Document Type : Original Article

Authors

1 Organization of Agriculture-Jahad of Isfahan, the Ministry of Agriculture-Jahad, Isfahan, Iran

2 Department of Plant Production and Genetics, Khuzestan Agricultural Sciences and Natural Resources University, Mollasani, Khuzestan, Iran

3 Professor, International Maize and Wheat Improvement Center (CIMMYT), Karaj, Iran

Abstract

Introduction
Heat stress during reproductive development is the main limitation in the production of wheat in most of the wheat fields in the world. It is important to recognize the physiological and molecular mechanisms associated with heat tolerance and the detection of screening methods in improving plants to tolerate heat. In Iran data from the effect of short-term heat wave on yield and physiological mechanisms do not exist under field conditions. The objective of this study was to determine the effects of short-term heat stress near flowering and early grain filling on grain yield and physiological parameters.

Materials and methods
This research was conducted at research farm of Khuzestan Agricultural Sciences and Natural Resources University, located 35 km northeast of Ahwaz, in 2014. Four wheat genotypes (Chamran. Maroon, Arvand and Atrak) were exposed to heat stress (maximum 35 °C) for a three-day in the field with a portable heat chamber at two different stages, near flowering (H1) and early grain set (H2). Chlorophyll content was measured using manual chlorophyll. Stomatal conductance was performed from the top three leaves of selected plants. Chlorophyll concentration was calculated using the Arnon method (1949). Cell membrane thermostability (CMTS) were calculated using the following equation: CMTS (%) = [1- (EC1 / EC2)] × 100
where EC1 and EC2 are the primary electrical conductivity (before the autoclave) and the secondary (after the autoclave), respectively. Proline was maesured using Bates method (1973). Ascorbate peroxidase (APX) activity was measured using Nakano and Asada method (1987). Catalase (CAT) activity was measured using Aebi method (1983), peroxidase activity (POX) was measured using Chance and Maehly method (1955), Superoxide dismutase was measured using Biochamp and Fridovich method (1971). Malon de aldehyde (MDA) was measured using Hess and Packer (1969). All calculations were performed using the SAS-9.4 statistical software. Factor analysis was done by using principal component analysis and Varimax rotation on the temporary factor.

Results and discussion
The results of analysis of variance showed that there were significant differences between heat stress levels and the traits of genotypes. Short-term heat stress (H1 or H2) decreased the average grain yield of Chamran, Maroon, Arvand and Atrak genotypes by 19.6, 18.6, 17.8 and 11.2 percent. Heat stress significantly caused to reduce water potential, chlorophyll fluorescence, leaf relative water content and cell membrane thermostability whereas stomatal conductance, chlorophyll destruction rate and proline content increased. Although there was no significant effect of short term heat stress on total chlorophyll concentration in this experiment, there was a negative correlation between grain yield and total chlorophyll concentration (r = -0.67 in H1 and r = -0.77 in H2). The activity of catalase, peroxidase and superoxide dismutase increased significantly in response to H1 or H2. The results of the factor analysis showed that four factors explained 86.7% and three factors explained 86.4% the variance among varieties in H1 and H2, respectively. Atrek, Chamran and Arvand varieties tolerated heat stress than Maroon variety by more photosynthesis persistency, higher metabolite content and more enzyme defense mechanism. It could be argued that cultivars with slower rate of leaf senescence after heat exposure and more enzymatic protection could be more tolerant to heat stress.

