水杨酸引发提高低温下水稻种子萌发活力的生理与分子效应

doi:10.3864/j.issn.0578-1752.2024.07.002

摘要/Abstract

摘要：

【目的】研究水杨酸（SA）引发对低温下水稻种子萌发活力的影响及生理响应，揭示SA引发对脱落酸（ABA）和赤霉素（GA）代谢途径相关基因以及细胞壁松弛基因的诱导模式，为水稻种子低温萌发研究提供理论依据。【方法】以籼型三系杂交水稻泰丰优208种子为材料，通过种子引发处理，分析SA对种子低温萌发活力及生理的影响，并通过qRT-PCR技术分析ABA、GA和扩展蛋白基因响应SA引发的表达模式。【结果】低温（15 ℃）显著推迟水稻种子萌发进程。在低温下萌发1 d种子中，其内源SA浓度是常温（28 ℃）下的1.7倍；但对于5 d的幼苗而言，低温下的SA浓度仅为常温下浓度的0.6%。SA引发可有效提高种子在低温下的萌发活力，尤其以2 000 μmol·L^-1 SA效果最为显著，该浓度显著提高了低温下种子的发芽指数、活力指数、芽长、根长、鲜重和干重，其中活力指数分别为未引发种子（CK1）和水引发种子（CK2）的3和2倍。在生理指标方面，SA引发提高了低温萌发过程中种子的可溶性糖、脯氨酸以及活性氧含量，增加了总淀粉酶、β-淀粉酶、超氧化物歧化酶（SOD）和过氧化氢酶（CAT）活性，降低了丙二醛（MDA）含量。与CK1相比，2 000 μmol·L^-1 SA引发将种子ABA含量降低了79%，同时将IAA和GA₁含量增加了32.2%和2.66倍。在基因表达方面，对于2 000 μmol·L^-1 SA引发的种子，ABA合成基因OsNCED2和OsNCED3的表达量分别比CK1降低了94.26%和90.24%；而ABA分解基因OsABA8’ox2和OsABA8’ox3的表达量分别为CK1的5.9和3.9倍。与CK1相比，SA引发显著上调了GA合成基因OsCPS1、OsKAO和OsGA20ox1的表达量，并显著下调GA分解基因OsGA2ox2和OsGA2ox6的表达量。在几个候选的细胞壁松弛因子扩展蛋白基因中，除了OsEXPB11外，其余几个同源基因均在一定程度上被引发而上调表达。与CK1相比，2 000 μmol·L^-1 SA引发分别使OsEXPA2、OsEXPB4和OsEXPB6的表达量上调12.2、5.9和6.1倍。【结论】SA引发可显著缓解低温对于水稻种子萌发和幼苗生长的影响。可能是由于SA提高SOD、CAT等抗氧化酶活性，降低MDA的产生，增加可溶性糖和脯氨酸的含量，进而增强种子和幼苗对于低温的耐受能力。另一方面，SA引发通过降低种子内源ABA含量，增加GA₁含量，增强总淀粉酶和β-淀粉酶活性，促进细胞壁松弛相关基因的表达，从而促进低温下的种子萌发和幼苗生长。

关键词: 水稻, 低温胁迫, 种子引发, 水杨酸, 生理指标, 基因表达

Abstract:

