Document Type : Original Article

Authors

1 Assistant Professor of Plant Science, Department of Biology, Faculty of Science, Yazd University, Yazd, Iran.

2 Full Professor of Faculty of Biological Sciences, Kharazmi University, Tehran, Iran- Department of Biology, Science and Research Branch, Islamic Azad University, Tehran, Iran.

3 Assistant Professor of Agriculture and Biotechnology Research Institute, Karaj, Iran.

4 Assistant Professor of department of Plant Science, Faculty of Bioogical Sciences, Kharazmi University, Tehran, Iran.

Abstract

Introduction
Drought stress is a major factor that reduces growth, development and production of plants especially in generative phase. Among crop plants, wheat (Triticum aestivum), which often experiences water-shortage conditions, is an appealing study system because there are so many natural genotypes differing in drought tolerance. Wheat as well as the most cereals is especially threatened by water deficit during flowering and grain filling period in relation to decreasing yield potential (Khanna-Chop and Selote., 2007). Phytohormones play an important role in plants tolerance of abiotic stresses by mediating a wide range of adaptive responses although some of them act differently. Abscisic acid (ABA) and cytokinin (CK) are two antagonist hormones that play useful roles when stresses as drought appear. But, recent data implied that abscisic acid (ABA)/CK ratios in xylem sap are important for stress signaling. Therefore, this experiment was established to study and compare the influence of exogenous treatments of individual CK and ABA and also CK and ABA interaction on flag leaves relative water content (LRWC), chlorophyll florescence and gas exchange in related to yield were studied in Triticum aestivum cv. Pishgam (drought-tolerant) and cv.MV-17 (drought-sensitive) under irrigation and drought conditions after post-anthesis stage.
Materials and methods
The influence of CK, ABA and their combination CK/ABA on LRWC, photochemical efficiency and gas exchange in Triticum aestivum cv. Pishgam (drought-tolerant) and cv.MV-17 (drought-sensitive) flag leaves and yield under irrigation and drought conditions during the grain-filling period. Factorial experiment was based on based on split plot in a randomized complete block design with three replications. Chlorophyll fluorescence as photochemical efficiency was assayed by OS1-FL, a pulse modulated fluorometer (OptiScience Corporation, Tyngsboro, MA). Gas exchange traits were determined at flowering using a portable gas exchange measuring system (Li 6400, Li-Cor, USA). For yield assay, 60 plants were selected randomly to assess grain weight 1000 and grain yield per plant. One-way ANOVA was applied to determine the significance of the results between different treatments and then Duncan multiple range tests (p < 0.05).
Results and discussion
Drought stress decreased LRWC in Pishgam cv. more than MV-17, significantly. Among hormonal treatments, the interaction of CK and ABA increased LRWC in both cultivars, significantly more than other hormonal treatments during drought stress. Chlorophyll florescence results showed that ФPSII and qP decreased and NPQ increased significantly in sensitive cv. more than tolerant cv. when irrigation was interrupted. CK and ABA interaction increased Fv/Fm, ФPSII and NPQ more than other hormonal treatments. Also CK/ABA application caused the significant increment in qP in Pishgam cv. Stomatal conductance (gs), net photosynthesis (A), transpiration (E) and Mesophyll conductance (MC) decreased in both cultivars when drought occurred while sub-stomatal CO2 concentration (Ci) and photosynthetic water use efficiency (PWUE) increased. These changes were more significant in sensitive cv. than tolerant cv. CK and ABA interaction increased Ci, A and PWUE in both cv. more than other hormonal treatments, while ABA treatment was the most effective hormonal treatment in reduction of gs, E and MC under drought condition. Yield reduced in both cultivars when drought stress appeared. Yield reduction was more in sensitive cv. CK and ABA combination was the most effective treatment for increment of yield, especially in sensitive cv. under drought stress. More Physiologic damages and yield reduction under drought exposure in susceptible cultivars were reviewed in the previous study (Chaves et al., 2003). CK effects on chlorophyll stability, protein synthesis in chloroplast and ABA effects on reduction of oxidative damage in photosynthesis system under drought and salt stress are reasons for production efficiency in plants (Pospisilova, 2003; Pospisilova et al., 2005). َAlso assesment of correlation coefficient showed the most significant relation between NPQ, A and PWUE with 1000-grain weight and grain yield.

Conclusion
It seems that CK and ABA combination is the best treatment in related to improvement of chlorophyll florescence, photosynthesis and yield efficiency in both cultivars under drought stress. Also, MV-17 (sensitive cv.) could use the hormonal treatment better than Pishgam (tolerant cv.) to improve photosynthesis and yield stability.

