Brassinosteroids make plant life easier under abiotic stresses mainly by modulating major components of antioxidant defense system

Frontiers in Environmental Science

Volumn 2, Issue JAN, 2015, Pages

  Vardhini, Bojjam V ^a   Anjum, Naser A ^b  

^a TELANGANA UNIVERSITY   (India)

^b UNIVERSITY OF AVEIRO   (Portugal)

Author keywords

Abiotic stress; Antioxidant defense system; Brassinosteroids; Reactive oxygen species; Tolerance

Indexed keywords

EID: 85030864322     PISSN: None     EISSN: 2296665X     Source Type: Journal    
DOI: 10.3389/fenvs.2014.00067     Document Type: Article

References (207)

1. Abbas, S., Latif, H. H., and Elsherbiny, E. A. (2013). Effect of 24-epibrassinolide on the physiological and genetic changes on two varieties of pepper under salt stress conditions. Pak. J. Bot. 45, 1273-1284.
2. Aghdam, M. S., Asghari, M., Farmani, B., Mohayeji, M., and Moradbeygi, H. (2012). Impact of postharvest brassinosteroids treatment on PAL activity in tomato fruit in response to chilling stress. Sci. Hort. 144, 116-120. doi: 10.1016/j.scienta.2012.07.008
3. Aghdam, M. S., and Mohammadkhani, N. (2014). Enhancement of chilling stress tolerance of tomato fruit by postharvest brassinolide treatment. Food Bioproc. Technol. 7, 909-914. doi: 10.1007/s11947-013-1165-x
4. Ahammed, G. J., Choudhary, S. P., Chen, S., Xia, X., Shi, K., Zhou, Y., et al. (2013a). Role of brassinosteroids in alleviation of phenanthrene-cadmium co-contamination-induced photosynthetic inhibition and oxidative stress in tomato. J. Exp. Bot. 64, 199-213. doi: 10.1093/jxb/ers323
5. Ahammed, G. J., Gao, C. J., Ogweno, J. O., Zhou, Y. H., Xia, X. J., Mao, W. H., et al. (2012). Brassinosteroids induce plant tolerance against phenanthrene by enhancing degradation and detoxification in Solanum lycopersicum L. Ecotoxicol. Environ. Saf. 80, 28-36. doi: 10.1016/j.ecoenv.2012.02.004
6. Ahammed, G. J., Ruan, Y. P., Zhou, J., Xia, X. J., Shi, K., Zhou, Y. H., et al. (2013). Brassinosteroid alleviates polychlorinated biphenyls-induced oxidative stress by enhancing antioxidant enzymes activity in tomato. Chemosphere 90, 2645-2653. doi: 10.1016/j.chemosphere.2012.11.041
7. Ahammed, G. J., Yuan, H. L., Ogweno, J. O., Zhou, Y. H., Xia, X. J., Mao, W. H., et al. (2012). Brassinosteroid alleviates phenanthrene and pyrene phytotoxicity by increasing detoxification activity and photosynthesis in tomato. Chemosphere 86, 546-555. doi: 10.1016/j.chemosphere.2011.10.038
8. Ahammed, G. J., Zhang, S., Shi, K., Zhou, Y. H., and Yu, J. Q. (2012). Brassinosteroid improves seed germination and early development of tomato seedling under phenanthrene stress. Plant Growth Regul. 68, 87-96. doi: 10.1007/s10725-012-9696-0
9. Ahmed, M. A., and Shalaby, M. A. F. (2013). Physiological role of brassinolide in improving yield of six wheat cultivars (Triticum aestivum L.) grown under newly reclaimed sandy soil. J. Appl. Sci. Res. 9, 6387-6393.
10. Alam, M. M., Hayat, S., Ali, B., and Ahmad, A. (2007). Effect of 28-homobrassinolide treatment on nickel toxicity in Brassica juncea. Photosynthetica 45, 139-142. doi: 10.1007/s11099-007-0022-4
11. Ali, B., Hasan, S. A., Hayat, S., Hayat, Q., Yadav, S., Fariduddin, Q., et al. (2008b). A role for brassinosteroids in the amelioration of aluminium stress through antioxidant system in mung bean (Vigna radiata L. Wilczek). Environ. Exp. Bot. 62, 153-159. doi: 10.1016/j.envexpbot.2007.07.014
12. Ali, B., Hayat, S., and Ahmad, A. (2007). 28-Homobrassinolide ameliorates the saline stress in chickpea (Cicer arietinum L.). Environ. Exp. Bot. 59, 217-223. doi: 10.1016/j.envexpbot.2005.12.002
13. Ali, B., Hayat, S., Fariduddin, Q., and Ahmad, A. (2008a). 24-Epibrassinolide protects against the stress generated by salinity and nickel in Brassica juncea. Chemosphere 72, 1387-1392. doi: 10.1016/j.chemosphere.2008. 04.012
14. Ali, Q., Athar, H. U. R., and Ashraf, M. (2006). Influence of exogenously applied brassinosteroids on the mineral nutrient status of two wheat cultivars grown under saline conditions. Pak. J. Bot. 38, 1621-1632.
15. Alyemeni, M. N., Hayat, S., Wijaya, L., and Anaji, A. (2013). Foliar application of 28-homobrassinolide mitigates salinity stress by increasing the efficiency of photosynthesis in Brassica juncea. Acta Bot. Bras. 27, 502-505. doi: 10.1590/S0102-33062013000300007
16. Anjum, N. A., Sofo, A., Scopa, A., Roychoudhury, A., Gill, S. S., Iqbal, M., et al. (2014). Lipids and proteins-major targets of oxidative modifications in abiotic stressed plants. Environ. Sci. Pollut. Res. doi: 10.1007/s11356-014-3917-1.
17. Anjum, N. A., Umar, S., and Chan, M. T. (2010). Ascorbate-Glutathione Pathway and Stress Tolerance in Plants. Dordrecht: Springer.
18. Anjum, N. A., Umar, S., and Ahmad, A. (2012). Oxidative Stress in Plants: Causes, Consequences and Tolerance. New Delhi: IK International Publishing House.
19. Anjum, S. A., Wang, L. C., Farooq, M., Hussain, M., Xue, L. L., and Zou, C. M. (2011). Brassinolide application improves the drought tolerance in maize through modulation of enzymatic antioxidants and leaf gas exchange. J. Agron. Crop Sci. 197, 177-185. doi: 10.1111/j.1439-037X.2010.00459.x
20. Anuradha, S., and Rao, S. S. R. (2007). Effect of 24-epibrassinolide on the growth and antioxidant enzyme activities in radish seedlings under lead toxicity. Indian J. Plant Physiol. 12, 396-400.
21. Anuradha, S., and Rao, S. S. R. (2007). The effect of brassinosteroids on radish (Raphanus sativus L.) seedlings growing under cadmium stress. Plant Soil Environ. 53, 465-472.
22. Anuradha, S., and Rao, S. S. R. (2009). Effect of 24-epibrassinolide on the photosynthetic activity of radish plants under cadmium stress. Photosynthetica 47, 317-320. doi: 10.1007/s11099-009-0050-3
23. Arora, N., Bhardwaj, R., Sharma, P., and Arora, H. K. (2008a). Effects of 28-homobrassinolide on growth, lipid peroxidation and antioxidative enzyme activities in seedlings of Zea mays L. under salinity stress. Acta Physiol. Plant 30, 833-839. doi: 10.1007/s11738-008-0188-9
24. Arora, N., Bhardwaj, R., Sharma, P., Arora, H. K., and Arora, P. (2008b). Amelioration of zinc toxicity by 28-homobrassinolide in Zea mays L. Can. J. Pure Appl. Sci. 2, 503-509.
