Effect of 2, 4-Brassinosteroid on Growth of Hordeum vulgare Seedlings under Cd-stress
-
摘要: 采用室内培养法,研究2,4-表油菜素内酯(2,4-EBL)对Cd胁迫下大麦幼苗的生长影响。结果表明,在所试Cd质量浓度(20 mg·L-1)和2,4-EBL质量浓度(0.2 mg·L-1)处理下,与对照相比,Cd处理使大麦幼苗生长受到抑制,叶片受到损伤,表现为根长、苗长、生物量、叶绿素含量、可溶性糖和可溶性蛋白含量、POD和CAT活性等指标下降,而叶片相对电导率、MDA、O2-含量、脯氨酸含量、SOD活性等指标增加。喷施2,4-EBL后,与单一Cd处理组相比,大麦幼苗根长、苗长、生物量和叶绿素含量显著增加,促进了光合作用和幼苗生长,降低了叶片相对电导率、MDA及O2-含量,而使渗透调节物质(可溶性糖、脯氨酸、可溶性蛋白)和抗氧化酶(SOD、POD、CAT)活性增加。表现出2,4-EBL对Cd胁迫后的大麦幼苗具有一定的缓解作用,增强了大麦在逆境中的适应能力。Abstract: Effect of 2, 4-brassinosteroid on the growth of barley (Hordeum vulgare) seedlings under Cd-stress were studied by an indoor culture experiment. In comparison with control, the treatment of Cd at 20 mg·L-1 inhibited the seedling growth and damaged the leaf physiology manifesting in the decreased root length and plant height, reduced contents on biomass, chlorophyll, soluble sugars and soluble protein, declined activities of POD and CAT, increased relative electrical conductivity, heightened contents of MDA, O2- and proline, and elevated activity of SOD. After spraying 2, 4-brassinosteroid at 0.2 mg·L-1, the Cd-stressed seedlings significantly increased on the root length, plant height, biomass and chlorophyll. The treatment promoted the photosynthesis and seedling growth, reduced the leaf relative conductivity, MDA and O2-, and facilitated an osmotic adjustment with the substances (e.g., soluble sugars, proline and soluble protein), and enhanced the antioxidant enzymatic activities on SOD, POD and CAT. It appeared that the plant hormone could alleviate the ill-effects of Cd-stress on and enrich the adaptability in the battle against the adversity for the barley seedlings.
-
Key words:
- Cd-stress /
- Hordeum vulgare /
- brassinosteroid /
- seedling growth
-
表 1 2, 4-表油菜素内酯对镉胁迫下大麦幼苗根长、苗长和生物量以及叶片损伤的影响
Table 1. Effects of 2, 4-brassinosteroid on root length, seedling height, biomass and leaf physiology of barley seedlings under Cd-stress
处理时间/d 处理组 根长/cm 苗长/cm 生物量/mg MDA含量/(μmol·g-1) 相对电导率/% O2-含量/(nmol·g-1) 3 CK 6.26±0.29b 7.92±0.34b 76.71±2.93b 2.21±0.08c 13.11±0.55c 34.51±2.24c EBL 7.12±0.22a 8.66±0.30a 84.09±3.05a 2.13±0.11cd 11.95±0.51d 32.17±2.16d Cd 4.89±0.17d 6.37±0.27c 59.45±2.46d 3.79±0.22a 33.29±1.23a 54.63±3.17a Cd+EBL 5.62±0.35c 7.14±0.41b 71.82±2.95c 2.97±0.16b 30.03±1.25b 39.43±2.43b 5 CK 10.07±0.32b 12.41±0.75b 116.24±4.62b 2.27±0.18c 14.47±0.61c 35.21±2.17c EBL 11.41±0.17a 13.16±0.64a 128.26±3.64a 2.06±0.20cd 12.28±0.54d 33.46±2.21cd Cd 7.79±0.18d 9.72±0.53d 97.71±3.52d 4.63±0.23a 37.62±2.09a 77.42±3.44a Cd+EBL 9.02±0.31c 10.96±0.65c 112.29±3.45c 3.82±0.14b 34.29±2.68b 58.68±3.36b 注:同列数据后不同小写字母表示差异显著(P < 0.05),表 2、3同。 表 2 2, 4-表油菜素内酯对镉胁迫下大麦幼苗渗透调节物质和抗氧化酶活性的影响
