Evaluation of Three Physiological Traits for Selecting Drought
Transcription
Evaluation of Three Physiological Traits for Selecting Drought
J. Agric. Sci. Technol. (2005) Vol. 7: 81-87 Evaluation of Three Physiological Traits for Selecting Drought Resistant Wheat Genotypes R. Amiri Fahliani1 and M. T. Assad 2∗ ABSTRACT Physiological traits are receiving increasing attention as screening tools for drought resistance. Two field experiments were conducted in 1998 at the Experimental Station of College of Agriculture, Shiraz University at Badjgah, to evaluate the effectiveness of leaf water potential, leaf osmotic potential and canopy temperature in screening resistant bread wheat (Triticum aestivum L.) genotypes. Nine wheat cultivars consisting of drought resistant, intermediate and susceptible genotypes were grown in two randomized complete block designs with three replications. The experiments only differed with respect to their irrigation regimes. Leaf water potentials and leaf osmotic potentials at three developmental stages -stem elongation, booting and flowering - under water stress conditions, and canopy temperature in non-stress conditions could discriminate between resistant and susceptible cultivars. Although the drought susceptibility index could partly discriminate between resistant and susceptible cultivars, it was not evaluated as a reliable index. The linear regression of grain yield on each trait was determined. The linear regressions of grain yield on leaf water potential; leaf osmotic potential and canopy temperature confirmed the above results. Keywords: Triticum aestivum, Leaf water potential, Osmotic potential, Turgor potential, Canopy temperature. water-stressed treatment were much lower than those of the well-watered control. They also concluded that osmotic adjustment did not contribute to the differences between cultivars in response to water stress. However, Blume (1989) suggested that induced osmotic adjustment under drought stress might be an important component of drought resistance in barley growth. Neumann (1995) rejected the notation that a stressinduced reduction in cellular turgor pressure is a primer cause of growth inhibition. Hoffman and Jobes (1978) reported that the relationship between crop yield and total leaf water potentials was negative and linear. Canopy temperature is another criterion, which has been considered effective in screening wheat (Blum et al., 1982; Pinter et al. 1990; Golestani Araghi and Assad, 1998) INTRODUCTION Soil water deficit depresses agricultural crop yield in many parts of the world. Plant breeders search for effective and repeatable criteria to screen germplasms, for drought resistance in segregating populations. Plant breeders have used selected physiological parameters that are important in the plantwater relations of bread wheat (Triticum aestivum L.) under stress conditions. Levitt (1972), and Matin et al. (1989) reported that total water potentials of plant tissue are different between drought-resistant and drought-susceptible genotypes. Moustafa et al. (1996) used leaf water and osmotic potentials to differentiate apparent drought tolerance among wheat cultivars. They reported that the leaf water potentials of the _____________________________________________________________________________ 1 Department of Agronomy, College of Agriculture, Yasuj University, Yasuj, Islamic Republic of Iran. Department of Agronomy, College of Agriculture, Shiraz University, Shiraz, Islamic Republic of Iran. ∗ Corresponding author. 