Tolerance of Eruca sativa to Stress by Four Types of Salt
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摘要:
目的 测定芸芥品系在单盐处理下萌发指标的变化,分析芸芥的耐盐极限值和临界值,明确芸芥对4种单盐胁迫耐受性的差异。 方法 以15芸芥4、11芸芥32-1和16芸芥9等3个品系为材料,分别采用浓度为30~300 mmol·L−1 NaCl、15~60 mmol·L−1 NaHCO3、30~150 mmol·L−1 Na2SO4和5~30 mmol·L−1 Na2CO3共4种单盐溶液对芸芥种子进行胁迫处理,测定发芽率、发芽指数和胚根长等6个指标,计算耐盐临界值和极限值,采用多指标综合评价芸芥在萌发期对不同钠盐胁迫的耐受程度。 结果 结果表明:(1)4种单盐胁迫下,胚根长的耐盐临界值和耐盐极限值均最小,可作为鉴定芸芥萌发期耐盐性的主要指标;(2)碱性盐Na+≥30 mmol·L−1和中性盐Na+≥180 mmol·L−1时,芸芥种子萌发及幼苗生长受到明显抑制;(3)在NaCl和Na2SO4胁迫下,15芸芥4耐盐性最强,11芸芥32-1居中,16芸芥9最弱;在Na2CO3胁迫下,15芸芥4耐盐性最强,16芸芥9居中,11芸芥32-1最弱;在NaHCO3胁迫下,16芸芥9耐盐性最强,15芸芥4居中,11芸芥32-1最弱;(4)15芸芥4对NaCl、Na2SO4和Na2CO3胁迫的耐性临界值和极限值均最高,其耐性临界值分别为158、88和16.3 mmol·L−1,耐性极限值分别为264、130、23 mmol·L−1;15芸芥4耐中性盐适应性较广;16芸芥9对NaHCO3胁迫的耐性临界值和极限值均最高,分别为42、64 mmol·L−1,该品系耐NaHCO3的适应性较广。 结论 Na2SO4对芸芥种子萌发的抑制效应强于NaCl,Na2CO3对芸芥种子萌发的抑制效应强于NaHCO3;芸芥耐NaCl和Na2SO4胁迫次序一致,均表现为15芸芥4>11芸芥32-1>16芸芥9,耐NaHCO3胁迫依次为16芸芥9>15芸芥4>11芸芥32-1,耐Na2CO3胁迫依次为15芸芥4>16芸芥9>11芸芥32-1。 Abstract:Objectives Eruca sativa seed germination and seedling tolerance to 4 types of salts were studied in a simulated experimentation for breeding and cultivar selection. Methods Three lines of E. sativa cultivars, 15 Yunjie 4, 11 Yunjie 32-1, and 16 Yunjie 9, under the stress by 4 types of salt in varied concentrations, including 30-300 mmol·L−1 NaCl, 30-150 mmol·L−1 Na2SO4, 15-60 mmol·L−1 NaHCO3, and 5-30 mmol·L−1 Na2CO3, were used in the experimentation. In the prepared media, the rate, potential, and index of seed germination as well as the weight, radicle length, and hypocotyl length of new shoots were determined. The critical salt concentration beyond which irrevocable damage occurred to the plant and the tolerance threshold to stress imposed by a specific salt at which a plant could still revert to normal conditions were obtained for a multiple indicator evaluation on the E. sativa lines. Results (1) The least critical concentration and the tolerance threshold to the exposure of individual salt by E. sativa were found when these indicators were gauged by the radical seedling length. Therefore, the measurement was selected for the evaluation. (2) At either the alkaline Na+ was greater or equal to 30 mmol·L−1 or the neutral Na+ greater or equal to 180 mmol·L−1, the seed germination and seedling growth were significantly inhibited. (3) Under NaCl or Na2SO4 stress, 15 Yunjie 4 displayed the strongest tolerance, followed by 11 Yunjie 32-1 and 16 