Abstract
The effects of 6-benzyladenine (6-BA) on plant growth, photosynthetic gas exchange, chlorophyll fluorescence and antioxidant systems of eggplant (Solanum melongena L.) under salt stress were investigated. Eggplant seedlings were exposed to 90 mM NaCl with four levels of 6-BA (5, 10, 20 and 50 μM) for 10 days. 6-BA at lower concentrations increased chlorophyll concentration, the net photosynthetic rate (P N), stomatal conductance (g s), and transpiration rate (E), intercellular CO2 concentration (C i) and water use efficiency (WUE), as well as the quantum efficiency of PSII photochemistry (ΦPSII), photochemical quenching (q p), and decreased non-photochemical quenching (NPQ), while higher concentrations reduced the effects or even exacerbated the occurrence of photosynthetic capacity. The activities of antioxidant enzymes such as superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX) increased significantly during salt treatments, and induced the increase of the activities of these enzymes at certain concentrations of 6-BA. 6-BA also reduced significantly malonaldehyde (MDA) contents and O ·−2 production. It was concluded that 6-BA could alleviate the detrimental effects of salt stress on plant growth by increasing photosynthetic efficiency and enhancing antioxidant enzyme systems in leaves at a proper concentration and of the varying 6-BA concentrations used, the most effective concentration for promoting growth was 10 μM under saline conditions.




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- ABA:
-
Abscisic acid
- APX:
-
Ascorbate peroxidase
- 6-BA:
-
6-Benzyladenine
- CAT:
-
Catalase
- C i :
-
Intercellular CO2 concentration
- CTK:
-
Cytokinin
- F m :
-
Maximal fluorescence
- F v :
-
Variable fluorescence
- g s :
-
Stomatal conductance
- MDA:
-
Malondialdehyde
- NBT:
-
Nitrotetrazolium blue chloride
- NPQ:
-
Non-photochemical quenching
- O ·−2 :
-
Superoxide anion radical
- P N :
-
Net photosynthetic rate
- POD:
-
Peroxidase
- ΦPSII:
-
Relative quantum efficiency of PSII photochemistry
- ROS:
-
Reactive oxygen species
- q p :
-
Photochemical quenching coefficient
- SOD:
-
Superoxide dismutase
- E :
-
Transpiration rate
References
Arora N, Bhardwaj R, Sharma P, Arora HK (2008) 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
Baker NR (1991) A possible role for photosystem II in environmental perturbations of photosynthesis. Physiol Plant 81:563–570
Baker NR (2008) Chlorophyll fluorescence: a probe of photosynthesis in vivo. Annu Rev Plant Biol 59:89–113
Blackman PG, Davies WJ (1984) Modification of the CO2 responses of maize stomata by abscisic acid and by naturally occurring and synthetic cytokinins. J Exp Bot 35:174–179
Boucaud J, Ungar IA (1976) Hormonal control of germination under saline conditions of three halophyte taxa in genus Suaeda. Physiol Plant 36:197–200
Brathe A, Andresen G, Gundersen LL, Malterud KE, Rise F (2002) Antioxidant activity of synthetic cytokinin analogues: 6-alkynyl and 6-alkenylpurines as novel 15-lipoxygenase inhibitors. Bioorgan Med Chem 10:1581–1586
Calatayud A, Barreno E (2004) Response to ozone in two lettuce varieties on chlorophyll a fluorescence, photosynthetic pigments and lipid peroxidation. Plant Physiol Biochem 42:549–555
Chakrabarti N, Mukherji S (2003) Alleviation of NaCl stress by pretreatment with phytohormones in Vigna radiata. Biol Plantarum 46:589–594
Chernyad’ev II (2009) The protective action of cytokinins on the photosynthetic machinery and productivity of plants under stress (review). App Biochem Microbiol 45:351–362
Dai QL, Chen CH, Feng B, Liu TT, Tian X, Gong YY, Sun YK, Wang J, Du SZ (2009) Effects of different NaCl concentration on the antioxidant enzymes in oilseed rape (Brassica napus L.) seedlings. Plant Growth Regul 59:273–278