Keywords

Aebi, H.E. 1983. Catalase. In: Bergmeyer H.U., Bergmeyer J., Grabi M. (eds.), Methods of Enzymatic Analysis. Third ed. Vol. 3. pp. 273-282, VCH Verlagsgesellschaft mbH, Germany.
Allakhverdive, S.I., Kreslavski, V.D., Klimov, V.V., Los, D.A., Carpentier, R., Mohanty, P., 2008. Heat Stress: An overview of molecular responses in photosynthesis. Photosynthesis Research. 98, 541-550.
Arnon, D.I., 1949. Copper enzymes in isolated chloroplasts. Polyphenoloxidase in Beta vulgaris. Plant Physiology. 24, 1-15.
Atteya, A.M., 2003. Alteration of water relations and yield of corn genotypes in response to drought stress. Journal of Plant Physiology. 29, 63-76.
Baker, N.R., Rosenquist, E., 2004. Application of chlorophyll fluorescence can improve crop production strategies: An examination of future possibilities. Journal of Experimental Botany. 55, 1607-1627.
Bates, L.S., Waldren, R.P., Teare, I.D., 1973. Rapid determination of free proline for water-stress studies. Plant and Soil. 39, 205-207.
Beachamp, C., Fridovich, F., 1971. Superoxide dismutase: cadmium assay and an assay applicable to acrylamide gels. Annual Biochemistry. 44, 276-27.
Blum, A., Mayer, J., Golan, G., 1988. The effect of grain number per ear (sink size) on source activity and its water relations in wheat. Journals Experiment of Botany. 39, 106 – 114.
Chance, B., Maehly, A.C., 1955. Assay of catalase and peroxidase. Methods in enzymology. 2, 764-775.
Farooq, M., Bramley, H., Palta, J.A., Siddique, H.M., 2011. Heat stress in wheat during reproductive and grain-filling phases. Critical Reviews in Plant Sciences. 30, 1–17.
Gibson, L.R., Paulsen, G.M., 1999. Yield components of wheat grown under high temperature stress during reproductive growth. Crop Science. 39, 1841–1846.
Harris, K., Subudhi, P.K., Borrell, A., Jordan, D., Rosenow, D., Nguyen, H.T., Klein, P., Klein, R., Mullet, J., 2007. Sorghum stay-green QTL individually reduce post-flowering drought-induced leaf senescence. Journal of Experimental Botany. 58, 327–338.
Havaux, M., Tardy, F., 1999. Loss of chlorophyll with limited reduction of photosynthesis as an adaptive response of Syrian barley landraces to high drought and heat stress. Australian Journal of Physiology. 26, 569-578.
Hays, D.B., Do, J.H., Mason, R.E., Morgan, G., and Finlayson, S.A., 2007. Heat stress induced ethylene production in developing wheat grains induces kernel abortion and increased maturation in a susceptible cultivar. Plant Science. 172, 1113–1123.
Heath, R.L., Packer, L., 1969. Photoperoxidation in isolated chloroplast. I. Kinetics and stoichiometry of fatty acid peroxidation. Archives of Biochemistry and Biophysics. 125, 189-198.
Jalal-Kamali, M.R., Duveiller, E., 2008.Wheat Production and Research in Iran: A Success Story. In: Raynolds, M.P., Pietragalla, J., Braun, H.J. (eds.). Proceeding of the international symposium on wheat yield potential: Challenges to international wheat breeding. pp. 54-58. CIMMYT, D.F. Mexico. 2008.
Jubany-Mari, T., Munne-Bosch, S., Alegre, L., 2010. Redox regulation of water stress responses in field-grown plants. Role of hydrogen peroxide and ascorbate. Plant Physiology and Biochemistry. 48, 351-358.
Kolchevskii, K.G., Kocharyan, N.I., Koroleva, Q.Y., 1990. Effect of salinity on photosynthetic characteristic and ion accumulation in C3 and C4 plant of Ararat plain. Photosynthetica. 31, 277-282.
Liu, C.M., Zhang, J.H., 2000. Heat–induced multiple effects on PSII in wheat plants. Journal of Plant Physiology. 156, 259-265.
Lopes, M.S., Reynolds, M.P., Jalal-Kamali, M.R., Moussa, M., Feltaous, Y., Tahir, I.S.A., Barma, N., Vargas, M., Mannes, Y., Baum, M., 2012. The yield correlations of selectable physiological traits in a population of advanced spring wheat lines grown in warm and drought environments. Field Crops Research. 128, 129–136.
Modarresi, M., Mohammadi, V., Zali, A., Mardi, M. 2010. Response of wheat yield and yield related traits to high temperature. Cereal Research Communications. 38, 23-31.
