 a	0.65 b
方差分析ANOVA
品种 Variety (V)		170.30 ^**	240.74 ^**	5.83 ^*	268.94 ^**	17.31 ^**	0.50 ns
处理 Treatment (T)		4.15 ns	5.21 ^*	4.60 ^*	2.44 ns	2.82 ns	18.12 ^**
V × T		1.44 ns	5.02 ^*	3.26 ns	3.18 ns	2.66 ns	0.95 ns

品种 Variety	处理 Treatment	总根长 Total root length	不同直径根长 Length of roots with different diameters
品种 Variety	处理 Treatment	总根长 Total root length	d₁ (0~0.50 mm)	d₂ (0.51~1.00 mm)	d₃ (1.01~1.50 mm)	d₄ (1.51~2.00 mm)	d₅ (2.01~2.50 mm)
春优84 Chunyou 84	对照 Control	3570.30 a	2091.56 a	1121.01 b	335.62 b	13.66 a	8.44 a
	低氧 Low oxygen	3575.39 a	2077.17 a	1041.31 b	434.81 a	12.56 ab	9.54 a
	中氧 Medium oxygen	3816.75 a	2061.05 a	1403.56 a	344.33 b	7.57 b	0.24 b
	高氧 High oxygen	3758.83 a	2225.89 a	1152.14 b	368.87 b	11.35 ab	0.57 b
秀水09 Xiushui 09	对照 Control	2411.08 a	1284.53 a	998.64 b	112.95 a	7.91 a	7.05 a
	低氧 Low oxygen	1856.02 b	749.97 c	985.88 b	112.73 a	0.23 c	7.22 a
	中氧 Medium oxygen	2574.64 a	1343.62 a	1066.98 b	157.33 a	5.00 b	1.71 b
	高氧 High oxygen	2496.34 a	1150.37 b	1200.66 a	136.05 a	8.61 a	0.65 b
方差分析ANOVA
品种 Variety (V)		170.30 ^**	240.74 ^**	5.83 ^*	268.94 ^**	17.31 ^**	0.50 ns
处理 Treatment (T)		4.15 ns	5.21 ^*	4.60 ^*	2.44 ns	2.82 ns	18.12 ^**
V × T		1.44 ns	5.02 ^*	3.26 ns	3.18 ns	2.66 ns	0.95 ns

品种 Variety	处理 Treatment	总根系表面积 Total root surface area	不同直径根表面积 Surface area of roots with different diameters
品种 Variety	处理 Treatment	总根系表面积 Total root surface area	d₁	d₂	d₃	d₄	d₅
春优84 Chunyou 84	对照 Control	517.84 b	113.35 c	248.56 b	125.79 bc	16.41 a	13.74 b
	低氧 Low oxygen	533.10 b	108.61 c	239.19 b	148.45 a	19.18 a	17.67 a
	中氧 Medium oxygen	587.37 a	163.31 a	306.14 a	116.67 c	1.09 b	0.17 c
	高氧 High oxygen	546.69 b	137.79 b	270.36 b	136.77 ab	1.39 b	0.39 c
秀水09 Xiushui 09	对照 Control	369.99 a	88.93 b	212.73 b	46.77 ab	11.04 a	10.52 b
	低氧 Low oxygen	322.59 b	52.55 c	215.13 b	37.57 b	0.12 c	17.23 a
	中氧 Medium oxygen	409.64 a	100.24 a	265.35 a	42.84 b	0.75 c	0.46 c
	高氧 High oxygen	404.58 a	96.93 ab	242.66 a	61.12 a	2.67 b	1.19 c
方差分析ANOVA
品种 Variety (V)		138.20 ^**	116.49 ^**	10.38 ^*	297.33 ^**	48.86 ^**	1.43 ns
处理 Treatment (T)		4.58 ^*	26.77 ^**	7.62 ^*	3.17 ns	53.25 ^**	185.68 ^**
V × T		1.01 ns	4.10 ns	0.22 ns	2.52 ns	30.29 ^**	2.08 ns

品种 Variety	处理 Treatment	总根系表面积 Total root surface area	不同直径根表面积 Surface area of roots with different diameters
品种 Variety	处理 Treatment	总根系表面积 Total root surface area	d₁	d₂	d₃	d₄	d₅
春优84 Chunyou 84	对照 Control	517.84 b	113.35 c	248.56 b	125.79 bc	16.41 a	13.74 b
	低氧 Low oxygen	533.10 b	108.61 c	239.19 b	148.45 a	19.18 a	17.67 a
	中氧 Medium oxygen	587.37 a	163.31 a	306.14 a	116.67 c	1.09 b	0.17 c
	高氧 High oxygen	546.69 b	137.79 b	270.36 b	136.77 ab	1.39 b	0.39 c
秀水09 Xiushui 09	对照 Control	369.99 a	88.93 b	212.73 b	46.77 ab	11.04 a	10.52 b
	低氧 Low oxygen	322.59 b	52.55 c	215.13 b	37.57 b	0.12 c	17.23 a
	中氧 Medium oxygen	409.64 a	100.24 a	265.35 a	42.84 b	0.75 c	0.46 c
	高氧 High oxygen	404.58 a	96.93 ab	242.66 a	61.12 a	2.67 b	1.19 c
方差分析ANOVA
品种 Variety (V)		138.20 ^**	116.49 ^**	10.38 ^*	297.33 ^**	48.86 ^**	1.43 ns
处理 Treatment (T)		4.58 ^*	26.77 ^**	7.62 ^*	3.17 ns	53.25 ^**	185.68 ^**
V × T		1.01 ns	4.10 ns	0.22 ns	2.52 ns	30.29 ^**	2.08 ns

品种 Variety	处理 Treatment	总根系体积 Total root volume	不同直径根体积 Volume of roots with different diameters
品种 Variety	处理 Treatment	总根系体积 Total root volume	d₁	d₂	d₃	d₄	d₅
春优84 Chunyou 84	对照 Control	8.86 bc	0.68 b	4.73 b	3.38 b	0.06 a	0.02 a
	低氧 Low oxygen	8.44 c	0.60 bc	4.55 b	3.10 b	0.14 a	0.03 a
	中氧 Medium oxygen	9.53 a	0.77 a	5.54 a	3.17 b	0.05 a	0.01 a
	高氧 High oxygen	9.22 ab	0.55 c	4.52 b	4.06 a	0.07 a	0.01 a
秀水09 Xiushui 09	对照 Control	5.47 ab	0.60 ab	3.78 c	1.00 ab	0.05 b	0.04 b
	低氧 Low oxygen	5.24 b	0.36 c	3.87 bc	0.87 b	0.01 c	0.13 a
	中氧 Medium oxygen	6.34 a	0.55 b	4.21 ab	1.39 a	0.11 a	0.07 b
	高氧 High oxygen	6.33 a	0.66 a	4.47 a	1.14 ab	0.03 bc	0.03 b
方差分析ANOVA
品种 Variety (V)		221.77 ^**	17.74^**	19.60 ^**	312.30 ^**	0.94 ns	14.59 ^**
处理 Treatment (T)		5.31^*	9.87 ^**	3.21 ns	0.23 ns	0.21 ns	5.74 ^*
V × T		0.28 ns	10.00 ^**	2.68 ns	0.23 ns	2.10 ns	2.89 ns