【Objective】The study investigated the impact of salicylic acid (SA) priming on the germination vigor and physiological response of rice seeds under low temperatures. It aimed to reveal the expression patterns of genes related to abscisic acid (ABA) and gibberellin (GA) metabolic pathways as well as cell wall relaxation genes by SA priming. This research provided a theoretical basis for the study of rice seed germination at low temperatures.【Method】Using indica three-line hybrid rice Taifengyou 208 seeds as materials, the effects of SA on seed germination vigor and physiology responses under low temperature were analyzed through seed priming treatment, and the expression patterns of genes related to ABA, GA and expansin in response to SA were analyzed by qRT-PCR.【Result】Low temperature (15 ℃) significantly delayed the germination process of rice seeds. In seeds germinated at low temperatures for one day, the endogenous SA concentration was 1.7 times higher than that at normal temperatures (28 ℃). However, for five-day-old seedlings, the SA concentration under low temperature was only 0.6% of that at normal temperatures. SA could effectively enhanced germination vigor of seeds at low temperature, with the most significant effects observed at 2 000 μmol·L^-1 SA. This concentration significantly increased the germination index, vigor index, shoot length, root length, fresh weight, and dry weight of seeds under low temperature conditions. Notably, the vigor index was three times that of non-primed seeds (CK1) and two times that of water-primed seeds (CK2). In terms of physiological indexes, SA priming increased the contents of soluble sugar, proline and active oxygen, enhanced the activities of total amylase, β-amylase, superoxide dismutase (SOD) and catalase (CAT), and decreased the content of malondialdehyde (MDA). Compared with CK1, 2 000 μmol·L^-1 SA decreased the ABA content by 79%, and increased the IAA and GA₁ contents by 32.2% and 2.66 times, respectively. In terms of gene expression, the expression levels of ABA synthesizing genes OsNCED2 and OsNCED3 were decreased by 94.26% and 90.24% compared with CK1 in seeds primed by 2 000 μmol·L^-1 SA, respectively, whereas the expression levels of ABA decomposing genes OsABA8’ox2 and OsABA8’ox3 were 5.9 and 3.9 times higher than that of CK1, respectively. Compared with CK1, SA priming significantly upregulated the expression of GA synthesizing genes OsCPS1, OsKAO and OsGA20ox1, while it significantly downregulated the expression of GA decomposing genes OsGA2ox2 and OsGA2ox6. In several candidate genes encoding cell wall relaxation protein, e.t. expansin, all but OsEXPB11 were significantly upregulated to some extent by priming. Compared with CK1, 2 000 μmol·L^-1 SA increased the expression levels of OsEXPA2, OsEXPB4 and OsEXPB6 to 12.2, 5.9 and 6.1 times, respectively.【Conclusion】SA priming can significantly alleviate the impact of low temperatures on rice seed germination and seedling growth, which is likely due to SA enhancing the activity of antioxidant enzymes such as SOD and CAT, reducing the production of MDA, and increasing the content of soluble sugars and proline, thereby strengthening the tolerance of seeds and seedlings to low temperatures. On the other hand, SA priming decreases endogenous ABA content, increases GA₁ content, enhances the activities of total amylase and β-amylase, and promotes the expression of genes related to cell wall relaxation, thus facilitating seed germination and seedling growth at low temperature.

Key words: rice, low temperature stress, seed priming, salicylic acid (SA), physiological index, gene expression

陈兵先, 张琪, 戴彰言, 周旭, 刘军. 水杨酸引发提高低温下水稻种子萌发活力的生理与分子效应[J]. 中国农业科学, 2024, 57(7): 1220-1236.

CHEN BingXian, ZHANG Qi, DAI ZhangYan, ZHOU Xu, LIU Jun. Physiological and Molecular Effects of Salicylic Acid on Rice Seed Germination at Low Temperature[J]. Scientia Agricultura Sinica, 2024, 57(7): 1220-1236.