Table 2. Effects of 2, 4-brassinosteroid on osmosis adjusting substances and antioxidant activities of barley seedlings under Cd-stress
处理时间/d 处理组 可溶性糖含量/(mg·g-1) 脯氨酸含量/(μg·g-1) 可溶性蛋白含量/(mg·g-1) SOD活性/(U·g-1) POD活性/(U·g-1·min-1) CAT活性/(U·g-1·min-1) 3 CK 6.03±0.34c 11.06±0.67d 35.32±2.66b 89.13±3.03cd 201.37±3.65bc 401.76±6.81ab EBL 6.95±0.41a 11.67±0.58c 36.29±2.73a 90.32±3.11c 202.11±4.73b 402.12±7.23a Cd 5.81±0.27d 17.26±0.92b 31.25±3.01d 139.06±4.14b 198.37±3.67d 285.92±3.75 d Cd+EBL 6.34±0.46b 17.81±0.87a 33.72±2.99c 168.96±4.26a 229.34±4.55a 353.17±4.64c 5 CK 7.11±0.29b 10.87±0.71d 36.24±3.24b 136.71±3.92cd 389.36±6.67bc 411.53±5.27ab EBL 8.06±0.49a 11.93±0.62c 36.88±4.17a 137.09±4.05c 390.04±6.81b 411.89±5.39a Cd 6.14±0.36d 14.96±0.73b 29.16±2.72d 187.34±4.65b 351.76±5.49d 286.28±3.79d Cd+EBL 6.59±0.35c 15.16±0.81a 31.41±3.69c 236.81±5.77a 409.12±6.92a 382.92±4.03c 表 3 2, 4-表油菜素内酯对镉胁迫下大麦幼苗叶绿素含量的影响
Table 3. Effects of 2, 4-brassinosteroid on chlorophyll content of barley seedlings under Cd-stress
处理时间/d 处理组 叶绿素a含量/(mg·g-1) 叶绿素b含量/(mg·g-1) 叶绿素(a+b)含量/(mg·g-1) 叶绿素a/b 3 CK 0.94±0.03b 0.53±0.01b 1.47±0.04b 1.77±0.05b EBL 1.03±0.02a 0.59±0.02a 1.62±0.06a 1.75±0.04bc Cd 0.71±0.04d 0.39±0.01d 1.1±0.04d 1.82±0.07a Cd+EBL 0.80±0.02c 0.46±0.02c 1.26±0.07c 1.74±0.03c 5 CK 0.99±0.04b 0.55±0.03b 1.54±0.04b 1.80±0.06b EBL 1.13±0.03a 0.66±0.03a 1.79±0.06a 1.71±0.03c Cd 0.65±0.02d 0.35±0.01d 1.00±0.03d 1.86±0.04a Cd+EBL 0.71±0.03c 0.42±0.02c 1.13±0.07c 1.69±0.03c -
[1] 郑洁, 王磊.油菜素内酯在植物生长发育中的作用机制研究进展[J].中国农业科技导报, 2014, 16(1):52-58. http://www.wanfangdata.com.cn/details/detail.do?_type=perio&id=zgnykjdb201401009 [2] CHOUDHARY S P, YU J Q, YAMAGUCHI-SHINOZAKI K, et al.Benefits of brassinosteroid crosstalk[J].Cell, 2012, 17(10):594-605. http://www.wanfangdata.com.cn/details/detail.do?_type=perio&id=ff2e8209407fcdfeb0a8c3748ca2b962 [3] PRITI K.Brassinosteroid-mediated stress responses[J].Plant Growth Regul, 2003, 22(4):289-292. doi: 10.1007/s00344-003-0058-z [4] ZHANG C, BAI M Y, CHONG K.Brassinosteroid-mediated regulation of agronomic traits in rice[J].Plant Cell Rep, 2014, 33(5):683-696. doi: 10.1007/s00299-014-1578-7 [5] 何俊瑜, 任艳芳, 王阳阳, 等.不同耐性水稻幼苗根系对镉胁迫的形态及生理响应[J].生态学报, 2011, 31(2):522-528. http://d.old.wanfangdata.com.cn/Periodical/stxb201102026 [6] HAYAT S, ALI B, HASAN S A, et al. Brassinosteroids enhanced the level of antioxidants under cadmium stress in Brassica juncea[J].Environmental and Experimental Botany, 2007, 60(1):33-41. doi: 10.1016/j.envexpbot.2006.06.002 [7] 郑爱珍.镉胁迫对芥蓝根系质膜过氧化及ATPase活性的影响[J].