Das C, Sengupta T, Chattopadhyay S, Setua M, Das NK, Saratchandra B (2002) Involvement of kinetin and spermidine in controlling salinity stress in mulberry (Morus alba L. cv. S1). Acta Physiol Plant 24:53–57
Davies PJ (2010) The plant hormones: their nature, occurrence, and functions. Plant Hormones A 1–15
Davies WJ, Zhang JH (1991) Root signal and the regulation of growth and development of plants in drying soil. Ann Rev Plant Physiol Plant Mol Biol 42:55–76
Dong HZ, Niu YH, Kong XQ, Luo Z (2009) Effects of early-fruit removal on endogenous cytokinins and abscisic acid in relation to leaf senescence in cotton. Plant Growth Regul 59:93–101
Dubey RS (2005) Photosynthesis in plants under stressful conditions. In: Pessarakli M (ed) Hand book photosynthesis, 2nd edn. CRC Press, Taylor and Francis Group, New York, pp 717–737
Durner J, Klessing DF (1996) Salicylic acid is a modulator of tobacco and mammalian catalases. J Biol Chem 271:28492–28502
Fadzilla NM, Finch RP, Burdon RH (1997) Salinity, oxidative stress and antioxidant responses in shoot cultures of rice. J Exp Bot 48:325–331
FAO (2000) Global network on integrated soil management for sustainable use of salt-affected soils. Available on: http://www.fao.org/ag/AGL/agll/spush/intro.htm
Gadallah MAA (1999) Effects of kinetin on growth, grain yield and some mineral elements in wheat plants growing under excess salinity and oxygen deficiency. Plant Growth Regul 27:63–74
Gan SS, Amasino RM (1995) Inhibition of leaf senescence by autoregulated production of cytokinin. Science 270:1986–1988
Genty B, Briatais JM, Baker NR (1989) The relationships between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochim Biophys Acta 990:87–92
Giannopolitis CN, Ries SK (1977) Superoxide dismutases. I. Occurrence in higher plants. Plant Physiol 59:309–314
Hammerschmidt R, Nuckles EM, Kuc J (1982) Association of enhanced peroxidase activity with induced systemic resistance of cucumber to Colletotrchum lagenarium. Physiol Plant Pathol 20:73–82
Hare PD, van Staden J (1997) The molecular basis of cytokinin action. Plant Growth Regul 23:41–78
Heath RL, Packer L (1968) Photoperoxidation in isolated chloroplasts. I. Kinetics and stoichiometry of fatty acid peroxidation. Arch Biochem Biophys 125:189–198
Hernandez JA, Almansa MS (2002) Short-term effects of salt stress on antioxidant systems and leaf water relations of pea leaves. Physiol Plant 115:251–257
Hernandez JA, Corpass FJ, Gomez M, del Rio LA, Sevilla F (1993) Salt-induced oxidative stress mediated by active oxygen species in pen leaf mitochondria. Physiol Plant 89:103–110
Heuer B, Meiri A, Shalevet J (1986) Salt tolerance of eggplant. Plant Soil 95:9–13
Horton P, Ruban AV, Walters RG (1996) Regulation of light harvesting in green plants. Annu Rev Plant Physiol Plant Mol Biol 47:655–684
Iqbal M, Ashraf M (2005) Presowing seed treatment with cytokinins and its effect on growth, photosynthetic rate, ionic levels and yield of two wheat cultivars differing in salt tolerance. J Integ Plant Biol 47:1315–1325
Iqbal M, Ashraf M, Jamil A (2006a) Seed enhancement with cytokinins: changes in growth and grain yield in salt stressed wheat plants. Plant Growth Regul 50:29–39
Iqbal M, Ashraf M, Jamil A, Rehman S (2006b) Does seed priming induce changes in the levels of some endogenous plant hormones in hexaploid wheat plants under salt stress? J Integ Plant Biol 48:181–189
Itai C, Vaadia Y (1965) Kinetin like activity in root exudate of water stressed sunflower plants. Physiol Plant 18:941–944
Javid MG, Sorooshzadeh A, Moradi F, Sanavy SAMM, Allahdadi I (2011) The role of phytohormones in alleviating salt stress in crop plants. Aust J Crop Sci 5:726–734
Jung S, Kim JS, Cho KY, Tae GS, Kang BG (2000) Antioxidant responses of cucumber (Cucumis sativus) to photoinhibition and oxidative stress induced by norflurazon under high and low PPFDs. Plant Sci 153:145–154