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参考文献 52

[1]	MARTHANDAN V, GEETHA R, KUMUTHA K, RENGANATHAN V G, KARTHIKEYAN A, RAMALINGAM J. Seed priming: A feasible strategy to enhance drought tolerance in crop plants. International Journal of Molecular Sciences, 2020, 21(21): 8258.
[2]	王慰亲. 种子引发促进直播早稻低温胁迫下萌发出苗的机理研究[D]. 武汉: 华中农业大学, 2019.
	WANG W Q. Mechanisms underlying the effects of seed priming on the establishment of direct-seeded early season rice under chilling stress[D]. Wuhan: Huazhong Agricultural University, 2019. (in Chinese)
[3]	胡亚丽, 聂靖芝, 吴霞, 潘姣, 曹珊, 岳娇, 罗登杰, 王财金, 李增强, 张辉, 吴启境, 陈鹏. 水杨酸引发对红麻幼苗耐盐性的影响. 中国农业科学, 2022, 55(14): 2696-2708. doi: 10.3864/j.issn.0578-1752.2022.14.002.
	HU Y L, NIE J Z, WU X, PAN J, CAO S, YUE J, LUO D J, WANG C J, LI Z Q, ZHANG H, WU Q J, CHEN P. Effect of salicylic acid priming on salt tolerance of kenaf seedlings. Scientia Agricultura Sinica, 2022, 55(14): 2696-2708. doi: 10.3864/j.issn.0578-1752.2022.14.002. (in Chinese)
[4]	YANG Z, ZHI P, CHANG C. Priming seeds for the future: Plant immune memory and application in crop protection. Frontiers in Plant Science, 2022, 13: 961840. doi: 10.3389/fpls.2022.961840
[5]	NIE L, SONG S, YIN Q, ZHAO T, LIU H, HE A, WANG W. Enhancement in seed priming-induced starch degradation of rice seed under chilling stress via GA-mediated α-amylase expression. Rice, 2022, 15(1): 19.
[6]	TANOU G, FOTOPOULOS V, MOLASSIOTIS A. Priming against environmental challenges and proteomics in plants: Update and agricultural perspectives. Frontiers in Plant Science, 2012, 3: 216. doi: 10.3389/fpls.2012.00216 pmid: 22973291
[7]	MIURA K, TADA Y. Regulation of water, salinity, and cold stress responses by salicylic acid. Frontiers in Plant Science, 2014, 5: 4. doi: 10.3389/fpls.2014.00004 pmid: 24478784
[8]	王立红, 李星星, 孙影影, 阿曼古丽·买买提阿力, 张巨松. 外源水杨酸对NaCl胁迫下棉花幼苗生长生理特性的影响. 西北植物学报, 2017, 37(1): 154-162.
	WANG L H, LI X X, SUN Y Y, MAIMAITIALI A M G L, ZHANG J S. Effects of exogenous salicylic acid on the physiological characteristics and growth of cotton seedlings under NaCl stress. Acta Botanica Boreali-Occidentalia Sinica, 2017, 37(1): 154-162. (in Chinese)
[9]	ALAM M, HAYAT K, ULLAH I, SAJID M, AHMAD M, BASIT A, AHMAD I, MUHAMMAD A, AKBAR S, HUSSAIN Z. Improving okra (Abelmoschus esculentus L.) growth and yield by mitigating drought through exogenous application of salicylic acid. Fresenius Environmental Bulletin, 2020, 29(1): 529-535.
[10]	AMJAD M, ZIAF K, IQBAL Q, AHMAD I, RIAZ M A, SAQIB Z A. Effect of seed priming on seed vigour and salt tolerance in hot pepper. Pakistan Journal of Agricultural Sciences, 2007, 44(3): 408-416.
[11]	LI Z, XU J, GAO Y, WANG C, GUO G, LUO Y, HUANG Y, HU W, SHETEIWY M S, GUAN Y, HU J. The synergistic priming effect of exogenous salicylic acid and H₂O₂ on chilling tolerance enhancement during maize (Zea mays L.) seed germination. Frontiers in Plant Science, 2017, 8: 1153. doi: 10.3389/fpls.2017.01153
[12]	侯林欣, 吕强, 黄明, 焦念元, 尹飞, 刘领, 吕梦, 付国占. 不同温度水杨酸引发对干旱胁迫下玉米种子发芽及幼苗生理特性的影响. 中国农学通报, 2021, 37(19): 13-21. doi: 10.11924/j.issn.1000-6850.casb2020-0504
	HOU L X, LÜ Q, HUANG M, JIAO N Y, YIN F, LIU L, LÜ M, FU G Z. SA priming of maize seeds at different temperatures under drought stress: Effects on seed germination and seedling physiological characteristics. Chinese Agricultural Science Bulletin, 2021, 37(19): 13-21. (in Chinese)
[13]	常云霞, 徐克东, 陈璨, 陈龙. 水杨酸对低温胁迫下大豆幼苗生长抑制的缓解效应. 大豆科学, 2012, 31(6): 927-931.
	CHANG Y X, XU K D, CHEN C, CHEN L. Salicylic acid mitigating the inhibition of low temperature stress to soybean seedlings. Soybean Science, 2012, 31(6): 927-931. (in Chinese)
[14]	NAZARI R, PARSA S, TAVAKKOL AFSHARI R, MAHMOODI S, SEYYEDI S M. Salicylic acid priming before and after accelerated aging process increases seedling vigor in aged soybean seed. Journal of Crop Improvement, 2020, 34(2): 218-237. doi: 10.1080/15427528.2019.1710734
[15]	HOSSEINIFARD M, STEFANIAK S, GHORBANI JAVID M, SOLTANI E, WOJTYLA Ł, GARNCZARSKA M. Contribution of exogenous proline to abiotic stresses tolerance in plants: A review. International Journal of Molecular Sciences, 2022, 23(9): 5186.
[16]	倪万潮, 束红梅, 郭书巧, 蒋璐, 何晓兰, 崔晓霞, 巩元勇. 不同水稻品种种子萌发生理特性差异研究. 中国农学通报, 2020, 36(2): 1-5. doi: 10.11924/j.issn.1000-6850.casb18080094
	NI W C, SHU H M, GUO S Q, JIANG L, HE X L, CUI X X, GONG Y Y. Seed germination of rice cultivars: Differences in physiological characteristics. Chinese Agricultural Science Bulletin, 2020, 36(2): 1-5. (in Chinese) doi: 10.11924/j.issn.1000-6850.casb18080094