生态学报, 2012, 32(2):483-488. http://d.old.wanfangdata.com.cn/Periodical/stxb201202018 [8] HASAN S A, HAYAT S, ALI B, et al.28-Homobrassinolide protects chickpea from cadmium toxicity by stimulating antioxidants[J].Environmental Pollution, 2008,151(1):60-66. http://www.wanfangdata.com.cn/details/detail.do?_type=perio&id=2baa0ccb697e9766bb9045f51a72c1d2 [9] 曾翔, 张玉烛, 王凯荣, 等.镉对水稻种子萌发的影响[J].应用生态学报, 2007, 18(7):1665-1668. doi: 10.3321/j.issn:1001-9332.2007.07.041 [10] 顾自奋, 黄志仁, 许如根, 等.近10年世界大麦生产概况[J].大麦科学, 2001, 18(1):1-4. doi: 10.3969/j.issn.1673-6486.2001.01.001 [11] 殷欣, 张海玲, 吴杨, 等.24-表油菜素内酯对镉胁迫下大豆苗期生理特性的影响[J].核农学报, 2016, 30(2):364-371. http://d.old.wanfangdata.com.cn/Periodical/hnxb201602021 [12] 高会玲, 刘金隆, 郑青松, 等.外源油菜素内酯对镉胁迫下菊芋幼苗光合作用及镉富集的调控效应[J].生态学报, 2013, 33(6):1935-1943. http://d.old.wanfangdata.com.cn/Periodical/stxb201306028 [13] 袁博, 金晓霞, 肖玮, 等.外源表油菜素内酯(2, 4-EBL)对镉(Cd)胁迫下番茄幼苗的缓解效应[J].分子植物育种, 2018, 16(9):2955-2965. http://d.old.wanfangdata.com.cn/Thesis/Y3402255 [14] 张义贤, 李晓科.镉、铅及其复合污染对大麦幼苗部分生理指标的影响[J].植物研究, 2008, 28(1):43-46. http://d.old.wanfangdata.com.cn/Periodical/zwyj200801010 [15] 肖志华, 张义贤, 张喜文, 等.外源铅、铜胁迫对不同基因型谷子幼苗生理生态特性的影响[J].生态学报, 2012, 32(3):889-897. http://d.old.wanfangdata.com.cn/Periodical/stxb201203025 [16] 李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社, 2000:184-185,258-261. [17] XIE Z, DUAN L, TIAN X, et al.Coronatine alleviates salinity stress in cotton by improving the antioxidative defense system and radical-scavenging activity[J].Journal of Plant Physiology, 2008,165(4):375-384. doi: 10.1016/j.jplph.2007.06.001 [18] 燕傲蕾, 吴亭亭, 王友保, 等.三种观赏植物对重金属镉的耐性与积累特性[J].生态学报, 2010, 30(9):2491-2498. http://d.old.wanfangdata.com.cn/Periodical/stxb201009030 [19] ARIFA K, FAHEEM A.Effect of exogenous application of 2, 4-epibrassinolideon growth, protein contents, and antioxidant enzyme activities of in vitro-grown Solanum tuberosum L.under salt stress, In Vitro Cellular & Devlopmental Biology-Plant[J].Dev Biol Plant, 2016, 52(1):1-11. [20] 王晓娟, 王文斌, 杨龙吴, 等.重金属镉(Cd)在植物体内的转运途径及其调控机制[J].生态学报, 2015, 35(23):7921-7929. http://d.old.wanfangdata.com.cn/Periodical/stxb201523035 [21] 吴杨, 高慧纯, 张必弦, 等.2, 4-表油菜素内酯对盐碱胁迫下大豆生育、生理及细胞超微结构的影响[J].中国农业科学, 2017, 50(5):811-821. http://www.cnki.com.cn/Article/CJFDTotal-ZNYK201705004.htm [22] HAUBRICK L L, ASSMANN S M. Brassinosteroids and plant function:some clues, more puzzles[J].Plant Cell and Environment, 2006, 29(3):446-457. doi: 10.1111/pce.2006.29.issue-3 [23] 王爱君, 王志敏, 汤青林, 等.超干处理对种子生物膜及种子活力的影响[J].南方农业:园林花卉版, 2011, 5(4):52-54. http://d.old.wanfangdata.com.cn/Periodical/nfny-lshyb201102016 [24] 乔琳, 李杰, 胡春红, 等.外施表油菜素内酯缓解玉米幼苗铅毒害机制研究[J].