Keywords

Ahmadi, A., Siosemardeh, A., 2005. Investigation on the physiological basis of grain yield and drought resistance in wheat: leaf photosynthetic rate, stomatal conductance and non stomatal limitation. International journal of Agriculture and Biology. 7, 807-811.
Alvarez, S., Marsh, E.L., Schroeder, S.G., Schachtman, D.P., 2008. Metabolomic and proteomic changes in the xylem sap of maize under drought. Plant, Cell and Environment. 31, 325-340.
Bilger, W., Bjorkman, O., 1990. Role of the xanthophyll cycle in photoprotection elucidated by measurements of light-induced absorbance changes, fluorescence and photosynthesis in leaves of Hedera canariensis. Photosynthesis Research. 25, 173-185.
Caruso, G., Cavaliere, C., Foglia, P., Gubbiotti, R., Samperi, R., Lagana, A., 2009. Analysis of drought responsive proteins in wheat (Triticum durum) by 2D-PAGE and MALDI-TOF mass spectrometry. Plant Science. 177, 570–576.
Chaves, M.M., Oliveira, M.M., 2004.  Mechanisms underlying plant resilience to water deficits: prospects for water-saving agriculture. Journal of Experimental Botany. 55, 2365-2384.
Chaves, M.M., Maroco, J.P., Periera, J.S., 2003. Understanding plant responses to drought from genes to the whole plant. Functional of Plant Biology. 30, 239-264.
Cornic, G., Fresneau, C., 2002. Photosynthetic carbon reduction and carbon oxidation cycles are the main electron sinks forPhotosystem II activity during a mild drought. Annals of Botany. 89, 887-894.
Dall’Osto, L., Fiore, A., Cazzaniga, S., Giuliano, G., Bassi, R., 2007. Different roles of a- and b-branch xanthophylls in photosystem assembly and photoprotection. Journal of Biological Chemistry. 282, 35056-36068.
 Demming-Adams, B., Adams, W.W., 1996. Xanthophyll cycle and light stress in nature: uniform response to excess direct sunlight among higher plant species. Planta.198, 460-470.
Fischer, R A., Rees, D., Sayre, K.D., Lu, Z.M., Candon, A.G., Saaverda, A.L., 1998. Wheat yield progress associated with higher stomatal conductance and photosynthetic rate, and cooler canopies. Crop Science. 38, 1467-1475.
Genty, B., Briantais, J.M., Baker, N.R., 1989. The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochimica et Biophysica Acta.990, 87-92.
Ghassemian, M., Lutes, J., Hur-Song, C., Lange, I., Chen, W., Zhu, T., Wang, X., Lange, M., 2008. Abscisic acid-induced modulation of metabolic and redox control pathways in Arabidopsis thaliana. Phytochemistry. 69, 2899-2911.
Khanna-Chop, R., Selote, D.S., 2007. Acclimation to drought stress generates oxidative stress tolerance in drought-resistant than-susceptible wheat cultivar under field conditions. Environmental and Experimental Botany. 60, 276-283.
Liang, J., Zhang, J., Cao, X., 2001. Singlet oxygen and photo-oxidative stress management in indica-japonica rice (Oryza sativa) hybrids. Physiologiae Plantarum. 112, 470-77. 
Lu, S., Su, W., Li, H., Guo, Z., 2009. Abscisic acid improves drought tolerance of triploid bermudagrass and involves H2O2- and NO-induced antioxidant enzyme activities. Plant, Cell and Environment. 31, 325-340.
Lu, C., Zhang, J., 1999. Effects of water stress on photosystem II photochemistry and its thermo stability in wheat plants. Journal of Experimental Botany.50, 1199-1206.
Mafakheri, A., Siosemardeh, A., Bahramnejad, B., Struik, P.C., Sohrabi, Y., 2010. Effect of drought stress on yield, proline and chlorophyll contents in three chickpea cultivars. Australian Journal of Crop Science. 4, 580-585.
Massacci, A., Nabiev, S.M., Pietrosanti, L., Nematov, S.K., Chernikova, T.N., Leipner, J., 2008. Response of the photosynthetic apparatus of cotton (Gossypium hirsutum) to the onset of drought stress under field conditions studied by gas-exchange analysis and chlorophyll fluorescence imaging. Plant Physiology and Biochemistry.46,189-195.
Pospisilova, J., Vagner, M., Malbeck, J., Travnickova, A., Batkova, P., 2005. Interactions between abscisic acid and cytokinins during water stress and subsequent rehydration. Biologia Plantarum. 49,533-540.
Pospisilova, J., 2003. Participation of phytohormones in the stomatal regulation of gas exchange during water stress. Biologia Plantarum. 46, 491-506.
Schachtman, D.P., Goodger, J.Q., 2008. Chemical root to shoot signaling under drought. Trends in Plant Science. 13, 281-287.
Subrahmanyam D., Subash, Y.S., Haris, A., Sikka, A.K., 2006. Influence of water stress on leaf photosynthetic characteristics in wheat cultivars differing in their susceptibility to drought.Photosynthetica.44, 125-129.
Terzi, R., Saglam A., Kutlu, N., Nar, H., Kadioglu, A., 2010. Impact of soil drought stress on photochemical efficiency of photosystem II and antioxidant enzyme activities of Phaseolus vulgaris cultivars. Turkish Journal of Botany. 34, 1-10.
Turner, N.C., 2004. Sustainable production of crops and pastures under drought in a Mediterranean environment. Annals of Applied Biology. 144, 139-147.
Turner, N.C., 1981. Techniques and experimental approaches for the measurement of plant water stress. Plant Soil. 58, 339-366.
Valliyodan, B., Nguyen, H.T., 2006. Understanding regulatory networks and engineering for enhanced drought tolerance in plants. Current Opinion in Plant Biology. 9,189-195.
wang, Z., Cao, W., Dai, T., Zhou, Q., 2001. Effects of exogenous hormones on floret development and grain set in wheat. Plant Growth Regulation. 40, 201-205.
Werner, T., Schmulling, T., 2009. Cytokinin action in plant development. Current Opinion in Plant Biology. 12, 527-538.
Yang, X., Chen, X., Ge, Q., Li, B., Tong, Y., Zhang, A., Li, Z., Kuang, T., Lu, C., 2006. Tolerance of photosynthesis to photoinhibition, high temperature and drought stress in flag leaves of wheat: A comparison between a hybridization line and its parents grown under field conditions. Plant Science. 171, 389-397. 
Zhang,Y.J., Zhao, C.J., Lui, L.Y., Wang, J.H., Wang, R.C., 2005. Chlorophyll fluorescence detected passively by difference reflectance spectra of wheat (Triticum aestivum L.) leaf. Journal of Integrative Plant Biology.47, 1228-1235.