[17]	CUI D, YIN Y, WANG J, WANG Z, DING H, MA R, JIAO Z. Research on the physio-biochemical mechanism of non-thermal plasma-regulated seed germination and early seedling development in Arabidopsis. Frontiers in Plant Science, 2019, 10: 1322. doi: 10.3389/fpls.2019.01322
[18]	ZHANG Y, CHEN B, XU Z, SHI Z, CHEN S, HUANG X, CHEN J, WANG X. Involvement of reactive oxygen species in endosperm cap weakening and embryo elongation growth during lettuce seed germination. Journal of Experimental Botany, 2014, 65(12): 3189-3200. doi: 10.1093/jxb/eru167 pmid: 24744430
[19]	HOLLOWAY T, STEINBRECHER T, PÉREZ M, SEVILLE A, STOCK D, NAKABAYASHI K, LEUBNER-METZGER G. Coleorhiza- enforced seed dormancy: A novel mechanism to control germination in grasses. New Phytologist, 2021, 229(4): 2179-2191. doi: 10.1111/nph.v229.4
[20]	YE N, ZHU G, LIU Y, ZHANG A, LI Y, LIU R, SHI L, JIA L, ZHANG J. Ascorbic acid and reactive oxygen species are involved in the inhibition of seed germination by abscisic acid in rice seeds. Journal of Experimental Botany, 2012, 63(5): 1809-1822. doi: 10.1093/jxb/err336 pmid: 22200664
[21]	LIU L, XIA W, LI H, ZENG H, WEI B, HAN S, YIN C. Salinity inhibits rice seed germination by reducing α-amylase activity via decreased bioactive gibberellin content. Frontiers in Plant Science, 2018, 9: 275. doi: 10.3389/fpls.2018.00275
[22]	CHEN B X, LI W Y, GAO Y T, CHEN Z J, ZHANG W N, LIU Q J, CHEN Z, LIU J. Involvement of polyamine oxidase-produced hydrogen peroxide during coleorhiza-limited germination of rice seeds. Frontiers in Plant Science, 2016, 7: 1219. doi: 10.3389/fpls.2016.01219
[23]	XIONG Q, MA B, LU X, HUANG Y H, HE S J, YANG C, YIN C C, ZHAO H, ZHOU Y, ZHANG W K, WANG W S, LI Z K, CHEN S Y, ZHANG J S. Ethylene-inhibited jasmonic acid biosynthesis promotes mesocotyl/coleoptile elongation of etiolated rice seedlings. The Plant Cell, 2017, 29(5): 1053-1072. doi: 10.1105/tpc.16.00981 pmid: 28465411
[24]	HUSSAIN S, KHALIQ A, ALI B, HUSSAIN H A, QADIR T, HUSSAIN S. Temperature extremes:Impact on rice growth and development// Plant Abiotic Stress Tolerance: Agronomic, Molecular and Biotechnological Approaches. Springer, 2019: 153-171.
[25]	BHANUPRAKASH K, YOGEESHA H S. Seed priming for abiotic stress tolerance: An overview// Abiotic Stress Physiology of Horticultural Crops. Springer, 2016: 103-117.
[26]	KIM S H, JANG D C, LEE J J, HEO J Y. Salicylic acid seed priming boosts germination in Brassica rapa ssp pekinensis under cold stress. Journal of Applied Horticulture, 2021, 23(3): 286-289. doi: 10.37855/jah.2021.v23i03.50
[27]	YACOUBI R, JOB C, BELGHAZI M, CHAIBI W, JOB D. Toward characterizing seed vigor in alfalfa through proteomic analysis of germination and priming. Journal of Proteome Research, 2011, 10(9): 3891-3903. doi: 10.1021/pr101274f pmid: 21755932
[28]	LIU H, ABLE A J, ABLE J A. Priming crops for the future: Rewiring stress memory. Trends in Plant Science, 2022, 27(7): 699-716. doi: 10.1016/j.tplants.2021.11.015
[29]	CHEN K, ARORA R. Priming memory invokes seed stress-tolerance. Environmental and Experimental Botany, 2013, 94: 33-45. doi: 10.1016/j.envexpbot.2012.03.005
[30]	DIAN R, LI S, SUN B, GUO A. Recent advances and new guidelines on hyperspectral and multispectral image fusion. Information Fusion, 2021, 69: 40-51. doi: 10.1016/j.inffus.2020.11.001
[31]	ELMASRY G, MANDOUR N, AL-REJAIE S, BELIN E, ROUSSEAU D. Recent applications of multispectral imaging in seed phenotyping and quality monitoring-An overview. Sensors, 2019, 19(5): 1090.
[32]	JISHA K C, VIJAYAKUMARI K, PUTHUR J T. Seed priming for abiotic stress tolerance: An overview. Acta Physiologiae Plantarum, 2013, 35: 1381-1396. doi: 10.1007/s11738-012-1186-5
[33]	ANJU U L, DODDAGOUDAR S R, PATTANASHETTI S K, BASAVE G, VIJAYKUMAR K. Influence of seed priming on seed germination, seedling growth, peroxidase activity, proline and total soluble sugar content of pearl millet (Pennisetum glaucum L.) under salinity stress. International Journal of Chemistry Studies, 2019, 7(5): 508-514.
[34]	周丽霞, 曹红星, 肖勇. 外源水杨酸对低温胁迫椰子幼苗生理特性的影响. 南方农业学报, 2017, 48(11): 2039-2045.
	ZHOU L X, CAO H X, XIAO Y. Effects of exogenous salicylic acid on physiological characteristics of Cocos nucifera L. young seedlings under cold stress. Journal of Southern Agriculture, 2017, 48(11): 2039-2045. (in Chinese)
[35]	李媛, 李武, 莫钊文, 闻祥成, 王抄抄, 徐刚红, 李妹娟, 聂俊, 唐湘如. 水杨酸和盐浸种对香稻和非香稻幼苗生理特性的影响. 华北农学报, 2014, 29(5): 168-174. doi: 10.7668/hbnxb.2014.05.029
