外源硝普钠对碱胁迫下水稻幼苗糖代谢的影响

doi:10.3864/j.issn.0578-1752.2026.02.004

摘要/Abstract

摘要：

【目的】 外源硝普钠（SNP）是一种常用的一氧化氮（NO）供体，在植物胁迫响应中具有重要的调控作用。探究外源SNP对碱胁迫下水稻幼苗糖代谢的影响，为揭示SNP增强水稻耐碱性的机制提供理论依据。 【方法】 以幼苗期水稻中花11和宁粳52为供试材料，设置对照（CK）、CK+SNP（50 µmol·L^-1）、碱胁迫（AS，将NaHCO₃与Na₂CO₃等比例混合配制成20 mmol·L^-1、pH 9.55的混合碱溶液）、AS+SNP（20 mmol·L^-1 AS+50 µmol·L^-1 SNP）4个处理，分析各处理对水稻幼苗期2个水稻品种糖代谢的影响。 【结果】 与AS处理相比，AS+SNP处理显著提高了水稻幼苗的苗高、根长、鲜重及干重；AS+SNP处理上调了叶绿素合成酶相关基因OsChlH、OsCAO1和OsCAO2的表达，从而增加了叶绿素的含量，促进光合作用；同时，上调表达蔗糖转化酶基因OsNIN1使果糖和蔗糖含量降低，葡萄糖含量升高，有助于糖酵解和三羧酸的良性循环，促进苹果酸和柠檬酸的积累，从而维持能量平衡；此外，通过诱导蔗糖磷酸合酶OsSPS1、赤霉素合成酶相关基因（OsGA3ox2、OsGA20ox1）、脯氨酸合成基因（OsP5CS）的表达，增加了赤霉素（GA）、脱落酸（ABA）、脯氨酸（Pro）和可溶性糖含量，并调节渗透胁迫和激素平衡，促进蔗糖从源向库的运输。 【结论】 外源喷施SNP通过上调叶绿素合成基因，促进碱胁迫下水稻幼苗叶绿素积累并维持光合效率，进而增强葡萄糖合成；同时，通过激活糖酵解和三羧酸循环，使糖类分解产物向苹果酸、柠檬酸等转化，促进有机酸积累。此外，外源SNP通过渗透调节和GA/ABA激素平衡，促进蔗糖运输及糖代谢通路活化，保障能量代谢通量，进而提高了水稻幼苗对碱胁迫的耐受性。

关键词: NO, 碱胁迫, 糖代谢, 基因表达, 生理响应, 水稻

Abstract:

【Objective】 Sodium nitroprusside (SNP), as an exogenous nitric oxide (NO) donor, plays a critical regulatory role in plant stress responses. This study aimed to investigate the effects of exogenous SNP on sugar metabolism in rice seedlings under alkaline stress, for providing a theoretical basis for elucidating the mechanism of SNP enhancing rice alkali tolerance. 【Method】 Seedlings of rice cultivars Zhonghua 11 and Ningjing 52 were used as test materials. Four treatments were applied: control (CK), CK+SNP (50 µmol·L^-1), alkaline stress (AS, a mixed alkaline solution containing NaHCO₃ and Na₂CO₃ at a 1:1 ratio, pH 9.55, 20 mmol·L^-1), and AS+SNP (20 mmol·L^-1 AS+50 µmol·L^-1 SNP). The effects of these treatments on sugar metabolism in the seedlings of the two rice cultivars were analyzed. 【Result】 Compared with AS treatment, AS+SNP treatment significantly increased rice seedling height, root length, fresh weight, and dry weight of rice seedlings. AS+SNP treatment upregulated the expression of chlorophyll synthase-related genes (OsChlH, OsCAO1, and OsCAO2), thereby increasing chlorophyll content and promoting photosynthesis. Concurrently, it upregulated the expression of the sucrose invertase gene OsNIN1, leading to decreased fructose and sucrose content but increased glucose content, which facilitated glycolysis and the tricarboxylic acid cycle, promoting the accumulation of malate and citrate to maintain energy balance. Additionally, SNP induced the expression of sucrose phosphate synthase OsSPS1, gibberellin synthase-related genes (OsGA3ox2, and OsGA20ox1), and proline synthesis gene (OsP5CS), increasing the levels of gibberellin (GA), abscisic acid (ABA), proline (Pro), and soluble sugar. This regulation helped maintain osmotic and hormonal balance and promoted sucrose transport from source to sink tissues. 【Conclusion】 Exogenous SNP application enhanced rice seedling tolerance to alkaline stress by upregulating chlorophyll biosynthesis genes, promoting chlorophyll accumulation, and sustaining photosynthetic efficiency, thereby facilitating glucose synthesis. Moreover, SNP activated glycolysis and the tricarboxylic acid cycle, directing sugar metabolites toward malate and citrate production, thus promoting organic acid accumulation. Additionally, exogenous SNP modulated osmotic regulation and GA/ABA hormonal balance, facilitating sucrose transport and sugar metabolic pathway activation, and ensuring efficient energy metabolism. These mechanisms collectively improved rice seedling resilience under alkaline stress.