Yunjie 9. On Na2CO3, 15 Yunjie 4 was most tolerant, which was followed by 16 Yunjie 9 and 11 Yunjie 32-1. NaHCO3 was best tolerated by 16 Yunjie 9 and followed by 15 Yunjie 4 and 11 Yunjie 32-1. And (4) among the 3 cultivars, 15 Yunjie 4 had the highest critical salt concentrations as well as tolerance thresholds in dealing with the various salts. It showed a threshold at 158 mmol·L−1 on NaCl, 88 mmol·L−1 on Na2SO4, and 16.3 mmol·L−1 on Na2CO3, and a critical concentration on NaCl at 264 mmol·L−1, on Na2SO4 at 30 mmol·L−1, and on Na2CO3 at 23 mmol·L−1. However, 16 Yunjie 9 exhibited a high tolerance to NaHCO3 with the greatest threshold at 42 mmol·L−1 and a critical concentration at 64 mmol·L−1. Conclusion The effect of stress exerted by NaCl on E. sativa seed germination was less sever than that by Na2SO4, and that by NaHCO3 lower than Na2CO3. The NaCl and Na2SO4 tolerance of the cultivars were consistently ranked as 15 Yunjie 4>11 Yunjie 32-1>16 Yunjie 9; the NaHCO3 tolerance as 16 Yunjie 9>15 Yunjie 4>11 Yunjie 32-1; and the Na2CO3 tolerance as 15 Yunjie 4>16 Yunjie 9>11 Yunjie 32-1. The information would provide a guideline for future studies on the cultivation and management of E. sativa farming on saline-alkali land. -
Key words:
- Eruca sativa /
- seed germination /
- salt stress /
- critical salt concentration /
- tolerance threshold
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图 1 NaCl溶液胁迫对3个芸芥品系种子萌发性状的影响
注:A.发芽势;B.发芽指数;C.发芽率;D. 幼苗鲜重;E.胚根长;F.胚轴长; 小写字母表示同一种盐不同浓度处理间5%水平差异显著(P<0.05),图2~4同。
Figure 1. Effects of NaCl on germination of 3 E. sativa lines
Notes:A: germination potential; B: germination index; C: germination rate; D: fresh weight; E: radical length; F: hypocotyl length. Data with different lowercase letters indicated significant difference (P<0.05). Same for Fig.2–4.
表 1 NaCl胁迫下3个芸芥品系种子萌发期耐盐临界值和极限值
Table 1. Tolerance thresholds and limits of 3 E. sativa lines on NaCl
品系
Lines耐盐临界值Salt tolerance threshold/(mmol·L−1) 耐盐极限值Salt tolerance limit/(mmol·L−1) 品系排序
Line sequenceA B C D E F 平均值
MeanA B C D E F 平均值
Mean15芸芥4 15yunjie4 244 173 141 212 174 168 185 336 239 209 306 247 245 264 1 11芸芥32-1 11 yunjie32-1 199 143 132 223 170 101 161 271 200 198 327 240 181 236 2 16芸芥9 16yunjie9 178 127 122 133 141 68 128 243 185 188 191 218 160 198 3 平均值 Mean 207 148 132 189 162 112 158 283 208 198 275 235 195 233 指标排序 Index sequence 1 4 5 2 3 6 1 4 5 2 3 6 注:A.发芽势;B.发芽指数;C.发芽率;D. 幼苗鲜重;E.胚根长;F.胚轴长. 表2~4同.