	LI Y, LI W, MO Z W, WEN X C, WANG C C, XU G H, LI M J, NIE J, TANG X R. Effects of pre-soaking with salicylic acid and salt on some physiological characteristics of the aromatic and non-aromatic rice seedlings. Acta Agriculturae Boreali-Sinica, 2014, 29(5): 168-174. (in Chinese) doi: 10.7668/hbnxb.2014.05.029
[36]	MITTLER R, ZANDALINAS S I, FICHMAN Y, VAN BREUSEGEM F. Reactive oxygen species signalling in plant stress responses. Nature Reviews Molecular Cell Biology, 2022, 23(10): 663-679. doi: 10.1038/s41580-022-00499-2
[37]	MIRANSARI M, SMITH D L. Plant hormones and seed germination. Environmental and Experimental Botany, 2014, 99: 110-121. doi: 10.1016/j.envexpbot.2013.11.005
[38]	杨楠, 曹亚从, 魏兵强, 王立浩. 单双子叶植物种子萌发和休眠的研究进展. 植物遗传资源学报, 2022, 23(5): 1249-1257. doi: 10.13430/j.cnki.jpgr.20220314001
	YANG N, CAO Y C, WEI B Q, WANG L H. Research progress on seed germination and dormancy of monocot and dicot plants. Journal of Plant Genetic Resources, 2022, 23(5): 1249-1257. (in Chinese)
[39]	EL-SHERIF N A. Salicylic acid and its crosstalk with other plant hormones under stressful environments//Managing Plant Stress Using Salicylic Acid: Physiological and Molecular Aspects. Wiley, 2022: 304-317.
[40]	帅海威, 孟永杰, 罗晓峰, 陈锋, 戚颖, 杨文钰, 舒凯. 生长素调控种子的休眠与萌发. 遗传, 2016, 38(4): 314-322.
	SHUAI H W, MENG Y J, LUO X F, CHEN F, QI Y, YANG W Y, SHU K. The roles of auxin in seed dormancy and germination. Hereditas, 2016, 38(4): 314-322. (in Chinese) doi: 10.1111/j.1601-5223.1952.tb02928.x
[41]	TENG Z, YU H, WANG G, MENG S, LIU B, YI Y, CHEN Y, ZHENG Q, LIU L, YANG J, DUAN M, ZHANG J, YE N. Synergistic interaction between ABA and IAA due to moderate soil drying promotes grain filling of inferior spikelets in rice. The Plant Journal, 2022, 109(6): 1457-1472. doi: 10.1111/tpj.v109.6
[42]	PARWEZ R, AFTAB T, GILL S S, NAEEM M. Abscisic acid signaling and crosstalk with phytohormones in regulation of environmental stress responses. Environmental and Experimental Botany, 2022, 199: 104885. doi: 10.1016/j.envexpbot.2022.104885
[43]	ZHU G, YE N, ZHANG J. Glucose-induced delay of seed germination in rice is mediated by the suppression of ABA catabolism rather than an enhancement of ABA biosynthesis. Plant and Cell Physiology, 2009, 50: 644-651. doi: 10.1093/pcp/pcp022 pmid: 19208695
[44]	CHEN B X, PENG Y X, GAO J D, ZHANG Q, LIU Q J, FU H, LIU J. Coumarin-induced delay of rice seed germination is mediated by suppression of abscisic acid catabolism and reactive oxygen species production. Frontiers in Plant Science, 2019, 10: 828. doi: 10.3389/fpls.2019.00828
[45]	徐振江, 陈兵先, 赵晟楠, 袁红霞, 郜珊珊, 张瑜, 陈靖, 王晓峰. 种子萌发过程中胚乳的突破性研究. 植物生理学报, 2012, 48(9): 853-863.
	XU Z J, CHEN B X, ZHAO S N, YUAN H X, GAO S S, ZHANG Y, CHEN J, WANG X F. Breaking through the endosperm during seed germination. Plant Physiology Journal, 2012, 48(9): 853-863. (in Chinese)
[46]	STEINBRECHER T, LEUBNER-METZGER G. The biomechanics of seed germination. Journal of Experimental Botany, 2017, 68(4): 765-783. doi: 10.1093/jxb/erw428 pmid: 27927995
[47]	CHEN B, MA J, XU Z, WANG X. Abscisic acid and ethephon regulation of cellulase in the endosperm cap and radicle during lettuce seed germination. Journal of Integrative Plant Biology, 2016, 58(10): 859-869. doi: 10.1111/jipb.12479
[48]	XU Z, YANG M, LI Z, XIAO J, YANG X, WANG H, WANG X. Tissue-specific pectin methylesterification and pectin methylesterase activities play a role in lettuce seed germination. Scientia Horticulturae, 2022, 301: 111134. doi: 10.1016/j.scienta.2022.111134
[49]	LIU C, LI L, CHEN B, WANG X. Suppression of α-l- arabinofuranosidase in the endosperm and atypical germination of lettuce seeds induced by sodium dichloroisocyanurate. Acta Physiologiae Plantarum, 2015, 37: 10. doi: 10.1007/s11738-014-1761-z
[50]	GONZÁLEZ-CALLE V, BARRERO-SICILIA C, CARBONERO P, IGLESIAS-FERNÁNDEZ R. Mannans and endo-β-mannanases (MAN) in Brachypodium distachyon: Expression profiling and possible role of the BdMAN genes during coleorhiza-limited seed germination. Journal of Experimental Botany, 2015, 66(13): 3753-3764. doi: 10.1093/jxb/erv168
[51]	COSGROVE D J. Loosening of plant cell walls by expansins. Nature, 2000, 407: 321-326. doi: 10.1038/35030000
[52]	CHEN F, BRADFORD K J. Expression of an expansin is associated with endosperm weakening during tomato seed germination. Plant Physiology, 2000, 124(3): 1265-1274. doi: 10.1104/pp.124.3.1265 pmid: 11080302