Kirkham MB, Gardner WR, Gerloff GC (1974) Internal water status of kinetin-treated, salt-stressed plants. Plant Physiol 53:241–243
Klaus A, Hirt H (2004) Reactive oxygen species: metabolism, oxidative stress and signal transduction. Ann Rev Plant Biol 55:373–399
Kudoyarova GR, Vysotskaya LB, Cherkozyanova A, Dodd IC (2007) Effect of partial rootzone drying on the concentration of zeatintype cytokinins in tomato (Solanum lycopersicum L.) xylem sap and leaves. J Exp Bot 58:161–168
Kuiper D, Schuit J, Kuiper PJC (1990) Actual cytokinin concentrations in plant tissue as an indicator for salt resistance in cereals. Plant Soil 123:243–250
Liu X, Huang B, Banowetz G (2002) Cytokinin effects on creeping bentgrass response to heat stress: 1. Shoot and root growth. Crop Sci 42:457–465
Lu C, Vonshak A (2002) Effects of salinity stress on photosystem II function in cyanobacterial Spirulina platensis cells. Physiol Plant 114:405–413
Masood A, Shah NA, Zeeshan M, Abraham G (2006) Differential response of antioxidant enzymes to salinity stress in two varieties of Azolla (Azolla pinnata and Azolla filiculoides). Environ Exp Bot 58:216–222
Maxwell K, Johnson GN (2000) Chlorophyll fluorescence—a practical guide. J Exp Bot 51:659–668
McCord JM (2000) The evolution of free radicals and oxidative stress. Am J Med 108:652–659
Merewitz EB, Gianfagna T, Huang B (2011) Protein accumulation in leaves and roots associated with improved drought tolerance in creeping bentgrass expressing an ipt gene for cytokinin synthesis. J Exp Bot 62:5311–5333
Mittler R (2002) Oxidative stress, antioxidants and stress tolerance. Trends Plant Sci 7:405–410
Moradi F, Ismail AM (2007) Responses of photosynthesis, chlorophyll fluorescence and ROS-scavenging systems to salt stress during seedling and reproductive stages in rice. Ann Bot 99:1161–1173
Naeem MS, Jin ZL, Wan GL, Liu D, Liu HB, Yoneyama K, Zhou WJ (2010) 5-Aminolevulinic acid improves photosynthetic gas exchange capacity and ion uptake under salinity stress in oilseed rape (Brassica napus L.). Plant Soil 332:405–415
Nakano Y, Asada K (1981) Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant Cell Physiol 22:867–880
Ogweno JO, Hu WH, Song XS, Shi K, Mao WH, Zhou YH, Yu JQ (2010) Photoinhibition-induced reduction in photosynthesis is alleviated by abscisic acid, cytokinin and brassinosteroid in detached tomato leaves. Plant Growth Regul 60:175–182
Palus KP, Thompson JE (1982) Effects of cytokinins and antioxidants on the susceptibility of membranes to ozone damage. Plant Cell Physiol 23:821–832
Pogany M, Elstner EF, Barna B (2003) Cytokinin gene introduction confers tobacco necrosis virus resistance and higher antioxidant levels in tobacco. Free Radical Res 37:15–16
Rashotte AM, Chae HS, Maxwell BB, Kieber JJ (2005) The interaction of cytokinin with other signals. Physiol Plant 123:184–194
Rivero RM, Kojima M, Gepstein A, Sakakibara H, Mittler R, Gepstein S, Blumwald E (2007) Delayed leaf senescence induces extreme drought tolerance in a flowering plant. Proc Natl Acad Sci USA 104:19631–19636
Rivero RM, Shulaev V, Blumwald E (2009) Cytokinin-dependent photorespiration and the protection of photosynthesis during water deficit. Plant Physiol 150:1530–1540
Rohacek K (2002) Chlorophyll fluorescence parameters: the definitions, photosynthetic meaning, and mutual relationships. Photosynthetica 40:13–29
Saied AS, Keutgen N, Noga G (2003) Effects of NaCl stress on leaf growth, photosynthesis and ionic contents of strawberry cvs Elsanta’ and ‘Korona’. In: Pardossi A, Serra G, Tognoni F (eds) International symposium on managing greenhouse crops in saline environment, International Society of Hortic Sci, Pisa, Italy, pp 67–73