25. Arora, P., Bhardwaj, R., and Kanwar, M. K. (2010a). 24-Epibrassinolide induced antioxidative defense system of Brassica juncea L. under Zn metal stress. Physiol. Mol. Biol. Plant 16, 285-293. doi: 10.1007/s12298-010-0031-9
26. Arora, P., Bhardwaj, R., and Kanwar, M. K. (2010b). 24-Epibrassinoliide regulated diminution of Cr metal toxicity in Brassica juncea L. plants. Braz. J. Plant Physiol. 22, 159-165. doi: 10.1590/S1677-04202010000300002
27. Arora, P., Bhardwaj, R., and Kanwar, M. K. (2012). Effect of 24-epibrassinolide on growth, protein content and antioxidative defense system of Brassica juncea L. subjected to cobalt ion toxicity. Acta Physiol. Plant 34, 2007-2017. doi: 10.1007/s11738-012-1002-2
28. Arteca, J. M., and Arteca, R. N. (2001). Brassinosteroid-induced exaggerated growth in hydroponically grown Arabidopsis plants. Physiol. Plant 112, 104-112. doi: 10.1034/j.1399-3054.2001.1120114.x
29. Asami, T., Mink, Y. K., Nagata, N., Yamagishi, K., Takatsuto, S., Fujioka, S., et al. (2000). Characterization of brassinozole, a triazole-type brassinosteroid biosynthesis inhibitor. Plant Physiol. 123, 93-100. doi: 10.1104/pp.123.1.93
30. Asgher, M., Khan, M. I. R., Anjum, N. A., and Khan, N. A. (2014). Minimizing toxicity of cadmium in plants-role of plant growth regulators. Protoplasma. doi: 10.1007/s00709-014-0710-4.
31. Ashraf, M., Akram, N. A., Arteca, R. N., and Foolad, M. R. (2010). The physiological, biochemical and molecular roles of brassinosteroids and salicylic acid in plant processes and salt tolerance. Critic. Rev. Plant Sci. 29, 162-190. doi: 10.1080/07352689.2010.483580
32. Avalbaev, A. M., Yuldashev, R. A., Fatkhutdinova, R. A., Urusov, F. A., Safutdinova, Y. V., and Shakirova, F. M. (2010). The influence of 24-epibrassinolide on the hormonal status of wheat plants under sodium chloride. Appl. Biochem. Microbiol. 46, 99-102. doi: 10.1134/S0003683810010175
33. Aydin, Y., Talas-Ogras, T., Ipekci-Altas, Z., and Gozukirmizi, N. (2006). Effects of brassinosteroid on cotton regeneration via somatic embryogenesis. Biologia (Bratislava) 61, 289-293. doi: 10.2478/s11756-006-0053-5
34. Bajguz, A. (2009). Brassinosteroid enhanced the level of abscisic acid in Chlorella vulgaris subjected to short-term heat stress. J. Plant Physiol. 166, 882-886. doi: 10.1016/j.jplph.2008.10.004
35. Bajguz, A. (2010). An enhancing effect of exogenous brassinolide on the growth and antioxidant activity in Chlorella vulgaris cultures under heavy metals stress. Environ. Exp. Bot. 68, 175-179. doi: 10.1016/j.envexpbot.2009. 11.003
36. Bajguz, A., and Piotrowska-Niczyporuk, A. (2014). Interactive effect of brassinosteroids and cytokinins on growth, chlorophyll, monosaccharide and protein content in the green alga Chlorella vulgaris (Trebouxiophyceae). Plant Physiol. Biochem. 80C, 176-183. doi: 10.1016/j.plaphy.2014.04.009
37. Behnamnia, M., Kalantari, K. M., and Rezanejad, F. (2009). Exogenous application of brassinosteroid alleviates drought-induced oxidative stress in Lycopersicon esculentum L. Gen. Appl. Plant Physiol. 35, 22-34.
38. Bishop, G. J., and Yakota, T. (2001). Plant steroid hormones, brassinosteroids: current highlights of molecular aspects on their synthesis,/metabolism, transport, perception and response. Plant Cell Physiol. 42, 114-120. doi: 10.1093/pcp/pce018
39. Borcioni, E., and Bonato-Negrelle, R. R. (2012). Application of analogue brassinosteroide (Biobras 16 (R)) germination and growth in vitro culture of zygotic embryos and acclimatization of bocaiuva. Ciencia Rural. 42, 270-275.
40. Bray, E. A., Bailey-Serres, J., and Weretilnyk, E. (2000). "Responses to abiotic stresses," in Biochemistry and Molecular Biology of Plants, eds W. Gruissem, B. Buchnnan, and R. Jones (Rockville, MD: American Society of Plant Physiologists), 1158-1249.
41. Cao, Y. Y., and Zhao, H. (2007). Protective roles of brassinolide in rice seedlings under heat stress. Zhongguo Shuidao Kexue (China Rice Science) 21, 525-529.
42. Carange, J., Longpré, F., Daoust, B., and Martinoli, M. G. (2011). 24-Epibrassinolide, a phytosterol from the brassinosteroid family, protects dopaminergic cells against MPP+-induced oxidative stress and apoptosis. J. Toxicol. 2011:392859. doi: 10.1155/2011/392859
43. Cheng, Y., Zhu, W., Chen, Y., Ito, S., Asami, T., and Wang, X. (2014). Brassinosteroids control root epidermal cell fate via direct regulation of a MYB-bHLH-WD40 complex by GSK3-like kinases. eLife 3:e02525. doi: 10.7554/eLife.02525
44. Choudhary, S. P., Kanwar, M., Bhardwaj, R., Gupta, B. D., and Gupta, R. K. (2011). Epibrassinolide ameliorates Cr (VI) stress via influencing the levels of indole-3-acetic acid, abscisic acid, polyamines and antioxidant system of radish seedlings. Chemosphere 84, 592-600. doi: 10.1016/j.chemosphere.2011. 03.056
45. Creelman, R. A., and Mullet, J. E. (1997). Oligosaccharins, brassinolides, and jasmonates: nontraditional regulators of plant growth, development, and gene expression. Plant Cell 9, 1211-1223. doi: 10.1105/tpc.9.7.1211
46. Dalio, R. J. D., Pinheiro, H. P., Sodek, L., and Haddad, C. R. B. (2013). 24-Epibrassinolide restores nitrogen metabolism of pigeon pea under saline stress. Bot. Stud. 54:9. doi: 10.1186/1999-3110-54-9
47. Das, T., Shukla, Y. M., Poonia, T. C., Meena, M., and Meena, M. D. (2013). Effects of brassinolide on physiological characteristics of rice (Oryza sativa L.) with different salinity levels. Ann. Biol. 29, 228-231.
48. Daur, I., and Tatar, O. (2013). Effects of gypsum and brassinolide on soil properties, and berseem (Trifolium alexandrinum L.) growth, yield and chemical composition grown on saline soil. Legume Res. 36, 306-311.