基因名称 Gene name	基因登录号 Gene ID	正向引物 Forward primer (5′-3′)	反向引物 Reverse primer (5′-3′)
OsNCED1	LOC_Os02g47510	GAACTTCGACTTCCCCGTGA	ACGTAGCACACCAAGTACCC
OsNCED2	LOC_Os12g24800	GAGCCTTGAGTTCGGTGTCA	CAGCAAAGCACCCTAGACCA
OsNCED3	LOC_Os03g44380	ATATGGCGACGATCACGACG	CGCGGAGAATCTCACCGAAT
OsNCED4	LOC_Os07g05940	TCGGGAGGTACGACTTCCAT	TTGAGGTACGGCTTGGACAC
OsNCED5	LOC_Os12g42280	CGAGCTCACCAAGTTCGAGT	TTGATGAAGGTGCCGTGGAA
OsABA8’ox1	LOC_Os02g47470	AAAACCAACATCAACGGCGG	GATTGCCAACCGTGGTCCTA
OsABA8’ox2	LOC_Os08g36860	CGTGTGTGTGGATGCAATGG	AGTGCTACACTAGGCACACC
OsABA8’ox3	LOC_Os09g28390	CTGGTCACTGGCTACAGGTG	TGCTACGCCATTGTCGTCAT
OsCPS1	LOC_Os02g17780	TCAAGAGACACCGCCAGTTC	ACAGTGCATGACCCTGGATG
OsKAO	LOC_Os06g 02019	CTCCTTCGTGTCCTTCCGTC	ACCACAGCTGAACCTTCCAC
OsGA20ox1	LOC_Os03g63970	TTCTTCCTCTGCCCGGAGAT	CATGTCGGCCCTGTAGTGG
OsGA3ox2	LOC_ Os01g08220	CTTCTGTGACGTGATGGAGGAG	CTCAAGAACAACCTCAGCAACTC
OsGA2ox1	LOC_Os05g06670	ACCCGCAGATCCTTAGCTTG	TGAACCCACATCTCCTTGCC
OsGA2ox2	LOC_Os01g22910	TGTTTGGTTGAGGGGTGAGT	CATTTTCCCCGATCAGTTGGT
OsGA2ox3	LOC_Os01g55240	ACTCGTTGCAGGTTCTGACC	GTGGCAATGGTGCAATCCTC
OsGA2ox6	LOC_Os04g44150	GGCTATCGGTGGCCTACTTC	TTGTCCTGACGTCTTCCTGC
OsEXPA2	LOC_Os01g60770	GCGGCCAGTTCTGATCGAGTA	GCAGCCTCAGAATAGCCAAAGC
OsEXPA4	LOC_Os05g39990	CCGTCTCCGACACCCACATAT	TGGACGAAGTCCAGAGAAGGAA
OsEXPB4	LOC_Os10g40730	CCCAACACATTCTACCGCTCCT	ACAGACCGACCACACAATCCC
OsEXPB6	LOC_Os10g40700	AATTTGCGTGGGATTGAGGTGT	TGGGTAGTACAGTGACAGTGGG
OsEXPB11	LOC_ Os02g44108	TGCAGTGCAGAGTTGCGGTAA	CAGAGACCGTGGAGGGAAGAAC
OsEXPLA1	LOC_Os03g04020	ACACGCACGAGTGGAAGTAGAA	TGCCGAGGGATTAGGAGGACT
OsGAPDH1	LOC_Os02g38920	GCAATCAAGGAGGAGGCTGA	ACGTGTCGCTCAAAGCAATG