Mojtabaei-Zamani, M. 2012. Investigation of some physiological mechanisms of heat tolerance during grain filling period in wheat. PhD dissertation, Faculty of Agriculture, Shahid Chamran University of Ahvaz, Iran. [In Persian with English Summary].
Morant-Manceau, A., Pradier, E., Tremblin, G. 2004. Osmotic adjustment, gas exchanges and chlorophyll fluorescence of a hexaploid triticale and its parental species salt stress. Journal of Plant Physiology. 169, 25-33.
Moshattati, A., Alami-Saied, K., Siadat, S.A., Bakhshandeh, A.M., Jalal-Kamali, M.R., 2010. Evaluation of terminal heat stress tolerance in spring bread wheat cultivars in Ahwaz conditions. Iran Journal Crop Science. 12, 85-99. [In Persian with English Summary].
Nakano, Y., Asada, K., 1987. Purification of ascorbate peroxidase in spinach chloroplasts; its inactivation in ascorbate-depleted medium and reactivation by monodehydroascorbate radical. Plant and Cell Physiology. 28, 131-140.
Pessarakli, M., 1999. Hand book of Plant and Crop Stress. Marcel Dekker Incorporation, 697P.
Reynolds, M.P., Balota, M., Delgado, M.I.B., Amani, I., Fischer, R.A., 1994. Physiological and morphological traits associated with spring wheat yield under hot, irrigated conditions. Australian Journal of Plant Physiology. 21, 717–730.
Reynolds, M.P., Nagarajan, S., Razzaque, M.A., Ageeb, O.A.A., 2001. Heat tolerance. In: Reynolds, M.P., Ortiz-Monasterio, J.I., McNab, A. (eds.), Application of Physiology in Wheat Breeding. Mexico, D. F. CIMMYT.
Reynolds, M.P., Singh, R.P., Ibrahim, A., Ageeb, O.A.A., Larque-Saavedra, A., Quick, J.S., 1998. Evaluating physiological traits to complement empirical selection for wheat in warm environments. Euphytica. 100, 85–94.
Roshanfekr-Dezfuli, H., Nabipour, M., Moradi, F., Mesgarbashi, M., 2011. Effect of temperature change on stomatal conductance and chlorophyll concentration in wheat. Crop Production (Scientific Journal of Agriculture), 34, 39-52. [In Persian with English Summary].
Sairam, R.K., Srivastava, G.C., Saxena, D.C. 2000. Increased antioxidant activity under elevated temperatures: a mechanism of heat stress tolerance in wheat genotypes. Biologyca Plantarum. 43, 245–251.
Schonfeld, M.A., Johnson, R.C., Carver, B.F., Mornhinwag, D.W. 1988.Water relations in winter wheat as drought resistance indicators. Crop Science: 28, 526-531.
Showler, A.T., Castro, B.A., 2010. Influence of drought stress on Mexican rice borer (Lepidoptera: Crambidae) oviposition preference in sugarcane. Crop Protection. 28, 722-727.
Swidzinski, J.A., Leaver, C.J., Sweetlove, L.J., 2004. A proteomic analysis of plant programmed cell death. Photochemistry, 65, 1829-1838
Talukder, A.S.M.H.M., Gill, G.S., McDonald, G.K., Hayman, P.T., Alexander, B.M., 2010. Field evaluation of sensitivity of wheat to high temperature stress near flowering and early grain set. In: Dove, H., Culvenor, R.A. (eds.), Food Security from Sustainable Agriculture. Proceedings of the 15th Australian Agronomy Conference. Lincoln, New Zealand.
Teskey, R., Wertin, T., Bauweraerts, I., Ameye, M., Mcguire, M.A., Steppe, K. 2015. Responses of tree species to heat waves and extreme heat events. Plant, Cell and Environment. 38, 1699–1712.
Turkan, I., Bor, M., Ozdemir, F., Koca, H. 2005. Differential responses of lipid peroxidation and antioxidants in the lea stress conditions. Plant Science. 163, 769-779.
Wahid, A., Gelani, S., Ashraf, M., Foolad, M.R., 2007. Heat tolerance in plants: an overview. Environmental Experiments Botany. 61, 199–223.
Wardlaw, I.F., Blumenthal, C., Larroque, O., Wrigley, C.W., 2002. Contrasting effects of chronic heat stress and heat shock on kernel weight and flour quality in wheat. Functional. Plant Biology. 29, 25–34.
Zadoks, J.C., Chang, T.T., Konzak, C.F. 1974. A decimal code for the growth stages of cereals. Weeds Research. 14, 415-421.
Zhang, J., Jiang, X.D., Li, T.L., Cao, X.J., 2014. Photosynthesis and ultrastructure of photosynthetic apparatus in tomato leaves under elevated temperature. Photosynthetica 52, 430–436.