49. Deng, Z. P., Zhang, X., Tang, W. Q., Oses-Prieto, J. A., Suzuki, N., Gendron, J. M., et al. (2007). A proteomics study of brassinosteroid response in Arabidopsis. Mol. Cell. Proteomics 6, 2058-2071. doi: 10.1074/mcp. M700123-MCP200
50. Ding, H. D., Zhu, X. H., Zhu, Z. W., Yang, S. J., Zha, D. S., and Wu, X. X. (2012). Amelioration of salt-induced oxidative stress in eggplant by application of 24-epibrassinolide. Biol. Plant 56, 767-770. doi: 10.1007/s10535-01 2-0108-0
51. Divi, U. K., and Krishna, P. (2010). Overexpression of the brassinosteroid biosynthetic gene AtDWF4 in Arabidopsis seeds overcomes abscisic acid-induced inhibition of germination and increases cold tolerance in transgenic seedlings. J. Plant Growth Regul. 29, 385-393. doi: 10.1007/s00344-010-9150-3
52. Divi, U. K., Rahman, T., and Krishna, P. (2010). Brassinosteroid-mediated stress tolerance in Arabidopsis shows interactions with abscisic acid, ethylene and salicylic acid pathways. BMC Plant Biol. 10:151. doi: 10.1186/1471-222 9-10-151
53. Domagalska, M. A., Sarnowska, E., Nagy, F., and Davis, S. J. (2010). Genetic analyses of interactions among gibberellin, abscisic acid, and brassinosteroids in the control of flowering time in Arabidopsis thaliana. PLoS ONE 5:e14012. doi: 10.1371/journal.pone.0014012
54. Dong, D., Li, Y., and Jiang, L. (2008). Effects of brassinosteroids on photosynthetic characteristics in soybean under aluminum stress. Zuowu Xuebao (Acta Agronomica Sinica) 34, 1673-1678. doi: 10.3724/SP.J.1006.2008. 01673
55. Durigan, D. R. J., Pinheiro, H. P., Sodek, L., and Haddad, C. R. B. (2011). The effect of 24-epibrassinolide and clotrimazole on the adaptation of Cajanus cajan (L.) Millsp to salinity. Acta Physiol. Plant 33, 1887-1896. doi: 10.1007/s11738-011-0732-x
56. Ekinci, M., Yildirim, E., Dursun, A., and Turan, M. (2012). Mitigation of salt stress in lettuce (Lactuca sativa L. var. Crispa) by seed and foliar 24-epibrassinolide treatments. Hortic. Sci. 47, 631-636.
57. El-Khallal, S. M. (2002). The influence of some phyto-growth regulators on the activity of antioxidant system in maize plant under water stressed conditions. Bull. Faculty Sci. Assiut. Univ. D Bot. 31, 183-197.
58. El-Khallal, S. M., Hathout, T. A., Ashour, A. E. R. A., and Kerrit, A. A. A. (2009). Brassinolide and salicylic acid induced antioxidant enzymes, hormonal balance and protein profile of maize plants grown under salt stress. Res. J. Agric. Biol. Sci. 5, 391-402.
59. El-Mashad, A., and Mohamed, H. (2012). Brassinolide alleviates salt stress and increases antioxidant activity of cowpea plants (Vigna sinensis). Protoplasma 249, 625-635. doi: 10.1007/s00709-011-0300-7
60. Fábregas, N., and Caño-Delgado, A. I. (2014). Turning on the microscope turret: a new view for the study of brassinosteroid signaling in plant development. Physiol. Plant 151, 172-183. doi: 10.1111/ppl.12130
61. Fariduddin, Q., Khalil, R. R. A. E., Mir, B. A., Yusuf, M., and Ahmad, A. (2013a). 24-Epibrassinolide regulates photosynthesis, antioxidant enzyme activities and proline content of Cucumis sativus under salt and/or copper stress. Environ. Monit. Assess. 185, 7845-7856. doi: 10.1007/s10661-013-3139-x
62. Fariduddin, Q., Khanam, S., Hasan, S. A., Ali, B., Hayat, S., and Ahmad, A. (2009a). Effect of 28-homobrassinolide on the drought stress-induced changes in photosynthesis and antioxidant system of Brassica juncea L. Acta Physiol. Plant 31, 889-897. doi: 10.1007/s11738-009-0302-7
63. Fariduddin, Q., Mir, B. A., Yusuf, M., and Ahmad, A. (2013b). Comparative roles of brassinosteroids and polyamines in salt stress tolerance. Acta Physiol. Plant 35, 2037-2053. doi: 10.1007/s11738-013-1263-4
64. Fariduddin, Q., Yusuf, M., Chalkoo, S., Hayat, S., and Ahmad, A. (2011). 28-Homobrassinolide improves growth and photosynthesis in Cucumis sativus L. through an enhanced antioxidant system in the presence of chilling stress. Photosynthetica 49, 55-64. doi: 10.1007/s11099-011-0022-2
65. Fariduddin, Q., Yusuf, M., Hayat, S., and Ahmad, A. (2009). Effect of 28-homobrassinolide on antioxidant capacity and photosynthesis in Brassica juncea plants exposed to different levels of copper. Environ. Exp. Bot. 66, 418-424. doi: 10.1016/j.envexpbot.2009.05.001
66. Farooq, M., Wahid, A., Lee, D. J., Cheema, S. A., and Aziz, T. (2010). Comparative time course action of the foliar applied glycinebetaine, salicylic acid, nitrous oxide, brassinosteroids and spermine in improving drought resistance of rice. J. Agron. Crop Sci. 196, 336-345. doi: 10.1111/j.1439-037X.2010.00422.x
67. Fedina, E. O. (2013). Effect of 24-epibrassinolide on pea protein tyrosine phosphorylation after salinity action. Russ. J. Plant Physiol. 60, 351-358. doi: 10.1134/S1021443713020088
68. Filova, A., Sytar, O., and Krivosudska, E. (2013). Effects of brassinosteroid on the induction of physiological changes in Helianthus annuus L. under copper stress. Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis 61, 623-629. doi: 10.11118/actaun201361030623
69. Friedrichsen, D. M., Nemhauser, J., Muramitsu, T., Maloof, N. N., Alonso, J., Ecker, J. R., et al. (2002). Three redundant brassinosteroids early response genes encode putative bHLH transcription factors required for normal growth. Genetics 162, 1445-1456.
70. Gao, W., Long, L., Zhu, L., Xu, L., Gao, W. H., Sun, L. Q., et al. (2013). Proteomic and virus-induced gene silencing (VIGS) analyses reveal that gossypol, brassinosteroids and jasmonic acid contribute to the resistance of cotton to Verticillium dahliae. Mol. Cell. Proteomics 12, 3690-3703. doi: 10.1074/mcp. M113.031013
71. Gill, S. S., and Tuteja, N. (2010). Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiol. Biochem. 48, 909-930. doi: 10.1016/j.plaphy.2010.08.016
72. Gomes, M. M. A., Torres Netto, A., Campostrini, E., Bressan-Smith, R., Zullo, M. A. T., Ferraz, T. M., et al. (2013). Brassinosteroid analogue affects the senescence in two papaya genotypes submitted to drought stress. Theor. Exp. Plant Physiol. 25, 186-195. doi: 10.1590/S2197-00252013000300003
73. González-Garcia, M. P., Vilarrasa-Blasi, J., Zhiponova, M., Divol, F., Mora-Garcia, S., Russinova., E., et al. (2011). Brassinosteroids control meristem size by promoting cell cycle progression in Arabidopsis roots. Development 138, 849-859. doi: 10.1242/dev.057331
74. Grove, M. D., Spencer, G. F., Rohwededer, W. K., Mandava, N. B., Worlet, J. F., and Warthen, J. C. Jr., et al. (1979). Brassinolide, a plant promoting steroid isolated from Brassica napus pollen. Nature 281, 121-124. doi: 10.1038/281216a0
75. Hartwig, T., Chuck, G. S., Fujioka, S., Klempien, A., Weizbauer, R., Potluri, D. P. V., et al. (2011). Brassinosteroid control of sex determination in maize. Proc. Natl. Acad. Sci. U.S.A. 108, 19814-19819. doi: 10.1073/pnas.1108359108