Key words: NO, alkaline stress, sugar metabolism, gene expression, physiological response, rice

廖婷璐, 石亚飞, 肖东浩, 舍杨梦斐, 郭富城, 杨九菊, 唐海江, 罗成科. 外源硝普钠对碱胁迫下水稻幼苗糖代谢的影响[J]. 中国农业科学, 2026, 59(2): 265-277.

LIAO TingLu, SHI YaFei, XIAO DongHao, SHE YangMengFei, GUO FuCheng, YANG JiuJu, TANG HaiJiang, LUO ChengKe. The Effect of Exogenous Nitroprusside on Sugar Metabolism in Rice Seedlings Under Alkaline Stress[J]. Scientia Agricultura Sinica, 2026, 59(2): 265-277.

图/表 8

表1

图1

图2

图3

图4

图5

图6

图7

参考文献 49

[1]	FAO. Global status of salt-affected soils. Rome: FAO, 2024.
[2]	冯起, 尹鑫卫, 朱猛, 张举涛, 刘蔚, 席海洋, 鱼腾飞, 杨林山, 刘文, 陆志翔. 统筹推进西北地区盐碱地综合治理利用: 现状、挑战与对策建议. 中国科学院院刊, 2024, 39(12): 2060-2073.
	FENG Q, YIN X W, ZHU M, ZHANG J T, LIU W, XI H Y, YU T F, YANG L S, LIU W, LU Z X. Overall promotion of integrated management and utilization of saline-alkali land in Northwest China: Conditions, challenges, and recommendations. Bulletin of Chinese Academy of Sciences, 2024, 39(12): 2060-2073. (in Chinese)
[3]	马国辉, 郑殿峰, 母德伟, 王奉斌, 戴其根, 魏中伟, 冯乃杰, 王才林. 耐盐碱水稻研究进展与展望. 杂交水稻, 2024, 39(1): 1-10.
	MA G H, ZHENG D F, MU D W, WANG F B, DAI Q G, WEI Z W, FENG N J, WANG C L. Research progress and prospect of saline-alkali tolerant rice. Hybrid Rice, 2024, 39(1): 1-10. (in Chinese)
[4]	王丽娟. 植物转化酶在糖代谢中的作用. 农业科学研究, 2010, 31(4): 70-75.
	WANG L J. Advances on the studies of invertase on sucrose metabolism in higher plant. Journal of Agricultural Science Research, 2010, 31(4): 70-75. (in Chinese)
[5]	郑广杰, 叶昌, 徐春梅, 陈松, 褚光, 陈里鹏, 章秀福, 王丹英. 低温胁迫对水稻胚芽中葡萄糖和水分状态的影响及其与种子出苗成活间的关系. 作物学报, 2024, 50(12): 3055-3068. doi: 10.3724/SP.J.1006.2024.42014
	ZHENG G J, YE C, XU C M, CHEN S, CHU G, CHEN L P, ZHANG X F, WANG D Y. Effect of low-temperature stress on glucose and water status in rice germ and its relationship with seedling emergence. Acta Agronomica Sinica, 2024, 50(12): 3055-3068. (in Chinese) doi: 10.3724/SP.J.1006.2024.42014
[6]	ROLLAND F, MOORE B, SHEEN J. Sugar sensing and signaling in plants. The Plant Cell, 2002, 14(Suppl_1): S185-S205. doi: 10.1105/tpc.010455
[7]	GONG X, LIU M L, ZHANG L J, RUAN Y Y, DING R, JI Y Q, ZHANG N, ZHANG S B, FARMER J, WANG C. Arabidopsis AtSUC2 and AtSUC4, encoding sucrose transporters, are required for abiotic stress tolerance in an ABA-dependent pathway. Physiologia Plantarum, 2015, 153(1): 119-136.
[8]	MA Q J, SUN M H, LU J, LIU Y J, HU D G, HAO Y J. Transcription factor AREB2 is involved in soluble sugar accumulation by activating sugar transporter and amylase genes. Plant Physiology, 2017, 174(4): 2348-2362. doi: 10.1104/pp.17.00502
[9]	李佳馨, 李霞, 谢寅峰. 外源海藻糖增强高表达转玉米C₄型PEPC水稻耐旱性的机制. 植物学报, 2021, 56(3): 296-314.
	LI J X, LI X, XIE Y F. Mechanism on drought tolerance enhanced by exogenous trehalose in C₄-PEPC rice. Chinese Bulletin of Botany, 2021, 56(3): 296-314. (in Chinese)
[10]	王丽华, 李改玲, 李晶, 左师宇, 曹鑫波, 佟昊阳, 魏湜. 外源糖对盐胁迫下小黑麦幼苗糖代谢的影响. 麦类作物学报, 2017, 37(4): 548-553.