2 81 ____________________________________________________________ Amiri Fahliani and Assad middle rows were used for grain yield determination, and data were recorded on the basis of 10 randomized selected plants in the second and seventh rows. Fertilizer was applied at the rate of 80 Kg/ha N and 70 Kg/ha P2O5. Crops received one half of N in urea form and total amount of P2O5 at planting, while the remaining N was applied at tillering stage. The two experiments differed with respect to their irrigation regimes. The non-stress experiment received water when 40±5 mm evaporation occurred from pan class A, while the stress experiment was not irrigated after plant establishment. The soil moisture status in the non-stress experiment was measured with a neutron probe (Troxler Model 2651). The effective rainfall during 1998-9 and total irrigation for each experiment are given in Table 1. Leaf water potential (ψw) was measured using a PMS pressure bomb (PMS Instrument Co., Corvallis, OR) at stem elongation, booting and flowering plant developmental stages based on Zeidak`s Code in both experiments. The youngest fully expanded leaf was detached and placed rapidly in a sample chamber and the pressure was recorded. For each developmental stage three randomly selected plants were used. Measurements were completed between 13.00 and 15.00 hours. To measure osmotic potential (ψs), the youngest fully expanded leaf of each of 10 randomly selected plants was used for each developmental stage. Leaves were placed in plastic bags and rapidly packed in and pearl millet (Singh and Kanemasu, 1983) genotypes resistant to drought. Plant breeders have used selected physiological parameters that play a role in the plant-water relations of wheat under stress conditions (Keim and Kronstad, 1981; Jaradat and Konzak, 1983; Seropian and Planchon, 1984; Turner, 1986 a, b; Blum, 1989; Matin et al. 1989). The objectives of this study were to evaluate leaf water potential, leaf osmotic potential, and canopy temperature in differentiating wheat cultivars for drought resistance and to find their relationships with grain yield. MATERIALS AND METHODS Two field experiments were conducted in 1998 at the Experimental Station of College of Agriculture, Shiraz University, at Badjgah (Iran). Nine hexaploid wheat cultivars were used in both experiments. According to the ranking proposed by the Seed and Plant Improvement Institute and later approved by Golestani Araghi and Assad (1998), three drought resistant cultivars (Omid, Roshan, and Kal-Haydary), three intermediate (Bayat, Niknejad, and M-75-5), and three drought sensitive ones (Falat, Darab, and Azadi Cross) cultivars were used. Cultivars were planted on 6th November 1998 in two experiments using randomized complete block designs each with three replications on a clay loam soil. Each plot consisted of eight 5m rows with the rows 25cm apart. The four Table 1. Precipitation distribution and total irrigation for each experiment. Month November December January February March April May June Total Total water used Effective Rainfall 19.7 126.8 26.1 165.8 60.4 50.7 5.0 454.5 Non-stress 140 200 100 440 894.5 82 Irrigation (mm) Stress 140 140 594.5 Selecting