76. Hasan, S. A., Hayat, S., and Ahmad, A. (2011). Brassinosteroids protect photosynthetic machinery against the cadmium induced oxidative stress in two tomato cultivars. Chemosphere 84, 1446-1451. doi: 10.1016/j.chemosphere.2011.04.047
77. Hasan, S. A., Hayat, S., Ali, B., and Ahmad, A. (2008). 28-Homobrassinolide protects chickpea (Cicer arietinum) from cadmium toxicity by stimulating antioxidants. Environ. Pollut. 151, 60-66. doi: 10.1016/j.envpol.2007.03.006
78. Haubrick, L. L., and Assmann, S. M. (2006). Brassinosteroids and plant function: some clues, more puzzles. Plant Cell Environ. 29, 446-457. doi: 10.1111/j.1399-3054.2006.00708.x
79. Haubrick, L. L., Torsethaugen, G., and Assmann, S. M. (2006). Effect of bassinolide, alone and in concert with abscisic acid, on control of stomatal aperture and potassium currents of Vicia faba guard cell protoplasts. Physiol. Plant 128, 134-143. doi: 10.1111/j.1399-3054.2006.00708.x
80. Hayat, S., Ali, B., Hasan, S. A., and Ahmad, A. (2007). Brassinosteroid enhanced the level of antioxidants under cadmium stress in Brassica juncea. Environ. Exp. Bot. 60, 33-41. doi: 10.1016/j.envexpbot.2006.06.002
81. Hayat, S., Alyemeni, M., and Hasan, S. (2012). Foliar spray of brassinosteroid enhances yield and quality of Solanum lycopersicum under cadmium stress. Saudi J. Biol. Sci. 19, 325-335. doi: 10.1016/j.sjbs.2012.03.005
82. Hayat, S., Hasan, S. A., Hayat, Q., and Ahmad, A. (2010a). Brassinosteroids protect Lycopersicon esculentum from cadmium toxicity applied as shotgun approach. Protoplasma 239, 3-14. doi: 10.1007/s00709-009-0075-2
83. Hayat, S., Hasan, S. A., Yusuf, M., Hayat, Q., and Ahmad, A. (2010b). Effect of 28-homobrassinolide on photosynthesis, fluorescence and antioxidant system in the presence or absence of salinity and temperature in Vigna radiata. Environ. Exp. Bot. 69, 105-112. doi: 10.1016/j.envexpbot.2010.03.004
84. Hayat, S., Yadav, S., Ali, B., and Ahmad, A. (2010c). Interactive effect of nitric oxide and brassinosteroids on photosynthesis and the antioxidant system of Lycopersicon esculentum. Russ. J. Plant Physiol. 57, 212-221. doi: 10.1134/S1021443710020081
85. Ho, M. O. (2003). Brassinosteroids accelerate the rate of cell division in isolated protoplasts of Petunia hybrida. J. Plant Biotechnol. 5, 63-67.
86. Hu, C. H., Guo, J., Chen, L., Li, J. P., Qiao, L., and He, L. (2014). Influence of exogenous brassinolide on growth and resistance of maize seedling with preservative stress. Hunan Nongye Daxue Xuebao 40, 113-116. doi: 10.1007/s11099-010-0071-y
87. Hu, W. H., Wu, Y., Zeng, J. Z., He, L., and Zeng, Q. M. (2010). Chill-induced inhibition of photosynthesis was alleviated by 24-epibrassinolide pretreatment in cucumber during chilling and subsequent recovery. Photosynthetica 48, 537-544. doi: 10.1007/s11099-010-0071-y
88. Hu, W. H., Yan, X. H., Xiao, Y. A., Zeng, J. J., Qi, H. J., and Ogweno, J. O. (2013). 24-Epibrassinosteroid alleviate drought-induced inhibition of photosynthesis in Capsicum annuum. Sci. Hortic. 150, 232-237. doi: 10.1016/j.scienta.2012. 11.012
89. Huang, B., Chu, C. H., Chen, S. L., Juan, H. F., and Chen, Y. M. (2006). A proteomics study of the mung bean epicotyl regulated by brassinosteroids under conditions of chilling stress. Cell Mol. Biol. Lett. 11, 264-278. doi: 10.2478/s11658-006-0021-7
90. Iqbal, N., Trivellini, A., Masood, A., Ferrante, A., and Khan, N. A. (2013). Current understanding on ethylene signaling in plants: the influence of nutrient availability. Plant Physiol. Biochem. 73, 128-138. doi: 10.1016/j.plaphy.201 3.09.011
91. Janeczko, A., Gullner, G., Skoczowski, A., Dubert, F., and Barna, B. (2007). Effects of brassinosteroid infiltration prior to cold treatment on ion leakage and pigment contents in rape leaves. Biol. Plant 51, 355-358. doi: 10.1007/s10535-00 7-0072-2
92. Janeczko, A., Hura, K., Skoczowski, A., Idzik, I., Biesaga-Koscielniak, J., and Niemczyk, E. (2009). Temperature-dependent impact of 24-epibrassinolide on the fatty acid composition and sugar content in winter oilseed rape callus. Acta Physiol. Plant 31, 71-79. doi: 10.1007/s11738-008-0202-2
93. Janeczko, A., Koscielniak, J., Pilipowicz, M., Szarek-Lukaszewska, G., and Skoczowski, A. (2005). Protection of winter rape photosystem 2 by 24-epibrassinolide under cadmium stress. Photosynthetica 43, 293-298. doi: 10.1007/s11099-005-0048-4
94. Janeczko, A., Oklestkova, J., Pociecha, E., Koscielniak, J., and Mirek, M. (2011). Physiological effects and transport of 24-epibrassinolide in heat-stressed barley. Acta Physiol. Plant 33, 1249-1259. doi: 10.1007/s11738-010-0655-y
95. Jiang, Y. P., Huang, L. F., Cheng, F., Zhou, Y. H., Xia, X. J., Mao, W. H., et al. (2013). Brassinosteroids accelerate recovery of photosynthetic apparatus from cold stress by balancing the electron partitioning, carboxylation and redox homeostasis in cucumber. Physiol. Plant 148, 133-145. doi: 10.1111/j.1399-3054.2012.01696.x
96. Jin, H., Do, J., Shin, S. J., Choi, J. W., Choi, Y. I., Kim, W., et al. (2014). Exogenously applied 24-epi brassinolide reduces lignification and alters cell wall carbohydrate biosynthesis in the secondary xylem of Liriodendron tulipifera. Phytochemistry 101, 40-51. doi: 10.1016/j.phytochem.2014.02.003
97. Kang, Y. Y., Guo, S. R., Duan, J. J., and Hu, X. H. (2006). Effects of 24-epibrassinolide on antioxidant system and anaerobic respiratory enzyme activities in cucumber roots under hypoxia stress. J. Plant Physiol. Mol. Biol. 32, 535-542.
98. Kang, Y. Y., Guo, S. R., Li, J., and Duan, J. J. (2009). Effect of root applied 24-epibrassinolide on carbohydrate status and fermentative enzyme activities in cucumber (Cucumis sativus L.) seedlings under hypoxia. Plant Growth Regul. 57, 259-269. doi: 10.1007/s10725-008-9344-x
99. Kanwar, M. K., Bhardwaj, R., Arora, P., Chowdhary, S. P., Sharma, P., and Kumar, S. (2012). Plant steroid hormones produced under Ni stress are involved in the regulation of metal uptake and oxidative stress in Brassica juncea L. Chemosphere 86, 41-49. doi: 10.1016/j.chemosphere.2011.08.048
100. Kanwar, M. K., Bhardwaj, R., Chowdhary, S. P., Arora, P., Sharma, P., and Kumar, S. (2013). Isolation and characterization of 24-epibrassinolide from Brassica juncea L. and its effects on growth, Ni ion uptake, antioxidant defense of Brassica plants and in vitro cytotoxicity. Acta Physiol. Plant 35, 1351-1362. doi: 10.1007/s11738-012-1175-8
101. Kapoor, D., Rattan, A., Gautam, V., Kapoor, N., and Bharadwaj, R. (2014). 24-Epibrassinolide mediated photosynthetic pigments and antioxidative defense systems of radish seedling under cadmium and mercury stress. J. Stress Physiol. Biochem. 10, 110-121.