Notes: A: germxsination potential; B: germination index; C: germination rate; D: fresh weight; E: radical length; F: hypocotyl length. Same for Table 2–4.表 2 Na2SO4胁迫下3个芸芥品系种子萌发期耐盐临界值和极限值
Table 2. Tolerance thresholds and limits of 3 E. sativa lines on Na2SO4
品系
Lines耐盐临界值Salt tolerance threshold/(mmol·L−1) 耐盐极限值Salt tolerance limit/(mmol·L−1) 品系排序
Line sequenceA B C D E F 平均值
MeanA B C D E F 平均值
Mean15芸芥4 15yunjie4 120 84 81 84 99 59 88 178 121 120 121 135 105 130 1 11芸芥32-1 11 yunjie32-1 104 82 72 88 84 74 84 168 118 112 125 125 112 127 2 16芸芥9 16yunjie9 97 65 60 81 74 61 73 161 91 87 114 102 90 108 3 平均值 Mean 107 77 71 84 86 65 82 169 110 106 120 120 102 121 指标排序 Index sequence 1 4 5 3 2 6 1 3 4 2 2 5 表 3 NaHCO3胁迫下3个芸芥品系种子萌发期耐盐临界值和极限值
Table 3. Tolerance thresholds and limits of 3 E. sativa lines on NaHCO3
品系
Lines耐盐临界值Salt tolerance threshold/(mmol·L−1) 耐盐极限值Salt tolerance limit/(mmol·L−1) 品系排序
Line sequenceA B C D E F 平均值
MeanA B C D E F 平均值
Mean15芸芥4 15yunjie4 36 34 27 39 36 21 32 65 50 45 56 53 41 52 2 11芸芥32-1 11 yunjie32-1 34 25 25 28 28 17 26 62 35 35 37 38 29 39 3 16芸芥9 16yunjie9 40 27 28 72 64 23 42 67 38 40 107 95 37 64 1 平均值 Mean 37 29 27 46 43 20 34 65 41 40 67 62 36 52 指标排序 Index sequence 3 4 5 1 2 6 2 4 5 1 3 6 表 4 Na2CO3胁迫下3个芸芥品系种子萌发期耐盐临界值和极限值
Table 4. Tolerance thresholds and limits of 3 E. sativa lines on Na2CO3
品系
Lines耐盐临界值Salt tolerance threshold/(mmol·L−1) 耐盐极限值Salt tolerance limit/(mmol·L−1) 品系排序
Line sequenceA B C D E F 平均值
MeanA B C D E F 平均值
Mean15芸芥4 15yunjie4 17 15 13 21 17 15 16.3 25 21 20 28 24 21 23 1 11芸芥32-1 11 yunjie32-1 16 13 14 13 14 10 13.3 22 18 19 19 20 15 18.8 3 16芸芥9 16yunjie9 15 14 14 15 19 12 14.8 22 19 20 22 26 18 21 2 平均值 Mean 16 14 13.7 16.3 16.7 12.3 14.8 23 19.3 19.7 23 23.3 18 21 指标排序 Index sequence 3 4 5 2 1 6 2 4 3 2 1 5 -
[1] WARWICK S I, GUGEL R K, GÓMEZ-CAMPO C, et al. Genetic variation in Eruca vesicaria (l.) cav [J]. Plant Genetic Resources: Characterization and Utilization, 2007, 5(3): 142−153. doi: 10.1017/S1479262107842675 [2] 王静, 许兴, 麻冬梅. 紫花苜蓿种质资源萌发期耐盐性鉴定 [J]. 核农学报, 2018, 32(10):1939−1948. doi: 10.11869/j.issn.100-8551.2018.10.1939WANG J, XU X, MA D M. Identification of salt-tolerant properties during germination period of Medicago sativa L. germplasm resources [J]. Journal of Nuclear Agricultural Sciences, 2018, 32(10): 1939−1948.(in Chinese) doi: 10.11869/j.issn.100-8551.2018.10.1939 [3] 曾玲玲, 季生栋, 王俊强, 等. 植物耐盐机理的研究进展 [J]. 黑龙江农业科学, 2009(5):156−159. doi: 10.3969/j.issn.1002-2767.2009.05.057ZENG L L, JI S D, WANG J Q, et al. Advances on the mechanism of plant salt tolerance [J]. Heilongjiang Agricultural Sciences, 2009(5): 156−159.(in Chinese) doi: 10.3969/j.issn.1002-2767.2009.05.057 [4] 阎旭东. 植物耐盐性鉴定及评价技术规程[M]. 