	WANG L H, LI G L, LI J, ZUO S Y, CAO X B, TONG H Y, WEI S. Effect of exogenous sugar on the sugar metabolism in Triticale seedling under salt stress. Journal of Triticeae Crops, 2017, 37(4): 548-553. (in Chinese)
[11]	MENG F Y, FENG N J, ZHENG D F, LIU M L, ZHANG R J, HUANG X X, HUANG A Q, CHEN Z M. Exogenous Hemin alleviates NaCl stress by promoting photosynthesis and carbon metabolism in rice seedlings. Scientific Reports, 2023, 13: 3497. doi: 10.1038/s41598-023-30619-7 pmid: 36859499
[12]	陈丽珊, 周红艳, 林伟伟. 外源褪黑素对盐胁迫下水稻苗期碳氮代谢的影响. 生态学杂志, 2023, 42(7): 1635-1643.
	CHEN L S, ZHOU H Y, LIN W W. Effects of exogenous melatonin on carbon and nitrogen metabolism of rice seedlings under salt stress. Chinese Journal of Ecology, 2023, 42(7): 1635-1643. (in Chinese)
[13]	BASIT F, ABBAS S, SHETEIWY M S, BHAT J A, ALSAHLI A A, AHMAD P. Deciphering the alleviation potential of nitric oxide, for low temperature and chromium stress via maintaining photosynthetic capacity, antioxidant defence, and redox homeostasis in rice (Oryza sativa). Plant Physiology and Biochemistry, 2024, 214: 108957. doi: 10.1016/j.plaphy.2024.108957
[14]	PIACENTINI D, RONZAN M, FATTORINI L, DELLA ROVERE F, MASSIMI L, ALTAMURA M M, FALASCA G. Nitric oxide alleviates cadmium- but not arsenic-induced damages in rice roots. Plant Physiology and Biochemistry, 2020, 151: 729-742. doi: S0981-9428(20)30173-X pmid: 32353678
[15]	吴坤, 邢承华, 饶玉春, 宋洪明, 陈晓阳, 蔡妙珍. 外源NO对铝毒下水稻根系生长和抗氧化系统的影响. 西北植物学报, 2014, 34(3): 536-542.
	WU K, XING C H, RAO Y C, SONG H M, CHEN X Y, CAI M Z. Effect of exogenous nitric oxide on root growth and antioxidant system in rice seedlings under aluminum toxicity. Journal of Northeast Agricultural University, 2014, 34(3): 536-542. (in Chinese)
[16]	方淑梅, 梁喜龙, 纪伟波, 林岩. 外源NO对盐碱胁迫下水稻幼苗生长抑制的缓解作用. 江苏农业科学, 2013, 41(8): 67-69.
	FANG S M, LIANG X L, JI W B, LIN Y. Alleviating effect of exogenous NO on growth inhibition of rice seedlings under saline-alkali stress. Jiangsu Agricultural Sciences, 2013, 41(8): 67-69. (in Chinese)
[17]	HSU Y T, LEE T M. Abscisic acid-dependent nitric oxide pathway and abscisic acid-independent nitric oxide routes differently modulate NaCl stress induction of the gene expression of methionine sulfoxide reductase A and B in rice roots. Journal of Plant Physiology, 2018, 231: 374-382. doi: S0176-1617(18)30403-6 pmid: 30388677
[18]	刘海艳, 杨丽洁, 丁艳锋, 李刚华, 王绍华, 刘正辉, 唐设, 刘仁梅, 蒋卫红. NO对水稻孕穗期干旱胁迫下叶片光合及产量的影响. 南京农业大学学报, 2017, 40(2): 195-202.
	LIU H Y, YANG L J, DING Y F, LI G H, WANG S H, LIU Z H, TANG S, LIU R M, JIANG W H. Effects of nitric oxide on photosynthesis and yield of rice under drought stress at booting stage. Journal of Northeast Agricultural University, 2017, 40(2): 195-202. (in Chinese)
[19]	闵炜芳, 方晶莹, 石亚飞, 摆小蓉, 舍杨梦斐, 田浩天, 罗成科. 硝普钠添加对碱胁迫下萌发期水稻碳氮代谢的影响. 植物营养与肥料学报, 2024, 30(5): 934-947.