102. Karlidag, H., Yildirim, E., and Turan, M. (2011). Role of 24-epibrassinolide in mitigating the adverse effects of salt stress on stomatal conductance, membrane permeability, and leaf water content, ionic composition in salt stressed strawberry (Fragaria x ananassa). Sci. Hort. 130, 133-140. doi: 10.1016/j.scienta.2011.06.025
103. Kartal, G., Temel, A., Arican, E., and Gozukirmizi, N. (2009). Effects of brassinosteroids on barley root growth, antioxidant system and cell division. Plant Growth Regul. 58, 261-267. doi: 10.1007/s10725-009-9374-z
104. Khan, N. A., Nazar, R., Iqbal, N., and Anjum, N. A. (2012). Phytohormones and Abiotic Stress Tolerance in Plants. Berlin; Heidelberg: Springer Verlag.
105. Kroutil, M., Hejtmankova, A., and Lachman, J. (2010). Effect of spring wheat (Triticum aestivum L.) treatment with brassinosteroids on the content of cadmium and lead in plant aerial biomass and grain. Plant Soil Environ. 56, 43-50.
106. Kumar, M., Sirhindi, G., Bhardwaj, R., Kumar, S., and Jain, G. (2010). Effect of exogenous H2O2 on antioxidant enzymes of Brassica juncea L. seedlings in relation to 24-epibrassinolide under chilling stress. Indian J. Biochem. Biophys. 47, 378-382.
107. Kurepin, L. V., Joo, S. H., Kim, S. K., Pharis, R. P., and Back, T. G. (2012). Interaction of brassinosteroids with light quality and plant hormones in regulating shoot growth of young sunflower and Arabidopsis seedlings. J. Plant Growth Regul. 31, 156-164. doi: 10.1007/s00344-011-9227-7
108. Kurepin, L. V., Qaderi, M. M., Back, T. G., Reid, D. M., and Pharis, R. P. (2008). A rapid effect of applied brassinolide on abscisic acid concentrations in Brassica napus leaf tissue subjected to short-term heat stress. Plant Growth Regul. 55, 165-167. doi: 10.1007/s10725-008-9276-5
109. Li, K. R., Wang, H. H., Han, G., Wang, Q. J., and Fan, J. (2008). Effects of brassinolide on the survival, growth and drought resistance of Robinia pseudoacacia seedlings under water-stress. New Forests 35, 255-266. doi: 10.1007/s11056-007-9075-2
110. Li, K. R., and Feng, C. H. (2011). Effects of brassinolide on drought resistance of Xanthoceras sorbifolia seedlings under water stress. Acta Physiol. Plant 33, 1293-1300. doi: 10.1007/s11738-010-0661-0
111. Li, L., and Van Staden, J. (1998). Effects of plant growth regulators on drought resistance of two maize cultivars. South Afr. J. Bot. 64, 116-120.
112. Li, L., and Van Staden, J. (1998). Effects of plant growth regulators on the antioxidant system in callus of two maize cultivars subjected to water stress. Plant Growth Regul. 24, 55-66.
113. Li, L., Van Staden, J., and Jager, A. K. (1998). Effects of plant growth regulators on the antioxidant system in seedlings of two maize cultivars subjected to water stress. Plant Growth Regul. 25, 81-87. doi: 10.1023/A:1010774725695
114. Li, Y. H., Liu, Y. J., Xu, X. L., Jin, M., An, L. Z., and Zhang, H. (2012b). Effect of 24-epibrassinolide on drought stress-induced changes in Chorispora bungeana. Biol. Plant 56, 192-196. doi: 10.1007/s10535-012-0041-2
115. Li, Z. Y., Xu, Z. S., He, G. Y., Yang, G. X., Chen, M., Li, L. C., et al. (2012a). A mutation in Arabidopsis BSK5 encoding a brassinosteroid-signaling kinase protein affects responses to salinity and abscisic acid. Biochem. Biophys. Res. Commun. 426, 522-527. doi: 10.1016/j.bbrc.2012.08.118
116. Liang, J., and Liang, Y. (2009). Effects of plant growth substances on water-logging resistance of oilseed rape seedling. Xinan Shifan Daxue Xuebao, Ziran Kexueban 34, 58-62.
117. Liu, H., Guo, T., Zhu, Y., Wang, C., and Kang, G. (2006). Effects of epi-brassinolide (epi-BR) application at anthesis on starch accumulation and activities of key enzymes in wheat grains. Acta Agron. Sinica 32, 924-930.
118. Liu, Y. J., Jiang, H. F., Zhao, Z. G., and An, L. Z. (2011). Abscisic acid is involved in brassinosteroids-induced chilling tolerance in the suspension cultured cells from Chorispora bungeana. J. Plant Physiol. 168, 853-862. doi: 10.1016/j.jplph.2010.09.020
119. Liu, Y. J., Zhao, Z. G., Si, J., Di, C. X., Han, J., and An, L. Z. (2009). Brassinosteroids alleviate chilling-induced oxidative damage by enhancing antioxidant defense system in suspension cultured cells of Chorispora bungeana. Plant Growth Regul. 59, 207-214. doi: 10.1007/s10725-009-9405-9
120. Lu, Z., Huang, M., Ge, D. P., Yang, Y. H., Cao, X. N., Qin, P., et al. (2003). Effect of brassinolide on callus growth and regeneration in Spartina patens (Poaceae). Plant Cell Tissue Org. Cult. 73, 87-89. doi: 10.1023/A:1022665210113
121. Lu, X. M., and Guo, S. R. (2013). Effects of brassinolide on the polyamines, ATPase activity and inorganic ion content in roots of cucumber seedlings under hypoxia stress. Shengtaixue Zazhi 32, 611-614.
122. Lu, X. M., and Yang, W. (2013). Alleviation effects of brassinolide on cucumber seedlings under NaCl stress. Ying Yong Sheng Tai Xue Bao 24, 1409-1414.
123. Lu, X., Chen, Y., Gong, W., and Chen, Y. (2006). Effect of brassinolide on the seedling growth and waterlogging resistance of soybean. Chinese Agric. Sci. Bull. 23, 37-38.
124. Ma, N., Liu, D., Zhang, C., Li, J., and Li, G. (2009). Regulation effect of exogenous hormones on growth and photosynthesis and yield of rapeseed (Brassica napus L.) after frozen. Zuowu Xuebao (Acta Agronomica Sinica). 35, 1336-1343. doi: 10.3724/SP.J.1006.2009.01336
125. Mahesh, B., Parshavaneni, B., Ramakrishna, B., and Rao, S. S. R. (2013). Effect of brassinosteroids on germination and seedling growth of radish (Raphanus sativus L.) under PEG-6000 induced water stress. Am. J. Plant Sci. 4, 2305-2313. doi: 10.4236/ajps.2013.412285
126. Malabadi, R. B., and Nataraja, K. (2007). 24-Epibrassinolide induces somatic embryogenesis in Pinus wallichiana A. B. Jacks. J. Plant Sci. 2, 171-178. doi: 10.3923/jps.2007.171.178
127. Manzano, S., Martínez, C., Megías, Z., Gómez, P., Garrido, D., and Jamilena, M. (2011). The role of ethylene and brassinosteroids in the control of sex expression and flower development in Cucurbita pepo. Plant Growth Regul. 65, 213-221. doi: 10.1007/s10725-011-9589-7
128. Mazorra, L. M., Holton, N., Bishop, G. J., and Nunez, M. (2011). Heat shock response in tomato brassinosteroid mutants indicates that thermotolerance is independent of brassinosteroid homeostasis. Plant Physiol. Biochem. 49, 1420-1428. doi: 10.1016/j.plaphy.2011.09.005
129. Mazorra, L. M., Nunez, M., Hechavarria, M., Coll, F., and Sanchez-Blanco, M. J. (2002). Influence of brassinosteroids on antioxidant enzymes activity in tomato under different temperatures. Biol. Plant 45, 593-596. doi: 10.1023/A:1022390917656
130. Meudt, W. J., Thompson, M. T., and Bennett, H. W. (1983). Investigations on mechanism of the brassinosteroid response. III. Techniques for potential enhancement of crop production. Proc. Plant Growth Regul. Soc. Am. 10, 312-318.
