Effect of Waterlogging on Yield and Yield Components of Wheat (Triticum aestivum L.)

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1 International Journal of Applied Agricultural Research ISSN Volume 5 Number 4 (2010) pp Research India Publications Effect of Waterlogging on Yield and Yield Components of Wheat (Triticum aestivum L.) Seyed Keyvan Marashi 1 * and G.S. Chinchanikar 2 1 Department of Agronomy, Islamic Azad University, Ahvaz Branch, Ahvaz, Iran 2 Department of Botany, University of Pune: , India *Corresponding Author Marashi_47@yahoo.com Abstract Waterlogging is an important factor influencing wheat yield. In the study, a field experiment was designed to evaluate the response of three wheat cultivars to waterlogging at three growth stages (stress at the first of tillering, stress at the beginning of stem elongation and stress at the commencement of booting stage) with three levels of waterlogging duration (0, 7, and 14 days). Results indicated that yield, yield components were significantly affected by cultivars, stages of waterlogging and waterlogging durations. Response of stress at the first of tillering was higher than other growth stages except 1000-kernel weight. Increasing waterlogging duration decreased yield, spike number per square meter, 1000-kernel weight and number of kernels per spike. Average yield loss was 71.6 kg/h or 1.5% for each day waterlogging which were mainly caused by a decrease in kernel number per spike and spike number per square meter which were reduced by 1.1% and 0.74% per day of waterlogging, respectively. To evaluate tolerant mechanisms of wheat cultivars, aerenchyma tissue and proline content were evaluated in a pot experiment under waterlogging duration (0, 7, 14 and 21 days) at seedling stage. Results indicated that aerenchyma formation and proline accumulation were important in determining waterlogging-tolerance in wheat cultivars. Keywords: Aerenchyma, proline, waterlogging tolerance, wheat. Introduction Waterlogging stress has been known to be major limiting factors for plant growth and yield during the waterlogged season. Aslam and Prathpar (2001) reported that wheat cultivation could be done successfully when the water table was 150 cm below soil surface. When the water table was 100 cm or less below soil surface, soil is saturated with water, the root growth decreased and got restricted to small region near the soil

2 562 Seyed Keyvan Marashi et al surface. Behaviour and perceptible variations of roots under waterlogging appeared after approximately 48 h.. Negative effects of waterlogging include denitrification and loss or leaching of nutrients (particularly N) out of the crop root zone, thus plant uptake of N,P,K are reduced (Stieger and Feller,1994; Hill et al., 2004). In waterlogged soil the functional relationship between roots and shoots was disturbed and carbon assimilation inhibited (Bradford and Hsiao, 1982; Taiz and Zeiger, 2003). Olgun et al. (2008) reported that, waterlogging treatment from 5 to 50 d after flowering of wheat reduced grain yield by % respectively. In other study, average yield losses in 15 wheat genotypes under 5 weeks waterlogging at the 3-4 leaf stage were about 44% (Collaku and Harrison, 2002). Collaku and Harrison (2002) also in evaluating of nine wheat genotypes to four levels of waterlogging treatment ( 0, 10, 20, and 30 d ) at 3 to 4 leaf stage revealed that response of genotypes was different for reduction of grain yield from 34% to 60%. It is observed that winter wheat with late-maturing duration was less sensitive to waterlogging than spring wheat with early-maturing duration (Samand et al., 2001). One mechanism that plant use to alleviate waterlogging effects is developing aerenchyma. But the ability of aerenchyma formation is different between plants (Watkin et al.,1998; Videmsek et al., 2006). Boru et al. (2003) reported that the percentage of root porosity in tolerant spring wheat under 2-3 weeks of hypoxic conditions was approximately two times more than sensitive genotype. The objectives of this study were to evaluate yield, yield components and tolerant mechanisms of wheat cultivars under waterlogging stress. Materials and Methods Field Experiment To evaluate effect of waterlogging conditions, three wheat cultivars were grown under no waterlogging (control), 7 and 14 days of continuous waterlogging, starting at the first of tillering stage, beginning of stem elongation and at the commencement of booting stage corresponding to stage 21(ZG 21), 31(ZG 31 ) and 45(ZG 45 ) of Zadoks scale, respectively (Zadoks et al.,1974). The field experiment was conducted in the Agricultural Research Station, Islamic Azad University, Iran during and cropping seasons. Experimental design was a factorial according to a randomized complete block design (RCBD) with three replications. Soil was kept saturated with water by continued irrigation for duration of waterlogging treatments. In order to prevent from penetration of water from one plot to other plots, 2 meter distance between plots was maintained. Each plots included seven rows, each six meters long and 20 cm apart. Cultivars were included Chamran, Dez and Vee/nac. The three wheat cultivars were spring wheat. Vee/nac cultivar had earlier maturing duration than other cultivars (two weeks). On the basis of soil composition amount of 150 Kg N ha -1 (1/2 at sowing stage and 1/2 at tillering stage), 100 Kg ha -1 P 2 O 5 (at sowing) and 100 Kg ha -1 K 2 0 (at sowing) were applied in each cropping year. Yield and yield components were estimated after physiological maturing by harvesting interior rows (the outer rows excluding at least 0.5 m from either end of the rows).