	MIN W F, FANG J Y, SHI Y F, BAI X R, SHE Y, TIAN H T, LUO C K. Effect of sodium nitroprusside addition on carbon and nitrogen metabolism of rice during germination under alkali stress. Plant Nutrition and Fertilizer Science, 2024, 30(5): 934-947. (in Chinese)
[20]	石亚飞, 闵炜芳, 摆小蓉, 舍杨梦斐, 田浩天, 罗成科. 外源物调节碱胁迫水稻生理特性及相关基因表达的效应. 植物营养与肥料学报, 2023, 29(5): 813-825.
	SHI Y F, MIN W F, BAI X R, SHE Y, TIAN H T, LUO C K. Effects of exogenous regulatory substances on physiological characteristics and gene expression of rice seedlings under alkali stress. Journal of Plant Nutrition and Fertilizers, 2023, 29(5): 813-825. (in Chinese)
[21]	路旭平, 李芳兰, 石亚飞, 张娟伟, 杨文伟, 罗成科, 田蕾, 李培富. 不同水稻品种幼苗响应碱胁迫的生理差异及胁迫等级构建. 生态环境学报, 2021, 30(8): 1757-1768. doi: 10.16258/j.cnki.1674-5906.2021.08.023
	LU X P, LI F L, SHI Y F, ZHANG J W, YANG W W, LUO C K, TIAN L, LI P F. Physiological differences of seedlings of different rice varieties in response to alkali stress and construction of stress levels. Ecology and Environment Sciences, 2021, 30(8): 1757-1768. (in Chinese) doi: 10.16258/j.cnki.1674-5906.2021.08.023
[22]	李芳兰. 碱胁迫对中花11种子萌发、幼苗生长的影响及外源物的缓解作用[D]. 银川: 宁夏大学, 2021.
	LI F L. Effects of alkali stress on seed germination and seedling growth of Zhonghua 11 and mitigation effects of exogenous substance[D]. Yinchuan: Ningxia University, 2021. (in Chinese)
[23]	LI M, YUE T R, HAN J T, WANG J Q, XIAO H J, SHANG F D. Exogenous glucose irrigation alleviates cold stress by regulating soluble sugars, ABA and photosynthesis in melon seedlings. Plant Physiology and Biochemistry, 2024, 217: 109214. doi: 10.1016/j.plaphy.2024.109214
[24]	李合生. 植物生理生化实验原理和技术. 北京: 高等教育出版社, 2000.
	LI H S. Principles and Techniques of Plant Physiological Biochemical experiment. Beijing: Higher Education Press, 2000. (in Chinese)
[25]	LU X P, MIN W F, SHI Y F, TIAN L, LI P F, MA T L, ZHANG Y X, LUO C K. Exogenous melatonin alleviates alkaline stress by removing reactive oxygen species and promoting antioxidant defence in rice seedlings. Frontiers in Plant Science, 2022, 13: 849553. doi: 10.3389/fpls.2022.849553
[26]	LIVAK K J, SCHMITTGEN T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2 (-Delta Delta C(T)) Method. Methods, 2001, 25(4): 402-408. doi: 10.1006/meth.2001.1262
[27]	ZHANG H T, LI J J, YOO J H, YOO S C, CHO S H, KOH H J, SEO H S, PAEK N C. Rice Chlorina-1 and Chlorina-9 encode ChlD and ChlI subunits of Mg-chelatase, a key enzyme for chlorophyll synthesis and chloroplast development. Plant Molecular Biology, 2006, 62(3): 325-337. doi: 10.1007/s11103-006-9024-z pmid: 16915519
[28]	LEE S, KIM J H, YOO E S, LEE C H, HIROCHIKA H, AN G. Differential regulation of chlorophyll a oxygenase genes in rice. Plant Molecular Biology, 2005, 57(6): 805-818. pmid: 15952067
[29]	HE L, LI M, QIU Z N, CHEN D D, ZHANG G H, WANG X Q, CHEN G, HU J, GAO Z Y, DONG G J, et al. Primary leaf-type ferredoxin 1 participates in photosynthetic electron transport and carbon assimilation in rice. The Plant Journal, 2020, 104(1): 44-58. doi: 10.1111/tpj.v104.1