131. Mitchell, J. W., Mandava, N. B., Worley, J. E., Plimmer, J. R., and Smith, M. V. (1970). Brassins: a family of plant hormones from rape pollen. Nature 225, 1065-1066. doi: 10.1038/2251065a0
132. Mousavi, E. A., Kalantari, K. M., and Jafari, S. R. (2009). Change of some osmolytes accumulation in water-stressed colza (Brassica napus L.) as affected by 24-epibrassinolide. Iranian J. Sci. Technol. Transac. A Sci. 33, 1-11.
133. Nagata, N., Asami, T., and Yoshida, S. (2001). Brassinozole, an inhibitor of brassinosteroids biosynthesis, inhibits development of secondary xylem in cress plants (Lepidium sativum). Plant Cell Biol. 42, 1006-1011. doi: 10.1093/pcp/pce122
134. Nakaya, M., Tsukaya, H., Murakami, N., and Kato, M. (2002). Brassinosteroids control the proliferation of leaf cells of Arabidopsis thaliana. Plant Cell Physiol. 43, 239-244. doi: 10.1093/pcp/pcf024
135. Nomura, T., Nakayama, N., Reid, J. B., Takeuchi, Y., and Yokota, T. (1997). Blockage of brassinosteroid biosynthesis and sensitivity cause dwarfism in Pisum sativum. Plant Physiol. 113, 31-37.
136. Nunez, M., Mazzafera, P., Mazorra, L. M., Siqueira, W. J., and Zullo, M. A. T. (2003). Influence of a brassinosteroid analog on antioxidant enzymes in rice grown in culture medium with NaCl. Biol. Plant 47, 67-70. doi: 10.1023/A:1027380831429
137. Ogweno, J. O., Hu, W. H., Song, X. S., Shi, K., Mao, W. H., Zhou, Y. H., et al. (2010). Photoinhibition-induced reduction in photosynthesis is alleviated by abscisic acid, cytokinin and brassinosteroid in detached tomato leaves. Plant Growth Regul. 60, 175-182. doi: 10.1007/s10725-009-9439-z
138. Ogweno, J. O., Song, X. S., Shi, K., Hu, W. H., Mao, W. H., Zhou, Y. H., et al. (2008). Brassinosteroids alleviate heat-induced inhibition of photosynthesis by increasing carboxylation efficiency and enhancing antioxidant systems in Lycopersicon esculentum. J. Plant Growth Regul. 27, 49-57. doi: 10.1007/s00344-007-9030-7
139. Özdemir, F., Bor, M., Demiral, T., and Türkan, I. (2004). Effects of 24-epibrassinolide on seed germination, seedling growth, lipid peroxidation, proline content and antioxidative system of rice (Oryza sativa L.) under salinity stress. Plant Growth Regul. 42, 203-211. doi: 10.1023/B:GROW.0000026509.25995.13
140. Peng, Z., Han, C., Yuan, L., Zhang, K., Huang, H., and Ren, C. (2011). Brassinosteroid enhances jasmonate-induced anthocyanin accumulation in Arabidopsis seedlings. J. Integ. Plant Biol. 53, 632-640. doi: 10.1111/j.1744-7909.2011.01042.x
141. Pokotylo, I. V., Kretynin, S. V., Khripach, V. A., Ruelland, E., Blume, Y. B., and Kravets, V. S. (2014). Influence of 24-epibrassinolide on lipid signalling and metabolism in Brassica napus. Plant Growth Regul. 73, 9-17. doi: 10.1007/s10725-013-9863-y
142. Rady, M. M. (2011). Effect of 24-epibrassinolide on growth, yield, antioxidant system and cadmium content of bean (Phaseolus vulgaris L.) plants under salinity and cadmium stress. Sci. Hortic. 129, 232-237. doi: 10.1016/j.scienta.2011.03.035
143. Rady, M. M., and Osman, A. S. (2012). Response of growth and antioxidativve system of heavy metal contaminated tomato plants 24-epibrassinolide. Afr. J. Agric. Res. 7, 3249-3254.
144. Raghu, K., Mahesh, K., Divya Sri, N., and Rao, S. S. R. (2014). Effect of brassinosteroids on the seed germination and seedling growth of radish (Raphanus sativus L) under arsenic toxicity stress. Intl. J. Dev. Res. 9, 1929-1933.
145. Ramakrishna, B., and Rao, S. S. R. (2013). Preliminary studies on the involvement of glutathione metabolism and redox status against zinc toxicity in radish seedlings by 28-homobrassinolide. Environ. Expt. Bot. 96, 52-58. doi: 10.1016/j.envexpbot.2013.08.003
146. Rao, S. S. R., Vardhini, B. V., Sujatha, E., and Anuradha, S. (2002). Brassinosteroids-new class of phytohormones. Curr. Sci. 82, 1239-1245.
147. Saka, H., Fujii, S., Imakawa, A. M., Kato, N., Watanabe, S., Nishizawa, T., et al. (2003). Effect of brassinolide applied at the meiosis and flowering stages on the levels of endogenous plant hormones during grain filling in rice plant (Oryza sativa L.). Plant Prod. Sci. 6, 36-42. doi: 10.1626/pps.6.36
148. Savaliya, D. D., Mandavia, C. K., and Mandavia, M. K. (2013). Role of brassinolide on enzyme activities in groundnut under water deficit stress. Indian J. Agric. Biochem. 26, 92-96. Shahbaz, M., and Ashraf, M. (2007). Influence of exogenous application of brassinosteroids on growth and mineral nutrients of wheat (Triticum aestivum L.) under saline conditions. Pak. J. Bot. 39, 513-522.
149. Shahbaz, M., and Ashraf, M. (2007). Influence of exogenous application of brassinosteroids on growth and mineral nutrients of wheat (Triticum aestivum L.) under saline conditions. Pak. J. Bot. 39, 513-522
150. Shahbaz, M., Ashraf, M., and Athar, H. U. R. (2008). Does exogenous application of 24-epibrassinolide ameliorate salt induced growth inhibition in wheat (Triticum aestivum L.)? Plant Growth Regul. 55, 51-64. doi: 10.1007/s10725-008-9262-y
151. Shahid, M. A., Pervez, M. A., Balal, R. M., Mattson, N. S., Rashid, A., Ahmad, R., et al. (2011). Brassinosteroid (24-epibrassinolide) enhances growth and alleviates the deleterious effects induced by salt stress in pea (Pisum sativum L.). Aust. J. Crop Sci. 5, 500-510.
152. Shang, Q., Song, S., Zhang, Z., and Guo, S. (2006). Exogenous brassinosteroid induced salt resistance of cucumber (Cucumis sativus L.) seedlings. Sci. Agric. Sinica 39, 1872-1877.
153. Sharma, I., Bhardwaj, R., and Pati, P. K. (2013a). Stress modulation response of 24-epibrassinolide against imidacloprid in an elite indica rice variety Pusa Basmati-1. Pesticide Biochem. Physiol. 105, 144-153. doi: 10.1016/j.pestbp.2013. 01.004
154. Sharma, I., Ching, E., Saini, S., Bhardwaj, R., and Pati, P. K. (2013b). Exogenous application of brassinosteroid offers tolerance to salinity by altering stress responses in rice variety Pusa Basmati-1. Plant Physiol. Biochem. 69, 17-26. doi: 10.1016/j.plaphy.2013.04.013
155. Sharma, I., Pati, P. K., and Bhardwaj, R. (2010). Regulation of growth and antioxidant enzyme activities by 28-homobrassinolide in seedlings of Raphanus sativus L. under cadmium stress. Indian J. Biochem. Biophys. 47, 172-177.