引发方式 Priming method	发芽势 Germination potential (%)	发芽率 Germination rate (%)	发芽指数 Germination index	活力指数 Vigor index	芽长 Seedling length (cm)	根长 Root length (cm)	鲜重 Fresh weight (g)	干重 Dry weigh (g)
CK1	43.75±3.11e	84.75±3.30d	15.26±2.84d	19.70±2.84e	1.13±0.29d	1.20±0.21d	0.44±0.01d	0.40±0.01bc
CK2	52.25±3.56d	84.25±3.20d	16.81±2.21cd	30.25±3.21d	1.21±0.07d	1.77±0.12c	0.47±0.01c	0.38±0.02c
50 μmol·L^-1 SA	55.00±3.92d	85.50±1.29d	16.17±1.90d	31.60±3.90d	1.47±0.21c	1.83±0.22c	0.43±0.02d	0.39±0.01c
500 μmol·L^-1 SA	90.75±4.27a	90.75±2.75c	17.32±1.98bc	36.37±4.98c	1.49±0.18c	2.10±0.22b	0.54±0.02b	0.42±0.02b
2000 μmol·L^-1 SA	83.25±3.30b	96.50±1.73a	21.59±2.67a	60.20±4.67a	1.65±0.05a	2.76±0.14a	0.63±0.02a	0.55±0.04a
5000 μmol·L^-1 SA	63.50±3.30c	92.50±1.29b	18.36±1.85b	48.40±3.05b	1.57±0.19b	2.69±0.24a	0.61±0.03a	0.54±0.02a