北京: 中国农业科学技术出版社, 2012. [5] DAI Q L, CHEN C, FENG B, et al. Effects of different NaCl concentration on the antioxidant enzymes in oilseed rape (Brassica napus L.) seedlings [J]. Plant Growth Regulation, 2009, 59(3): 273−278. doi: 10.1007/s10725-009-9402-z [6] 刘正祥, 张华新, 杨秀艳, 等. 植物对氯化钠和硫酸钠胁迫生理响应研究进展 [J]. 世界林业研究, 2015, 28(4):17−23.LIU Z X, ZHANG H X, YANG X Y, et al. Research progress on physiological responses of plants to NaCl and Na2SO4 stress [J]. World Forestry Research, 2015, 28(4): 17−23.(in Chinese) [7] 孙万仓. 中国芸芥的分布、类型划分及油菜-芸芥杂交亲和性研究[D]. 长沙: 湖南农业大学, 2000.SUN W C. Studies on the distribution, chassification of Chinese Yunjie and intergenus crossing-compatibility of Eruca sativa mill × rapeseeds (Brassica)[D]. Changsha: Hunan Agricultural University, 2000. (in Chinese) [8] HILDA A, BABAK D H, AZRA A A. Hilda Amiripour, Babak Delnavaz Hashemloian, Azra Ataei Azimi. Morpho–physiological responses of Rocket (Eruca sativa L.) varieties to sodium sulfate (Na2SO4) stress: an experimental approach [J]. Acta Physiol Plant, 2016, 38: 246. doi: 10.1007/s11738-016-2262-z [9] 范惠玲, 白生文, 刘钊, 等. 混合盐碱胁迫对芸芥生长发育和生理性状的影响 [J]. 中国油料作物学报, 2018, 40(4):544−551. doi: 10.7505/j.issn.1007-9084.2018.04.011FAN H L, BAI S W, LIU Z, et al. Effects of mixed salt-alkaline on growth and physiological characteristics in Eruca sativa [J]. Chinese Journal of Oil Crop Sciences, 2018, 40(4): 544−551.(in Chinese) doi: 10.7505/j.issn.1007-9084.2018.04.011 [10] 范惠玲, 刘秦, 白生文, 等. 不同生态型芸芥中有机物质对盐胁迫的响应 [J]. 中国农学通报, 2017, 33(3):52−56. doi: 10.11924/j.issn.1000-6850.casb16030179FAN H L, LIU Q, BAI S W, et al. Response of organic substance to salt stress in different Eruca sativa mill. ecotypes [J]. Chinese Agricultural Science Bulletin, 2017, 33(3): 52−56.(in Chinese) doi: 10.11924/j.issn.1000-6850.casb16030179 [11] 范惠玲, 刘秦, 白生文, 等. 26份芸芥种质的耐盐性评价 [J]. 干旱地区农业研究, 2019, 37(3):15−22, 59. doi: 10.7606/j.issn.1000-7601.2019.03.02FAN H L, LIU Q, BAI S W, et al. Evaluation of salt tolerance of 26 Eruca sativa Mill [J]. Agricultural Research in the Arid Areas, 2019, 37(3): 15−22, 59.(in Chinese) doi: 10.7606/j.issn.1000-7601.2019.03.02 [12] 范惠玲, 白生文, 朱雪峰, 等. 油菜及其近缘种种子萌发期耐盐碱性差异 [J]. 作物杂志, 2019(3):178−184.FAN H L, BAI S W, ZHU X F, et al. Difference of salt-alkaline tolerance of three rape and its two relatives at germination stage [J]. Crops, 2019(3): 178−184.(in Chinese) [13] 李宏, 程平, 郑朝晖, 等. 盐旱胁迫对3种新疆造林树木种子萌发的影响 [J]. 西北植物学报, 2011, 31(7):1466−1473.LI H, CHENG P, ZHENG Z H, et al. Seeds germination of three common afforestating trees under salt and drought stress in Xinjiang [J]. Acta Botanica Boreali-Occidentalia Sinica, 2011, 31(7): 1466−1473.(in Chinese) [14] 裴毅, 杨雪君, 陈晓芬, 等. 氯化钠和碳酸氢钠胁迫对白芥种子萌发的影响 [J]. 时珍国医国药, 2015, 26(7):1750−1752.PEI Y, YANG X J, CHEN X F, et al. Effects of NaCl and NaHCO3 stress on seed gemination of Sinapis alba L [J]. Lishizhen Medicine and Materia Medica Research, 2015, 26(7): 1750−1752.(in Chinese) [15] 白小明, 王靖婷, 贺佳圆, 等. 8个野生早熟禾种子萌发期耐盐性研究 [J]. 