156. Sharma, I., Pati, P. K., and Bhardwaj, R. (2011a). Effect of 24-epibrassinolide on oxidative stress markers induced by nickel-ion in Raphanus sativus L. Acta Physiol. Plant 33, 1723-1735. doi: 10.1007/s11738-010-0709-1
157. Sharma, I., Pati, P. K., and Bhardwaj, R. (2011b). Effect of 28-homobrassinolide on antioxidant defence system in Raphanus sativus L. under chromium toxicity. Ecotoxicology 20, 862-874. doi: 10.1007/s10646-011-0650-0
158. Sharma, N., Hundal, G. S., Sharma, I., and Bharadwaj, R. (2012). Effect of 24-epibrssinolide on protein content and activities of glutathione-S-transferase and poly phenol oxidase in Raphanus sativus L. plants under cadmium and mercury metal stress. Terrest. Aquat. Toxicol. 6, 1-7.
159. Sharma, P., Bhardwaj, R., Arora, N., Arora, H. K., and Kumar, A. (2008). Effects of 28-homobrassinolide on nickel uptake, protein content and antioxidative defence system in Brassica juncea. Biol. Plant 52, 767-770. doi: 10.1007/s10535-008-0149-6
160. Singh, I., and Shono, M. (2005). Physiological and molecular effects of 24-epibrassinolide, a brassinosteroid on thermotolerance of tomato. Plant Growth Regul. 47, 111-119. doi: 10.1007/s10725-005-3252-0
161. Song, S., Liu, W., Guo, S., Shang, Q., and Zhang, Z. (2006). Salt resistance and its mechanism of cucumber under effects of exogenous chemical activator. Yingyong Shengtai Xuebao 17, 1871-1876.
162. Strnad, M., and Kohout, L. (2003). A simple brassinolide analogue 2a, 3a-dihydroxy-17b-(3-methyl butynyloxyl) 7-oxa-B-homo-5a androstan-6-one which induces bean second node splitting. Plant Growth Regul. 40, 39-47. doi: 10.1023/A:1023091918892
163. Suzuki, Y., Saso, K., Fujioka, S., Yoshida, S., Nitasaka, E., Nagata, S., et al. (2003). A dwarf mutant strain of Pharbitis nil, Uzukobito (kobito), has defective brassinosteroids biosynthesis. Plant J. 36, 401-410. doi: 10.1046/j.1365-313X.2003.01887.x
164. Tabur, S., and Demir, K. (2009). Cytogenetic response of 24-epibrassinolide on the root meristem cells of barley seeds under salinity. Plant Growth Regul. 58, 119-123. doi: 10.1007/s10725-008-9357-5
165. Talaat, N. B., and Shawky, B. T. (2013). 24-Epibrassinolide alleviates saltinduced inhibition of productivity by increasing nutrients and compatible solutes accumulation and enhancing antioxidant system in wheat (Triticum aestivum L.). Acta Physiol. Plant. 35, 729-740. doi: 10.1007/s11738-01 2-1113-9
166. Tao, Y. Z., Zheng, J., Xu, Z. M., Zhang, X. H., Zhang, K., and Wang, G. Y. (2004). Functional analysis of ZmDWF1, a maize homolog of Arabidopsis brassinosteroids biosynthetic DWF1/DIM gene. Plant Sci. 167, 743-751. doi: 10.1016/j.plantsci.2004.05.012
167. Upreti, K. K., and Murti, G. S. R. (2004). Effects of brassinosteroids on growth, nodulation, phytohormone content and nitrogenase activity in French bean under water stress. Biol. Plant 48, 407-411. doi: 10.1023/B:BIOP.0000041094.13342.1b
168. Vardhini, B. V. (2011). Studies on the effect of brassinolide on the antioxidative system of two varieties of sorghum grown in saline soils of Karaikal. Asian Aust. J. Plant Sci. Biotechnol. 5, 31-34.
169. Vardhini, B. V. (2012). Application of brassinolide mitigates daline stress of certain metabolites of sorghum grown in Karaikal. J. Phytol. 4, 1-3.
170. Vardhini, B. V. (2012). Effect of brassinolide on certain enzymes of sorghum plant grown in saline soils of Karaikal. J. Phytol. 4, 30-33.
171. Vardhini, B. V. (2013). Comparative study of Sorghum vulgare Pers. Grown in two experimental sites by brassinolide application at vegetative, flowering and grain filling stage. Proc. Andhra Pradesh Akad. Sci. 15, 75-79.
172. Vardhini, B. V. (2013). "Brassinosteroids role for amino acids, peptides and amines modulation in stressed plants-a review," in Plant Adaptation to Environmental Change: Significance of Amino Acids and their Derivatives, eds N. A. Anjum, S. S. Gill, and R. Gill (Wallingford, CT: CAB International), 300-316.
173. Vardhini, B. V., Anuradha, S., and Rao, S. S. R. (2006). Brassinosteroids-a great potential to improve crop productivity. Indian J. Plant Physiol.11, 1-12.
174. Vardhini, B. V., and Rao, S. S. R. (1998). Effect of brassinosteroids on growth, metabolite content and yield of Arachis hypogaea. Phytochemistry 48, 927-930. doi: 10.1016/S0031-9422(97)00710-3
175. Vardhini, B. V., and Rao, S. S. R. (1999). Effect of brassinosteroids on nodulation and nitrogenase activity in groundnut (Arachis hypogaea. L). Plant Growth Regul. 28, 165-167. doi: 10.1023/A:1006227417688
176. Vardhini, B. V., and Rao, S. S. R. (2002). Acceleration of ripening of tomato pericarp discs by brassinosteroids. Phytochemistry 61, 843-847. doi: 10.1016/S0031-9422(02)00223-6
177. Vardhini, B. V., and Rao, S. S. R. (2003). Amelioration of water stress by brassinosteroids on seed germination and seedling growth of three varieties of sorghum. Plant Growth Regul. 41, 25-31. doi: 10.1023/A:1027303518467
178. Vardhini, B. V., and Rao, S. S. R. (2003). Influence of brassinosteroids on growth, yield, metabolite content and enzyme activities of tomato (Lycopersicon esculentum Mill.). Proc. Natl. Acad. Sci. India B Biol. Sci. 73, 307-315.
179. Vardhini, B. V., and Rao, S. S. R. (2005). Influence of brassinosteroids on seed germination and seedling growth of sorghum under water stress. Indian J. Plant Physiol. 10, 381-385.
180. Vardhini, B. V., Rao, S. S. R., and Rao, K. V. N. (2008). "Effect of brassinolide on growth, yield, metabolite content and enzyme activities of tomato (Lycopersicon esculentum) Mill," in Recent Advances in Plant Biotechnology and its Applications, eds S. K. Ashwani Kumar and I. K. Sopory (New Delhi: International Publishing House Ltd.), 133-139.