3 Effect of Waterlogging on Yield and Yield Components 563 Pot Experiment This experiment was conducted in Botany department, University of Pune-India (during cropping season). Four periods of waterlogging were applied as treatments: 0 (control), 7, 14 and 21 d. wheat cultivars included were Chamran, Dez and Vee/nac. In order to apply waterlogging treatments, cultivars were planted in pots made of PVC (30 cm height, 16 cm diameter, containing farm soil) and they were waterlogged at seedling stage (ZG 11 ) in a basin filled with water. Water was periodically added to keep the level 2 cm above the soil level. Aerenchyma tissue and proline content were determined after the end of waterlogging treatments. Aerenchyma tissue was assessed in hand sections of adventitious roots behind the root tip (Watkin et al., 1998; Videmsek et al., 2006). Estimation of proline performed according to method of Bates et al. (1973). The data so collected were analyzed statistically by using computer software MSTATC. Duncan's multiple range tests at alpha level 5% was computed to compare the significant differences among means. Results Grain Yield Comparison of grain yield showed significant differences for tested cultivars (Table 1). Among three wheat cultivars, the maximum and the minimum of grain yield observed in Chamran and Vee/nac cultivars. Effect of waterlogging at different growth stages was significantly difference (Table1). More negative effects of waterlogging observed when waterlogging were at early growth stages of wheat. Average grain yield losses at the first of tillering (ZG 21 ), beginning of stem elongation (ZG 31 ) and commencement of booting (ZG 45 ) were 18.2%, 11.3% and 4.8 %, respectively. A significant effects for prolonging of waterlogging duration were observed for grain yield (Table1). Yield losses for 7d (W 7 ) and 14 d (W 14 ) waterlogging were 13.3% and 21.1%, respectively. In our study, average yield reduction for each day waterlogging was 71.6 kg/h or 1.5 %. Meanwhile percentage of reduction for first week was more than second week of waterlogging. Table 1: Mean values for grain yield, number of spikes per square meter, number of kernels per spike, 1000-kernel weight. Treatment Grain yield ( kg/ha ) Spike number per square meter Kernel number per spike kernel weight (g) Cultivars Chamran a a 27.1 a 40.7 b Dez b b 24.9 b 42.4 a Vee/nac c c 22.4 c 37.3 c Stage of waterlogging

4 564 Seyed Keyvan Marashi et al ZG c b 23.2 c 40.7 a ZG b b 24.4 b 40.5 a ZG a a 26.7 a 39.3 b Duration of waterlogging 0 day (W 0 ) (control) a a 27.1 a 40.9 a 7 days (W 7 ) b b 24.2 b 40.1ab 14 days (W 14 ) c b 23.1 c 39.5 b Means with different letters are significantly different at P=0.05, using Duncan's Multiple Range Test. Number of Spikes per Square Meter The number of spikes per square meter in wheat cultivars were significantly different. The maximum and the minimum of spike number per square meter obtained in Chamran and Vee/nac cultivars (Table1). Also there was significant influence of waterlogging on spike number per square meter. It was most affected when waterlogging was imposed at ZG 21 and ZG 31. Reduction in spike number per square meter at ZG 21, ZG 31 and ZG 45 were 10.2, 7.5 and 0.5%, respectively. Results also indicated that, extending of waterlogging duration caused a decrease in spike number per square meter. The maximum of spike number per square meter was obtained from control (432.3) whereas W 14 gave the lowest one (387.0) (Table 1). As compared with control, reduction in spike number per square meter in W 7 and W 14 were 7.6 and 10.4%, respectively. Number of Kernels per Spike Screening of wheat cultivars showed a significant difference for kernel number per spike (Table 1). Among three cultivars, the maximum and the minimum of kernel number per spike observed in Chamran and Vee/nac cultivars by 27.1 and 22.4, respectively. There was significant effect of waterlogging at different growth stages on kernel number per spike (Table1). Number of kernels per spike was most affected when waterlogging was at first of tillering stage (ZG 21 ). In our study, the percentage reduction in kernel per spike at ZG 21, ZG 31 and ZG 45 were 16.8, 10.0 and 1.5%, respectively. Results also showed that increasing of waterlogging duration from 0 to 14 d reduced the number of kernels per spike (Table1). Number of kernels per spike decreased from 10.7% in 7d waterlogging to 14.8% in 14d waterlogging. One Thousand Kernel Weight One thousand kernel weight is an important yield components in the wheat. The results showed that, one thousand kernel weight was significantly different for three wheat cultivars (Table1). In this study Dez cultivar was superior as compared to Chamran and Vee/nac cultivars. Differences between stages of waterlogging were found to be significant in one thousand kernel weight (Table 1). Less seed weight was obtained from commencement of booting stage (ZG 45 ). Reduction of one thousand kernel weight at the first tillering (ZG 21 ), stem elongation (ZG 31 ) and booting stages