草地学报, 2013, 21(3):546−555.BAI X M, WANG J T, HE J Y, et al. Salt resistance of eight wild Poa L. during seed germination stages [J]. Acta Agrestia Sinica, 2013, 21(3): 546−555.(in Chinese) [16] 韩清芳, 李崇巍, 贾志宽. 不同苜蓿品种种子萌发期耐盐性的研究 [J]. 西北植物学报, 2003, 23(4):597−602. doi: 10.3321/j.issn:1000-4025.2003.04.014HAN Q F, LI C W, JIA Z K. The study on salt tolerance of different alfalfa varieties in germinating period [J]. Acta Botanica Boreali-Occidentalia Sinica, 2003, 23(4): 597−602.(in Chinese) doi: 10.3321/j.issn:1000-4025.2003.04.014 [17] 吴凤萍, 韩清芳, 贾志宽. 4个白花苜蓿品系种子萌发期耐盐性研究 [J]. 草业科学, 2008, 25(8):57−62. doi: 10.3969/j.issn.1001-0629.2008.08.011WU F P, HAN Q F, JIA Z K. Preliminary study on the salt tolerance of four alfalfa lines in seed germination period [J]. Pratacultural Science, 2008, 25(8): 57−62.(in Chinese) doi: 10.3969/j.issn.1001-0629.2008.08.011 [18] 李昀, 沈禹颖, 阎顺国. NaCl胁迫下5种牧草种子萌发的比较研究 [J]. 草业科学, 1997, 14(2):50−53.LI Y, SHEN Y Y, YAN S G. Comparative studies of effect of NaCl stress on the seed germination of 5 forage species [J]. Pratacultural Science, 1997, 14(2): 50−53.(in Chinese) [19] 陈德明. 盐渍环境中的植物耐盐性及其影响因素 [J]. 土壤学进展, 1994, 22(5):22−29.CHEN D M. Plant salt tolerance under salinity condition and influencing factors [J]. Progress in soil science, 1994, 22(5): 22−29.(in Chinese) [20] 祁栋灵, 韩龙植, 张三元. 水稻耐盐/碱性鉴定评价方法 [J]. 植物遗传资源学报, 2005, 6(2):226−230, 235. doi: 10.3969/j.issn.1672-1810.2005.02.021QI D L, HAN L Z, ZHANG S Y. Methods of characterization and evaluation of salt or alkaline tolerance in rice [J]. Journal of Plant Genetic Resources, 2005, 6(2): 226−230, 235.(in Chinese) doi: 10.3969/j.issn.1672-1810.2005.02.021 [21] 王佳佳, 谷思玉. NaCl与Na2CO3+NaHCO3对玉米萌发期胁迫效应的比较 [J]. 作物杂志, 2012(2):138−141. doi: 10.3969/j.issn.1001-7283.2012.02.037WANG J J, GU S Y. Comparison of NaCl and Na2CO3+NaHCO3 stress on corn germination [J]. Crops, 2012(2): 138−141.(in Chinese) doi: 10.3969/j.issn.1001-7283.2012.02.037 [22] 王清华, 杨建平, 张中华, 等. 盐胁迫对不同品种辣椒种子萌发特性的影响 [J]. 西北农业学报, 2007, 16(3):136−140. doi: 10.3969/j.issn.1004-1389.2007.03.035WANG Q H, YANG J P, ZHANG Z H, et al. Effect of salt stress on germination characteristics of different pepper varieties [J]. Acta Agriculturae Boreali-Occidentalis Sinica, 2007, 16(3): 136−140.(in Chinese) doi: 10.3969/j.issn.1004-1389.2007.03.035 [23] 张振霞, 刘萍, 杨中艺. 25个多年生黑麦草品种萌发期对盐胁迫的抗性研究 [J]. 草业科学, 2007, 24(2):14−19. doi: 10.3969/j.issn.1001-0629.2007.02.004ZHANG Z X, LIU P, YANG Z Y. Salt tolerance of 25 Lolium perenne varieties at germination stage [J]. Pratacultural Science, 2007, 24(2): 14−19.(in Chinese) doi: 10.3969/j.issn.1001-0629.2007.02.004 [24] 李孔晨, 皇甫江云, 卢欣石. 一年生黑麦草萌发期耐盐性综合评价 [J]. 草地学报, 2010, 18(3):388−393, 398.LI K C, HUANGFU J Y, LU X S. Salt tolerance evaluation of 10 varieties of Lolium multif lam. at germination period [J]. Acta Agrestia Sinica, 2010, 18(3): 388−393, 398.(in Chinese)