181. Vogler, F., Schmalzl, C., Englhart, M., Bircheneder, M., and Sprunck, S. (2014). Brassinosteroids promote Arabidopsis pollen germination and growth. Plant Reproduction 27, 153-167. doi: 10.1007/s00497-014-0247-x
182. Wang, B., Zhang, J., Xia, X., and Zhang, W. H. (2011). Ameliorative effect of brassinosteroid and ethylene on germination of cucumber seeds in the presence of sodium chloride. Plant Growth Regul. 65, 407-413. doi: 10.1007/s10725-011-9595-9
183. Wang, H. H., Feng, T., Peng, X. X., Yan, M. L., Zhou, P. L., and Tang, X. K. (2009). Ameliorative effects of brassinosteroid on excess manganese-induced oxidative stress in Zea mays L. leaves. Agric. Sci. China 8, 1063-1074. doi: 10.1016/S1671-2927(08)60314-4
184. Wang, M., Jiang, W. J., and Yu, H. J. (2010). Effects of exogenous epibrassinolide on photosynthetic characteristics in tomato (Lycopersicon esculentum Mill) seedlings under weak light stress. J. Agric. Food Chem. 58, 3642-3645. doi: 10.1021/jf9033893
185. Wang, Q., Ding, T., Gao, L., Pang, J., and Yang, N. (2012). Effect of brassinolide on chilling injury of green bell pepper in storage. Sci. Hortic. 144, 195-200. doi: 10.1016/j.scienta.2012.07.018
186. Wang, X. H., Shu, C., Li, H. Y., Hu, X. Q., and Wang, Y. X. (2014). Effects of 0.01% brassinolide solution application on yield of rice and its resistance to autumn low-temperature damage. Acta Agriculturae Jiangxi 26, 36-38.
187. Weng, Q., Zhou, B. L., Yu, Y., and Fu, Y. W. (2007). Effects of exogenous ABA, BR, and ETH on changes of lycopene contents in fruit of tomato. Shenyang Nongye Daxue Xuebao 38, 784-787.
188. Xi, Z., Wang, Z., Fang, Y., Hu, Z., Hu, Y., Deng, M., et al. (2013). Effects of 24-epibrassinolide on antioxidation defense and osmoregulation systems of young grapevines (V. vinifera L.) under chilling stress. Plant Growth Regul. 71, 57-65. doi: 10.1007/s10725-013-9809-4
189. Xia, X. H., Huang, Y. Y., Wang, L., Huang, L. F., Yu, Y. L., Zhou, Y. H., et al. (2006). Pesticides-induced depression of photosynthesis was alleviated by 24-epibrassinolide pretreatment in Cucumis sativus L. Pesticide Biochem. Physiol. 86, 42-48. doi: 10.1016/j.pestbp.2006.01.005
190. Xia, X. J., Gao, C. J., Song, L. X., Zhou, Y. H., Shi, K., and Yu, J. Q. (2014). Role of H2O2 dynamics in brassinosteroid-induced stomatal closure and opening in Solanum lycopersicum. Plant Cell Environ. 37, 2036-2050. doi: 10.1111/pce.12275
191. Xia, X. J., Wang, Y. J., Zhau, Y. H., Tao, Y., Mao, W. H., Shi, K., et al. (2009a). Reactive oxygen species are involved in brassinosteroid-induced stress tolerance in cucumber. Plant Physiol. 150, 801-814. doi: 10.1104/pp.109. 138230
192. Xia, X. J., Zhang, Y., Wu, J. X., Wang, J. T., Zhou, Y. H., Shi, K., et al. (2009b). Brassinosteroids promote metabolism of pesticides in cucumber. J. Agric. Food Chem. 57, 8406-8413. doi: 10.1021/jf901915a
193. Xia, X. J., Zhou, Y. H., Ding, J., Shi, K., Asami, T., Chen, Z., et al. (2011). Induction of systemic stress tolerance by brassinosteroid in Cucumis sativus. New Phytol. 191, 706-720. doi: 10.1111/j.1469-8137.2011.03745.x
194. Yamamoto, R., Fujioka, S., Demura, T., Takatsuto, S., Yoshida, S., and Fukuda, H. (2001). Brassinosteroid levels increase drastically prior to morphogenesis of tracheary elements. Plant Physiol. 125, 556-563. doi: 10.1104/pp.125. 2.556
195. Yuan, G. F., Jia, C. G., Li, Z., Sun, B., Zhang, L. P., Liu, N., et al. (2010). Effect of brassinosteroids on drought resistance and abscisic acid concentration in tomato under water stress. Sci. Hortic. 126, 103-108. doi: 10.1016/j.scienta.2010.06.014
196. Yuan, L., Shu, S., Sun, J., Guo, S., and Tezuka, T. (2012). Effects of 24-epibrassinolide on the photosynthetic characteristics, antioxidant system, and chloroplast ultrastructure in Cucumis sativus L. under Ca(NO3)2 stress. Photosynthesis Res. 112, 205-214. doi: 10.1007/s11120-012-9774-1
197. Yusuf, M., Fariduddin, Q., and Ahmad, A. (2011a). 28-Homobrassinolide mitigates boron induced toxicity through enhanced antioxidant system in Vigna radiata plants. Chemosphere 85, 1574-1584. doi: 10.1016/j.chemosphere.2011. 08.004
198. Yusuf, M., Fariduddin, Q., and Ahmad, A. (2012). 24-Epibrassinolide modulates growth, nodulation, antioxidant system, and osmolyte in tolerant and sensitive varieties of Vigna radiata under different levels of nickel: a shotgun approach. Plant Physiol. Biochem. 57, 143-153. doi: 10.1016/j.plaphy.2012. 05.004
199. Yusuf, M., Fariduddin, Q., Ahmad, I., and Ahmad, A. (2014). Brassinosteroidmediated evaluation of antioxidant system and nitrogen metabolism in two contrasting cultivars of Vigna radiata under different levels of nickel. Physiol. Mol. Biol. Plant. 20, 449-460. doi: 10.1007/s12298-014-0259-x
200. Yusuf, M., Fariduddin, Q., Hayat, S., Hasan, S. A., and Ahmad, A. (2011b). Protective response of 28-homobrassinolide in cultivars of Triticum aestivum with different levels of nickel. Arch. Environ. Contam. Toxicol. 60, 68-76. doi: 10.1007/s00244-010-9535-0
201. Zadvornova, Y. V., Alekseichuk, G. N., Laman, N. A., Khripach, V. A., and Grut, S. (2005). Effect of brassinosteroids on activation of the cell cycle during germination of Brassica oleracea L. seeds. Doklady Natsional'noi Akademii Nauk Belarusi 49, 70-73.
202. Zhang, M. C., Zhai, Z. X., Tian, X. L., Duan, L. S., and Li, Z. H. (2008). Brassinolide alleviated the adverse effect of water deficits on photosynthesis and the antioxidant of soybean (Glycine max L.). Plant Growth Regul. 56, 257-264. doi: 10.1007/s10725-008-9305-4
203. Zhang, S., Hu, J., Zhang, Y., Xie, X. J., and Knapp, A. (2007). Seed priming with brassinolide improves lucerne (Medicago sativa L.) seed germination and seedling growth in relation to physiological changes under salinity stress. Aust. J. Agric. Res. 58, 811-815. doi: 10.1071/AR06253
204. Zhang, Y. P., Zhu, X. H., Ding, H. D., Yang, S. J., and Chen, Y. Y. (2013). Foliar application of 24-epibrassinolide alleviates high-temperature-induced inhibition of photosynthesis in seedlings of two melon cultivars. Photosynthetica 51, 341-349. doi: 10.1007/s11099-013-0031-4
205. Zhao, Y., Liang, Z. Y., and Yang, Y. J. (2013). Effects of exogenous brassinosteroid on Cd tolerance in Solatium nigrum seedlings. Zhongguo Shengtai Nongye Xuebao 21, 872-876.
206. Zhiponova, M. K., Vanhoutte, I., Boudolf, V., Betti, C., Dhondt, S., Coppens, F., Mylle, E., et al. (2013). Brassinosteroid production and signaling differentially control cell division and expansion in the leaf. New Phytol. 197, 490-502. doi: 10.1111/nph.12036
207. Zhou, A., Wang, H., Walker, J. C., and Li, J. (2004). BRL1, a leucine-rich repeat receptor-like protein kinase, is functionally redundant with BRI1 in regulating Arabidopsis brassinosteroid signaling. Plant J. 40, 399-409. doi: 10.1111/j.1365-313X.2004.02214.x