Drought Resistant Wheat Genotypes ____________________________________ reduced under stress conditions, but the reductions in spike length were not significant. Susceptible cultivars on average showed higher drought susceptibility indices than intermediate and resistant cultivars, however there was some misclassification, especially in respect to intermediate and resistant cultivars. The leaf water potentials (ψw) of cultivars at three developmental stages are given in Table 3. The ψw values decreased with maturation in both environments and cultivars showed significant differences in all stages (p<0.01). Table 3 indicates that leaf water potential apparently discriminated between drought resistant, intermediate, and susceptible cultivars at the three stages in both environmental conditions. Drought resistant cultivars showed lower ψw values as compared to sensitive ones. This is in agreement with the results obtained by others (Barlow et al., 1980; Keim and Kronstad, 1981; Matin et al. 1989; Entz and Flower, 1990; Moustafa et al., 1996). The linear relationship between ψw and grain yield was significant under stress condition only (Table 4). Although ψw in all stages and conditions could classify cultivars in respect to drought resistance, however, Table 3 shows that ψw values in stress conditions were more effective. The results obtained from linear regressions also confirmed these results. The leaf osmotic potential (ψs) of cultivars at different developmental stages in stress and non-stress conditions are given in Table 3. The trend of variation in ψs values in different developmental stages was similar to that of ψw. Cultivars were significantly different with regard to values in all stages, in both environments (p<0.01). The ψs values of drought resistant cultivars were, on average, lower than those of drought susceptible ones in all conditions, indicating that ψs was an effective technique in screening resistant genotypes. Other investigators (Grumet et al., 1987; Blume, 1989; Musick et al., 1994) also reported that drought resistant cultivars had lower ψs values as compared to a box in order to avoid water-loss as vapor from the sample and maintained at -15°C for five hours. The frozen samples were then thawed for approximately 30 minutes and the freezing point (T) of collected saps was measured using a digital thermometer (Model ET - 2001). The osmotic potential then was calculated (Kramer, 1995) by: ψs = (-T / 1.86) × 2.27 The canopy temperature (TC) of each plot was measured at three development al stages at 13.00 to 15.00 hours in both experiments using an infrared thermometer (Kane-May Model Infratrace 800). The instrument was pointed down at three random points in each plot and held at an oblique angle to the canopy surface to minimize the influence of soil exposure. The drought susceptibility index (S) was also determined by the following equation (Fischer and Mourer, 1978): S = [1-( y D / y p)] / D Where y D and y p , are the grain yield of each cultivar at stress and non-stress conditions respectively, and D = 1- ( Y D / Y P). Y D and Y P are the mean yield of all the cultivars under stress and non-stress conditions. Analysis of variance in all the measurements was conducted by Statistical Analysis System (SAS, 1985). Means were separated using the least significant difference (LSD). The regression of