[30]	AZZOUZ-OLDEN F, HUNT A G, DINKINS R. Transcriptome analysis of drought-tolerant sorghum genotype SC56 in response to water stress reveals an oxidative stress defense strategy. Molecular Biology Reports, 2020, 47(5): 3291-3303. doi: 10.1007/s11033-020-05396-5
[31]	金姣姣, 刘自刚, 米文博, 徐明霞, 邹娅, 徐春梅, 赵彩霞. 利用RNA-Seq鉴定调控甘蓝型油菜叶片光合特性的低温胁迫应答基因. 生物技术通报, 2022, 38(4): 126-142. doi: 10.13560/j.cnki.biotech.bull.1985.2021-1048
	JIN J J, LIU Z G, MI W B, XU M X, ZOU Y, XU C M, ZHAO C X. Identification of low temperature stress-responsive genes regulating photosynthetic characteristics in the leaves of Brassica napus by RNA-seq. Biotechnology Bulletin, 2022, 38(4): 126-142. (in Chinese)
[32]	张荣, 刘淋茹, 付凯霞, 武紫君, 宋一凡, 王璐媛, 侯阁阁, 贺利, 冯伟, 段剑钊, 王永华, 郭天财. 干旱胁迫下外源褪黑素对冬小麦小花发育及碳营养代谢的调控. 中国农业科学, 2024, 57(23): 4644-4657. doi:10.3864/j.issn.0578-1752.2024.23.006.
	ZHANG R, LIU L R, FU K X, WU Z J, SONG Y F, WANG L Y, HOU G G, HE L, FENG W, DUAN J Z, WANG Y H, GUO T C. Regulatory of exogenous melatonin on floret development and carbon nutrient metabolism in winter wheat under drought stress. Scientia Agricultura Sinica, 2024, 57(23): 4644-4657. doi:10.3864/j.issn.0578-1752.2024.23.006. (in Chinese)
[33]	谷闻东, 刘春娟, 李邦, 刘畅, 周宇飞. 外源色氨酸对低氮胁迫下高粱苗期叶片碳氮平衡和衰老特性的影响. 中国农业科学, 2023, 56(7): 1295-1310. doi:10.3864/j.issn.0578-1752.2023.07.008.
	GU W D, LIU C J, LI B, LIU C, ZHOU Y F. Effects of exogenous tryptophan on C/N balance and senescence characteristics of sorghum seedlings under low nitrogen stress. Scientia Agricultura Sinica, 2023, 56(7): 1295-1310. doi:10.3864/j.issn.0578-1752.2023.07.008. (in Chinese)
[34]	FANG S M, HOU X, LIANG X L. Response mechanisms of plants under saline-alkali stress. Frontiers in Plant Science, 2021, 12: 667458. doi: 10.3389/fpls.2021.667458
[35]	DU J G, LI W L, WANG Z, CHEN Z H, WANG C, LU W, XIONG A S, TAN G F, ZHENG Y X, LI M Y. Effects of exogenous melatonin on drought stress in celery (Apium graveolens L.): Unraveling the modulation of chlorophyll and glucose metabolism pathways. BMC Genomics, 2024, 25(1): 1104. doi: 10.1186/s12864-024-11054-y
[36]	DU X L, FENG N J, ZHENG D F, LIN Y, ZHOU H, LI J H, YANG X H, HUO J X, MEI W Q. Effects of exogenous Uniconazole (S3307) on oxidative damage and carbon metabolism of rice under salt stress. BMC Plant Biology, 2025, 25(1): 541. doi: 10.1186/s12870-025-06467-0
[37]	李俊伟, 刘景辉, 王俊英, 郭来春, 王春龙, 任长忠. 盐与碱胁迫对燕麦离子平衡和有机酸含量的影响. 西北植物学报, 2022, 42(10): 1700-1710.
	LI J W, LIU J H, WANG J Y, GUO L C, WANG C L, REN C Z. Effect of salt and alkali stress on oat ion balance and organic acid content. Acta Botanica Boreali-Occidentalia Sinica, 2022, 42(10): 1700-1710. (in Chinese)
[38]	刘建新, 刘瑞瑞, 刘秀丽, 贾海燕, 卜婷, 李娜. 外源硫化氢调控盐碱胁迫下裸燕麦叶片糖和酚酸代谢反应. 中国生态农业学报(中英文), 2023, 31(3): 463-477.