Shabala S, Cuin TA (2008) Potassium transport and plant salt tolerance. Physiol Plant 133:651–669
Sharma N, Gupta NK, Gupta S, Hasegawa H (2005) Effect of NaCl salinity on photosynthetic rate, transpiration rate, and oxidative stress tolerance in contrasting wheat genotypes. Photosynthetica 43:609–613
Shigeoka S, Ishikawa T, Tamoi M, Miyagawa Y, Takeda T, Yabuta Y, Yoshimura K (2002) Regulation and function of ascorbate peroxidase isoenzymes. J Exp Bot 53:1305–1319
Smirnoff N (1993) The role of active oxygen in the response of plants to water deficit and desiccation. New Phytol 125:27–58
Stepien P, Klobus G (2006) Water relations and photosynthesis in Cucumis sativus L. leaves under salt stress. Biol Plant 50:610–616
Takei K, Takahashi T, Sugiyama T, Yamaya T, Sakakibara H (2002) Multiple routes communicating nitrogen availability from roots to shoots: signal transduction pathway mediated by cytokinin. J Exp Bot 53:971–977
Thomas TH (1992) Some reflections on the relationship between endogenous hormones and light-mediated seed dormancy. Plant Growth Regul 11:239–248
van Kooten O, Snel J (1990) The use of chlorophyll fluorescence nomenclature in plant stress physiology. Photosynth Res 25:147–150
Walker MA, Dumbroff EB (1981) Effects of salt stress on abscisic acid and cytokinin levels in tomato. ZPfl anzenphysiol 101:461–470
Wang AG, Luo GH (1990) Quantitative relation between the reaction of hydroxylamine and superoxide anion radicals in plants. Plant Physiol Commun 6:55–57
Wang W, Vinocur B, Altman A (2003) Plant responses to drought, salinity and extreme temperatures: towards genetic engineering for stress tolerance. Planta 218:1–14
Willits DH, Peet MM (2001) Measurement of chlorophyll fluorescence as a heat stress indicator in tomato: laboratory and greenhouse comparisons. J Am Soc Hort Sci 126:188–194
Wu FB, Wu LH, Xu FH (1998) Chlorophyll meter to predict nitrogen sidedress requirements for short-season cotton (Gossypium hirsutum L.). Field Crops Res 56:309–314
Wu XX, Zhang YP, Zha DS (2010) Effect of NaCl stress on growth and photosynthetic characteristics of eggplant seedlings. Acta Agricu Zhejiangensis 22:193–197
Xia JR, Li YJ, Zou DH (2004) Effects of salinity stress on PSII in Ulva lactuca as probed by chlorophyll fluorescence measurements. Aquat Bot 80:129–137
Yamaguchi T, Blumwald E (2005) Developing salt-tolerant crop plants: challenges and opportunities. Trends Plant Sci 10:615–620
Yu JQ, Huang LF, Hu WH, Zhou YH, Mao WH, Ye SF, Nogues S (2004) A role for brassinosteroids in the regulation of photosynthesis in Cucumis sativus. J Exp Bot 55:1135–1143
Zahir ZA, Asghar HN, Arshad M (2001) Cytokinin and its precursors for improving growth and yield of rice. Soil Biol Biochem 33:405–408
Zhang RH, Li J, Guo SR, Takafumi Tezuka (2009) Effects of exogenous putrescine on gas-exchange characteristics and chlorophyll fluorescence of NaCl-stressed cucumber seedlings Photosynth Res 100:155–162
Zhu Z, Wei G, Li J, Qian Q, Yu J (2006) Silicon alleviates salt stress and increases antioxidant enzymes activity in leaves of salt-stressed cucumber (Cucumis sativus L.). Plant Sci 167:527–533
Acknowledgments
This work was supported by the Natural Science Foundation of Shanghai (No. ZR1426800) and Shanghai Municipal Committee of Agriculture (2009-2-1).
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Communicated by Z.-L. Zhang.
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Wu, X., Zhu, Z., Li, X. et al. Effects of cytokinin on photosynthetic gas exchange, chlorophyll fluorescence parameters and antioxidative system in seedlings of eggplant (Solanum melongena L.) under salinity stress. Acta Physiol Plant 34, 2105–2114 (2012). https://doi.org/10.1007/s11738-012-1010-2
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DOI: https://doi.org/10.1007/s11738-012-1010-2