grain yield on each physiological index was also determined. To compare the effects of stress and non-stress, and cultivars by moisture conditions interaction, a combined analysis of variance was used. RESULTS AND DISCUSSION The mean grain yield and some related components for both experiments are showed in Table 2. Cultivars did not differ significantly in respect to grain yield under non-stress conditions, however, the differences were significant under stress conditions (p<0.01). This emphasized the different responses of cultivars to drought conditions. 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Table 3 indicates that ψs values in stress conditions could classify genotypes better compared with non-stress condition. The regressions of grain yield on ψs were significant under stress condition (Table 4). This indicated that leaf osmotic potential might be a good trait for selecting wheat genotypes resistant to draught in stress conditions. The differences in the canopy temperature (Tc) of cultivars were highly significant in both moisture conditions (p<0.01) and growth stages, except stem-elongation in non-stress conditions (Table 3). This exception may be due to similar transpiration activity of all cultivars under well-watered environmental conditions. Except for stem elongation in non-stress condition, the linear regressions of grain yield on canopy temperature were not significant in other cases (Table 4). The canopy temperature of cultivars in flowering, under non-stress conditions, could help discriminate between resistant and susceptible cultivars better than at other stages. Pinter et al. (1990) and Golestani Araghi and Assad (1998) also reported that (Ta-Tc) is a valuable technique in screening drought resistant genotypes. 1. Barlow, E. W. R., Lee, J. W., Munns, R. and Smart, M.G. 1980. Water Relations of Developing Wheat Grain. Aust. J. Plant Physiol., 7: 519-525. 2. Blum, A. 1989. Osmotic Adjustment and Growth of Barley Genotypes under Drought Stress. Crop Sci. 29: 230-233. 3. Blum, A., Mayer, J. and Gozlan, G. 1982. Infrared Thermal Sensing of Plant Canopies as a Screening Technique for Dehydration Avoidance in Wheat. Field Crop Res., 5: 137-146. 4. Entz, M. H. and Flower, D. B. 1990. Influence of genotype, water and N on leaf water relations in No-till winter wheat. Can. J. Plant Sci., 70: 431-441. 5. Fischer, R. A. and Maurer, R. 1978. Drought Resistance in Spring Wheat Cultivars. 1: Grain Yield Responses. Aust. J. Agr. Res., 29: 897-912. 6. Golestani Araghi, S.and Assad, M. T. 1998. Evaluation of four Screening Techniques for Drought Resistance and Their Relationship to Yield Reduction Ratio in Wheat. Euphytica, 103: 293-299. 7. Grumet, R., Albrechtensen, R. S. and Hanson, A. D. 1987. Growth and Yield of Barley Isopopulations Differing in Solute Potential. Crop Sci., 27: 991-995. 8. Hoffman, G. J. and Jobes J. A. 1978. Growth and Water Relations of Cereal Crops as Influenced by Salinity and Relative Humidity. Agron. J., 70: 765-769. 9. Jaradat, A. and Konzak, C. F. 1983. Screening of Wheat Genotypes for Drought Tolerance: 1. Excised-leaf Water Retention. Cereal Res. Com., 11: 179-186. 10. Keim, D. L. and Kronstad, W. E. 1981. Drought Response of Winter Wheat Cultivars Grown Under Field Stress Conditions. Crop Sci., 21: 11-15. 11. Kramer, P. J., and J. S. Boyer. 