	LIU J X, LIU R R, LIU X L, JIA H Y, BU T, LI N. Exogenous hydrogen sulfide modulates metabolic responses of sugar and phenolic acid in naked oat leaves under saline-alkali stress. Chinese Journal of Eco-Agriculture, 2023, 31(3): 463-477. (in Chinese)
[39]	LEMOINE R, LA CAMERA S, ATANASSOVA R, DÉDALDÉCHAMP F, ALLARIO T, POURTAU N, BONNEMAIN J L, LALOI M, COUTOS-THÉVENOT P, MAUROUSSET L, et al. Source-to-sink transport of sugar and regulation by environmental factors. Frontiers in Plant Science, 2013, 4: 272. doi: 10.3389/fpls.2013.00272 pmid: 23898339
[40]	SCOFIELD G N, HIROSE T, AOKI N, FURBANK R T. Involvement of the sucrose transporter, OsSUT1, in the long-distance pathway for assimilate transport in rice. Journal of Experimental Botany, 2007, 58(12): 3155-3169. pmid: 17728297
[41]	马福林, 马玉花. 干旱胁迫对植物的影响及植物的响应机制. 宁夏大学学报(自然科学版), 2022, 43(4): 391-399.
	MA F L, MA Y H. Effect of drought stress on plants and their response mechanism. Journal of Ningxia University (Natural Science Edition), 2022, 43(4): 391-399. (in Chinese)
[42]	CHEN Q C, HU T, LI X H, SONG C P, ZHU J K, CHEN L Q, ZHAO Y. Phosphorylation of SWEET sucrose transporters regulates plant root: Shoot ratio under drought. Nature Plants, 2022, 8(1): 68-77. doi: 10.1038/s41477-021-01040-7
[43]	MATHAN J, SINGH A, RANJAN A. Sucrose transport in response to drought and salt stress involves ABA-mediated induction of OsSWEET13 and OsSWEET15 in rice. Physiologia Plantarum, 2021, 171(4): 620-637. doi: 10.1111/ppl.v171.4
[44]	DINLER B S, ANTONIOU C, FOTOPOULOS V. Interplay between GST and nitric oxide in the early response of soybean (Glycine max L.) plants to salinity stress. Journal of Plant Physiology, 2014, 171(18): 1740-1747. doi: 10.1016/j.jplph.2014.07.026
[45]	ZHANG A Y, ZHANG J, ZHANG J H, YE N H, ZHANG H, TAN M P, JIANG M Y. Nitric oxide mediates brassinosteroid-induced ABA biosynthesis involved in oxidative stress tolerance in maize leaves. Plant & Cell Physiology, 2011, 52(1): 181-192.
[46]	SANG J R, ZHANG A Y, LIN F, TAN M P, JIANG M Y. Cross-talk between calcium-calmodulin and nitric oxide in abscisic acid signaling in leaves of maize plants. Cell Research, 2008, 18(5): 577-588. doi: 10.1038/cr.2008.39 pmid: 18364679
[47]	LIANG Y F, ZHAO J Y, YANG R, BAI J Y, HU W X, GU L X, LIAN Z Y, HUO H Q, GUO J, GONG H J. PROCERA interacts with JACKDAW in gibberellin-enhanced source-sink sucrose partitioning in tomato. Plant Physiology, 2024, 197(1): kiaf024.
[48]	FRESCHI L. Nitric oxide and phytohormone interactions: Current status and perspectives. Frontiers in Plant Science, 2013, 4: 398. doi: 10.3389/fpls.2013.00398 pmid: 24130567
[49]	CHEN L C, SUN S H, SONG C P, ZHOU J M, LI J Y, ZUO J R. Nitric oxide negatively regulates gibberellin signaling to coordinate growth and salt tolerance in Arabidopsis. Journal of Genetics and Genomics, 2022, 49(8): 756-765.