Effect of Waterlogging on Yield and Yield Components 565 (ZG 45 ) were 0.5%, 1% and 3.9%, respectively. One thousand kernel weight was significantly influenced by duration of waterlogging (Table 1).

9% compared to control, respectively. Aerenchyma Formation Cross sectional area of roots of three wheat cultivars are shown in fig 1.

5 Effect of Waterlogging on Yield and Yield Components 565 (ZG 45 ) were 0.5%, 1% and 3.9%, respectively. One thousand kernel weight was significantly influenced by duration of waterlogging (Table 1). A decrease in one thousand kernel weight as a result of increasing duration of waterlogging was observed. Reduction in one thousand kernel weight in W 7 and W 14 were 1.9% and 3.9% compared to control, respectively. Aerenchyma Formation Cross sectional area of roots of three wheat cultivars are shown in fig 1. Screening of cross-sectional area showed that three wheat cultivars differed in their ability to develop aerenchyma tissue under waterlogging condition. The maximum of aerenchyma formation occurred in Chamran cultivar and the minimum was in Vee/nac cultivar. This is suggested that aerenchyma formation in Chamran cultivar is more importance in determining waterlogging-tolerance than Vee/nac cultivar, since it enhances the internal movement of gases. Figure 1: Cross sectional area of roots in three wheat cultivars after 21 d waterlogging condition a) No waterlogging (control) b) Chamran c) Dez d)vee/nac. Proline Content Results for proline content into the roots and leaves are presented in fig 2. Results showed that: 1) Proline content into the roots is more than leaves 2) proline content increased under prolonging waterlogging duration 3) Increasing of proline content into the roots was more than leaves under waterlogging duration 4) The maximum of proline content obtained in Dez and Chamran cultivars and the minimum was in Vee/nac cultivar. (a) (b) Figure 2: Comparison of proline content in three wheat cultivars under waterlogging duration a) Roots b) Leaves.

6 566 Seyed Keyvan Marashi et al Discussion Waterlogging tolerance is a generic trait in plants (Sheikh et al., 2008; Setter et al., 2009). Comparison of three wheat cultivars under waterlogging treatments showed that Chamran and Dez cultivars were the best for yield and all yield components and the weakest was Vee/nac cultivar. This was due to 1) more ability of aerenchyma formation and proline accumulation in Chamran and Dez cultivars. Watkin et al. (1998); Videmsek et al. (2006); Olgun et al. (2008) also suggested to differential ability of cultivars to aerenchyma formation and proline production under waterlogging stress. 2) Earlier maturing duration of Vee/nac as compared to other cultivars. Samand et al. (2001) also suggested to less sensitivity of winter wheat with late-maturing duration than spring wheat with early-maturing duration. Results for waterlogging at different growth stages showed that yield and all yield components except 1000-kernel weight were more affected by waterlogging at early growth stages. It was seemed that yield components that appeared at early growth season are more sensitive to waterlogging. This was contrast with findings of Bakshandeh (1989); Samand et al. (2001) and Belford et al. (2008). The maximum 1000 kernel weight was obtained at ZG21. This was due to less seeds competition at grain filling period for dry matter accumulation as a result of reduction of spike number per square meter and number of seeds per spike. Samand et al. (2001) who found that if waterlogging treatment was applied before the anthesis stage, the seed weight of waterlogged wheat was not different. Decrease in seed weight support the findings of Olgun et al. (2008). They showed that reduction in seed weight for 5 to 50 d waterlogging after flowering stage were from 8.4% to 89.4%. Screening of duration of waterlogging showed that grain yield decreased under prolonging of waterlogging duration. This was due to decline in yield components namely spike number per square meter and number of seeds per spike and kernel weight under waterlogging duration. Other studies also have suggested to reduction of yield and all yield components with increasing of waterlogging duration (Collaku and Harrison, 2002; Lakzadeh et al., 2002 ; Olgun et al., 2008). References [1] Aslam, M., and Prathpar, S. A., 2001, "Waterlogging and salinity in the rice wheat zone of Sind, Pakistan," Food Products Press, New York, USA, pp [2] Bakshandeh, A., 1989, "Effect of waterlogging at early stage of crop development on the growth of wheat (Triticum aestivum L.)," Ph.D. Thesis, University of Reading, London, England. [3] Bates, LE.,1973, "Rapid determination of free proline for water stress studies," Plant and Soil, 39: [4] Belford, R.K., Cannell, R.Q., and Thomson, R.J., 2008, "Effects of single and multiple waterloggings on the growth and yield of winter wheat on clay soil," Journals of the Science of Food and Agriculture, 36:

7 Effect of Waterlogging on Yield and Yield Components 567 [5] Bradford, K.J., and Hsiao, T.C.,1982, "Stomatal behavior and water relations of waterlogged Tomato plants," Plant Physiology, 70: [6] Boru, G., Ginkel, M. V., Trethowan, R. M., Boersma, L., and Kronstad, W. E., 2003, "Oxygen use from solution by wheat genotypes differing in tolerance to waterlogging," Euphytica, 132(2) [7] Collaku, A., and Harrison, S.A., 2002, losses in wheat due to waterlogging," Crop Science, 42: [8] Hill, N., Moyes, N., McTaggart, R., Anderson, W.K., and Tugwell, R., 2004, "Waterlogging and nitrogen management for wheat in high rainfall cropping areas of southern western Australia," 4 th Australian Agronomy Conference, Australia, pp [9] Lakzadeh, A., Kashani, A., Bakhshandeh, A., and Mameghani, R., 2002, "Effects of waterlogging in difference growth stages on tiller number of wheat (Chamran cultivar), " 7 th Congress of Agronomy and Plant Breeding, Karaj, Iran, pp [10] Olgun, M., Kumlay, A. M., Adiguzel, M.C., and Caglar, A., 2008, "The effect of waterlogging in wheat (T. aestivum L.)," Plant Soil Science, 58: [11] Samand, A., Meisner, C.A., Saifuzzaman, M., and Ginkel, V. M., 2001, "Waterlogging tolerance. In: Application of physiology in wheat breeding," M.P. Reynolds, J.I. Ortiz-Monasterio and A. Mcnab, CIMMYT- Mexico, pp: [12] Setter, T. L., Waters, I., Sharma, S. K., Singh, K. N., Kulshreshtha, N., Yaduvanshi, N. P. S., Ram, P. C., Singh, B. N., Rane, J., McDonald, G., Khabaz-Saberi, H., Biddulph, T. B., Wilson, R., Barclay, I., McLean R., and Cakir, M., 2009, "Review of wheat improvement for waterlogging tolerance in Australia and India: the importance of anaerobiosis and element toxicities associated with different soils," Annals of Botany, 103(2): [13] Sheikh, F., Kalateh Arabi, M., Soghi, H., Taghi Bazi, M., Abroudi, and A. M., 2008, "The effect of waterlogging stress at filling stage on yield and yield component of wheat (Triticum aestivum L.)," Iranian Society of Agronomy and plant breeding, Gurgan University of Agricultural Science and Natural Resource, Iran, 1(1): [14] Stieger, P.A., and Feller, U., 1994, "Nutrient accumulation and translocation in maturing wheat plants grown on waterlogging soil", Plant and Soil, 160: [15] Taiz, L., and Zeiger, E., 2003, "Plant physiology," 3 rd edition, New Delhi- Banglore Panima Publishing Corporation, pp: [16] Videmsek, U., Turk, B., Vodnik, D., 2006, "Root aerenchyma formation and function," Acta Agriculture Slovenica, pp: [17] Watkin, E., Campbell, J.T., and Greenway, H., 1998, "Root development and aerenchyma formation in two wheat cultivars and one triticale cultivar grown in stagnant agar and aerated nutrient solution," Annals of Botany, 81: [18] Zadoks, J.C., Chang, T.T., Konzak, C.F., 1974, "A decimal code for the growth stage of cereals," Weed Res, 14:

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A comparative study of grain yield and yield components of wheat (Triticum aestivum L.) in response to waterlogging condition

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