1995. Water Relations of Plant and Soils. Academic press, Inc., New York, 494 pp. 12. Levitt, J. 1972. Response of Plants to Environmental Stress. Academic Press, New York. 13. Matin, M. A., Brown, J. H. and Ferguson, H. 1989. Leaf Water Potential, Relative Water Content, and Diffusive Resistance as Screening Techniques for Drought Resistance in barley. Agron. J., 81: 100-105. 14. Moustafa, M. A., Boersma, L. and Kronstad, W. E. 1996. Response of four Spring Wheat CONCLUSION This study showed that leaf water potentials and leaf osmotic potentials of wheat plants at the three developmental stages (stem elongation, booting, and flowering) under water stress conditions, and canopy temperature in non-stress conditions were the best criteria for screening drought resistant genotypes. Although the drought susceptibility index could partly discriminate between resistant and susceptible cultivars, it was not evaluated as a consistent reliable criterion alone. For more reliability of results use of more genotypes and seasons are recommended. REFERENCES 87 ____________________________________________________________ Amiri Fahliani and Assad 18. SAS Institute, 1985. SAS User’s Guide Statistics. Cary, NC: SAS Institute. 19. Seropian, C. and Planchon, C. 1984. Physiological Response of Six Bread Wheat and Durum Wheat Genotypes to Water Stress. Euphytica., 33: 757-767. 20. Singh, P. and Kanemasu, E. T. 1983. Leaf and Canopy Temperature of Pearl Millet Genotypes Under Irrigated and Nonirrigated Conditions. Agron. J., 75: 497-501. 21. Turner, N. C. 1986a. Adaptation to Water Deficits: a Changing Perspective. Aust. J. Plant Physiol., 13: 175-190. 22. Turner, N. C. 1986b. Crop Water Deficits: a Decade of Progress. Adv. Agron., 39: 1-15. Cultivars to Drought Stress. Crop Sci., 36: 982-986. 15. Musick, J. T., Jones, O. R., Stewart, B. A. and Dusek, D. A. 1994. Water-yield Relationships for Irrigated and Dryland Wheat in the U.S. Southern Plains. Agron. J., 86: 980986. 16. Neumann, P. M. 1995. The Role of Cell Wall Adjustment in Plant Resistance to Water Deficits. Crop Sci., 35: 1258-1266. 17. Pinter, P. S., Jr. Zipoli, G., Reginato, R. J., Jackson, R. D., Idso, S. B. and Hopman, J. P. 1990. Canopy Temperature as an Indicator of Differential Water use and Yield Performance Among Wheat Cultivars. Agric. Water manage., 18: 35-48. ارزﻳﺎﺑﻲ ﺳﻪ ﻣﻌﻴﺎر ﻓﻴﺰﻳﻮﻟﻮژﻳﻚ ﺟﻬﺖ ﮔﺰﻳﻨﺶ ژﻧﻮﺗﻴﭗﻫﺎي ﮔﻨﺪم ﻣﻘﺎوم ﺑﻪ ﺧﺸﻜﻲ ر .اﻣﻴﺮي ﻓﻬﻠﻴﺎﻧﻲ و م .ت .آﺳﺎد ﭼﻜﻴﺪه ﺻﻔﺎت ﻓﻴﺰﻳﻮﻟﻮژﻳﻚ ﺑﻪ ﻋﻨﻮان اﺑﺰاري ﺟﻬﺖ ﺗﺸﺨﻴﺺ ژﻧﻮﺗﻴﭗﻫﺎي ﻣﻘﺎوم ﺑﻪ ﺧﺸﻜﻲ ﻣﻮرد ﺗﻮﺟﻪ روز اﻓﺰون ﻗﺮار ﮔﺮﻓﺘﻪ اﺳﺖ .دو آزﻣﺎﻳﺶ در ﺳﺎل 1377در اﻳﺴﺘﮕﺎه ﭘﮋوﻫﺸﻲ داﻧﺸﻜﺪه ﻛﺸﺎورزي ،داﻧﺸﮕﺎه ﺷﻴﺮاز ﺟﻬﺖ ارزﻳﺎﺑﻲ ﺗﺎﺛﻴﺮ ﭘﺘﺎﻧﺴﻴﻞ آب ﺑﺮگ ،ﭘﺘﺎﻧﺴﻴﻞ اﺳﻤﺰي ﺑﺮگ ،و درﺟﻪ ﺣﺮارت ﺳﺎﻳﻪ اﻧﺪاز ﺑﺮ ﺗﺸﺨﻴﺺ ژﻧﻮﺗﻴﭗﻫﺎي ﻣﻘﺎوم ﮔﻨﺪم ) (Triticum aestivum L.اﻧﺠﺎم ﮔﺮﻓﺖ .ﻧﻪ رﻗﻢ ﮔﻨﺪم ﻣﻘﺎوم ،ﻧﻤﻴﻪ ﻣﻘﺎوم و ﺣﺴﺎس ﺑﻪ ﺧﺸﻜﻲ در دو ﻃﺮح ﺟﺪاﮔﺎﻧﻪ ﺑﻠﻮكﻫﺎي ﻛﺎﻣﻞ ﺗﺼﺎدﻓﻲ ﺑﻪ ﻛﺎر ﺑﺮده ﺷﺪ .ﻳﻜﻲ از آزﻣﺎﻳﺶﻫﺎ ﺗﺤﺖ ﺷﺮاﻳﻂ ﻣﻄﻠﻮب و دﻳﮕﺮي ﺗﺤﺖ ﺗﻨﺶ آﺑﻲ ﻗﺮار ﮔﺮﻓﺖ .ﺻﻔﺎت ﻓﻴﺰﻳﻮﻟﻮژﻳﻚ ﻣﺰﺑﻮر در ﺳﻪ ﻣﺮﺣﻠﻪ رﺷﺪ )ﻃﻮﻳﻞ ﺷﺪن ﺳﺎﻗﻪ ،ﻏﻼف رﻓﺘﻦ و ﮔﻞ دﻫﻲ( ﮔﻴﺎه اﻧﺪازهﮔﻴﺮي ﺷﺪ .رﮔﺮﺳﻴﻮن ﺷﺎﺧﺺ ﺣﺴﺎﺳﻴﺖ ﺑﻪ ﺧﺸﻜﻲ ﺑﺮ ﻫﺮ ﻛﺪام از ﺻﻔﺎت ﻧﻴﺰ ﺗﻌﻴﻴﻦ ﺷﺪ .ﭘﺘﺎﻧﺴﻴﻞ آب ﺑﺮگ و ﭘﺘﺎﻧﺴﻴﻞ اﺳﻤﺰي ﺑﺮگ در ﻫﺮ ﺳﻪ ﻣﺮﺣﻠﻪ رﺷﺪ ﺗﺤﺖ ﺷﺮاﻳﻂ آﺑﻲ و درﺟﻪ ﺣﺮارت ﺳﺎﻳﻪ اﻧﺪازه ﺗﺤﺖ ﺷﺮاﻳﻂ ﻣﻄﻠﻮب در ﺗﺸﺨﻴﺺ ژﻧﻮﺗﻴﭗﻫﺎي ﺣﺴﺎس و ﻣﻘﺎوم ﻣﻮﺛﺮ ﺑﻮدﻧﺪ .ﻫﺮ ﭼﻨﺪ ﺷﺎﺧﺺ ﺣﺴﺎﺳﻴﺖ ﺑﻪ ﺧﺸﻜﻲ ﺗﺎ ﺣﺪودي در ﮔﺮوه ﺑﻨﺪي ارﻗﺎم ﺑﻪ ﺣﺴﺎس و ﻣﻘﺎوم ﻣﻮﺛﺮ ﺑﻮد ،وﻟﻲ ﻣﻌﻴﺎر ﻗﺎﺑﻞ اﻋﺘﻤﺎدي ﺗﺸﺨﻴﺺ داده ﻧﺸﺪ .رﮔﺮﺳﻴﻮن ﺧﻄﻲ ﺷﺎﺧﺺ ﺣﺴﺎﺳﻴﺖ ﺑﻪ ﺧﺸﻜﻲ ﺑﺮ ﺗﻤﺎم ﻣﻌﻴﺎرﻫﺎ ﺗﻌﻴﻴﻦ ﺷﺪ. 88