基因名称 Gene symbol	RGAP ID RGAP ID	引物 Primer sequence (5′—3′) (forward/reverse)	产物长度 Product size (bp)
OsP5CS	LOC_Os05g38150	AATGACAGTTTAGCAGGAC/ACCACTATACAACCCATCC	87
OsNIN1	LOC_Os03g20020	CTATTCTGCTTTGCGTTGT/CTGAGCCTGTAGTTGATGG	87
OsSUS5	LOC_Os04g24430	GCATTCGGGCTAACGGTCAT/ATTCCTGCCTCCCTGTCGT	140
OsSPS1	LOC_Os01g69030	GGCACAGCAAGACACTCCC/CGCCACGAACTAGACCATG	134
OsYGL1	LOC_Os05g28200	CTGTTGGTGGGTCCTTGCTT/TAGCTCGCACCAAGAGCAAA	97
OsCAO1	LOC_Os10g41780	TGCTCATCAAGCCTTCCTTCAGGTG/GCGTTGTTCTTTGCATAGTCAGCTC	115
OsCAO2	LOC_Os10g41760	CTGCTCGTCAAGCCTTCCTCCTCCT/CCCTCATGTTGTTCTTGGCATGTTG	130
OsFd1	LOC_Os08g01380	CAGGCGGAGGAGGAAGGGAT/TAGGCGTGGCAGGTGAGGAC	161
OsChlH	LOC_Os03g20700	CCAATCCGTAACCCGAAGGT/CAATAATTTTGGCGCTCTTCAA	103
OsNCED2	LOC_Os12g24800	TCCGCAATGGTCCCAACC/ACAGTGTTGACGAGGCCG	111
OsABA8ox2	LOC_Os08g36860	TGGCCAAGCTGGAGATGC/ACGGGCTGTACTCCACCT	96
OsGA3ox2	LOC_Os01g08220	TGTTCTTGAGGGCGCTGG/GTCGCCGTCATCCTCTCG	85
OsGA20ox1	LOC_Os03g63970	GGTGCATGGACGCCTTCT/CTTCCACGGCAGCTTGGA	125