核农学报, 2014, 28(11):2126-2131. doi: 10.11869/j.issn.100-8551.2014.11.2126 [25] SANTOS L R, BATISA B L, LOBATO A K S.Brassinos-teroids mitigate cadmium toxicity in cowpea plants[J].Photosynthetica, 2017, 55(4):1-15. [26] SYED A H B, WANG R F, IMRUL M A, et al.Genotype-dependent effect of exogenous 24-epibrassinolide on chromium-induced changes in ultrastructure and physicochemical traits in tobacco seedlings[J]. Environ Sci Pollut Res, 2016, 23(18):1-10. http://www.wanfangdata.com.cn/details/detail.do?_type=perio&id=3bd9f43c55ed401f44905c8f1dd2f0c0 [27] 靳开川, 何金环.油菜素内酯在植物抗逆中的作用及信号传导机制综述[J].江苏农业科学, 2017, 45(14):4-7. http://d.old.wanfangdata.com.cn/Periodical/jsnykx201714002 [28] HASAN S A, HAYAT S, AHMAD A. Brassinosteroids protect photosynthetic machinery against the cadmium induced oxidative stress in two tomato cultivars[J].Chemosphere, 2011, 84(10):1446-1451. doi: 10.1016/j.chemosphere.2011.04.047 [29] 尹博, 王秀峰, 姜春辉, 等.外源油菜素内酯对番茄铜胁迫的缓解效应[J].植物营养与肥料学报, 2012, 18(1):162-168. http://d.old.wanfangdata.com.cn/Thesis/Y2116982 [30] XIA X J, HUANG L F, ZHOU Y H. Brassinosteroids promote photosynthesisand growth by enhancing activation of Rubisco and expression of photosynthetic genes in Cucumis sativus L.[J].Planta, 2009,230:1185-1196. doi: 10.1007/s00425-009-1016-1 [31] DING J, SHI K, ZHOU Y H.Effects of root and foliar applications of 2, 4-epibrassinolide on fusarium wilt and antioxidant metabolism in cucumber roots[J].Hort Science, 2009, 44:1340-1345. doi: 10.1007%2Fs10658-008-9401-3 [32] 张义贤, 付亚萍, 肖志华, 等.铅胁迫对不同基因型谷子幼苗生理特性及基因组DNA多态性的影响[J].农业环境科学学报, 2013, 32(3):478-484. http://www.wanfangdata.com.cn/details/detail.do?_type=perio&id=nyhjbh201303011 [33] ANURADHA S, RAO S.The effect of brassinosteroids on radish seedlings growing under Cadmium stress[J].Plant Soil and Environment, 2007, 53(11):465-472. [34] 钟杨, 陈洪国.芸苔素内酯水剂对辣椒幼苗水分胁迫生理的影响[J].北方园艺, 2011(2):42-44. http://d.old.wanfangdata.com.cn/Periodical/bfyany201102014 [35] 魏湜, 吉彪, 李赞, 等.盐胁迫下油菜素内酯对玉米幼苗生理特性的影响[J].东北农业大学学报, 2018, 49(5):9-16. doi: 10.3969/j.issn.1005-9369.2018.05.002 [36] 郑爱珍.镉胁迫对芥蓝根系质膜过氧化及ATPase活性的影响[J].生态学报, 2012, 32(2):483-488. http://d.old.wanfangdata.com.cn/Periodical/stxb201202018 [37] 李涛涛, 高永峰, 马瑄, 等.外源油菜素内酯对三种杨树在干旱、盐和铜胁迫下光合生理的影响[J].基因组学与应用生物学, 2016, 35(1):218-226. http://www.wanfangdata.com.cn/details/detail.do?_type=perio&id=jyzxyyyswx201601034 [38] FARIDUDDIN Q, MIR B A, YUSUF M, et al.2, 4-epibrassinolide and/or putrescine trigger physiological and biochemical responses for the salt stress mitigation in Cucumis sativus L[J].Photosynthetica, 2014, 52(3):464-474. doi: 10.1007/s11099-014-0052-7 [39] BAJGUZ A, HAYAT S.Effects of brassinosteroids on the plant responses to environmental stresse[J].Plant Physiology and Biochemistry, 2009, 47(1):1-8.