International Journal of Water Resources and Arid Environments 4(1): 08-14, 2015 ISSN PSIPW, 2015

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International Journal of Water Resources and Arid Environments 4(1): 08-14, 2015 ISSN 2079-7079 PSIPW, 2015 Using Bermudagrass (Cynodon dactylon L.) In Urban Desert Landscaping and as a Forage Crop for Sustainable Agriculture in Arid Regions and Combating Desertification Mohammad Pessarakli School of Plant Sciences, Forbes Bldg., Room 303, College of Agriculture and Life Sciences, The University of Arizona, Tucson, AZ 85721, USA Abstract: Bermudagrass (Cynodon dactylon L.), cv. Arizona Common, was studied in a greenhouse experiment to evaluate its growth responses in terms of shoot and root lengths, shoot and root fresh and dry weights and percentage of grass visual green cover under control and salt stress conditions viz., sodium chloride (NaCl). Plants were grown under control (distilled water) and four levels of salt (75, 150, 225 and 300 mm NaCl 1 equivalent to 4.38, 8.77, 13.15 and 17.53 g L sodium chloride, respectively), using a hydroponics system. Results showed that the adverse effect of salinity on the root length started to appear from the third harvest, generally at the higher levels of salinity and it was more pronounced as the exposure time to salinity progressed. Shoot length was more severely affected under salt stress compared to root length. The effect of salinity on shoot length was shown from the first harvest. Generally, for any harvest, only the highest NaCl level significantly reduced the shoot fresh and dry weights compared to that of the control or any other salinity levels. The root fresh and dry weights were decreased under salinity stress compared with the control treatment, while there was no significant difference detected between the root fresh or dry weights of different salinity treatments. All the grasses showed excellent recovery when transferred into the normal 1/2 strength Hoagland solution after the termination of the salinity stress period. Bermudagrass proved to have a satisfactory growth under the salinity levels of the experiment higher than the soil salinity of the harsh desert conditions. This indicates that Bermudagrass can effectively be used for cultivation under desert saline soils. Therefore, this grass could be recommended for sustainable production under harsh desert conditions with high soil salinity levels, limited water resources and drought conditions and effectively combating desertification processes. Key words: Bermudagrass Combating desertification Sustainable agriculture INTRODUCTION Continuous desertification of the arable lands due to urbanization, global warming and shortage of water mandates use of marginal lands and saline soils and low quality/saline water, particularly associated with desert regions, for irrigation and cultivation of plant species with high degrees of salt and drought tolerance, especially in regions experiencing water shortage. Cultivation and agricultural practices on saline soils and using low quality/saline water for irrigation imposes more salt stress on plants which are already under stress in these regions characterized with saline soils and shortage of water. Thus, there is an urgent need for finding salt/drought tolerant plant species to survive/sustain under such stressful conditions. Since the native plants are already growing under such conditions and are adapted to these stresses, these plants are the most suitable candidates to be manipulated under the minimum cultural practices and minimum inputs (water, fertilizer and other agrichemicals) for use under these stressful conditions. If stress tolerant species/genotypes/ cultivars of these native plants are successfully identified, there would be a substantial savings in cultural practices and inputs in using these plants by the growers and will result in substantial savings in the local, regional and the national currencies Corresponding Author: Mohammad Pessarakli, School of Plant Sciences, Forbes Bldg., Room 303, College of Agriculture and Life Sciences, The University of Arizona, Tucson, AZ 85721, USA. 8

of the countries. In previous investigations [1-4] indicated Bermudagrass grows worldwide as a turf that Bermudagrass (Cynodon dactylon L.) is a halophytic grass/landscaping plant, forage/pasture grass, or as a grass species that has a great potential to be used under sustainable desert cover grass species for soil erosion harsh and stressful environmental conditions, control in a wide range of climates, soils and characterized with arid regions, yet perform very environmental conditions. It is a warm-season grass satisfactory growth. This grass has multiple usages, species, a common major turfgrass/ landscaping plant or including animal feed, soil conservation and stabilization sustainable desert grass species used in the Southwest against erosion and use for lawns/parks/ recreation areas USA and other arid regions. The species is very tolerant as desert landscaping plant. to salinity, drought and heat stress, all characteristics of One strategy to enhance plant survival and recovery the Southwest or other desert areas. Despite its fairly well from salt stress is to use cultivars with superior salinity stress tolerance, the species is vigorously studied to find tolerance [5-9]. However, development of salt-tolerant more tolerant cultivars with more efficient nutrient and cultivars is not simple because the trait is quantitative water use [1, 23, 24]. These characteristics can be (controlled by many physiological mechanisms and beneficial for this plant species growth in arid and semigenes) [10, 11] and lacks a standardized screening arid regions where the soils are usually saline/sodic and protocol at both intra- and inter-species levels [5, 7, 12]. water is limited for irrigation and other agricultural uses. Therefore, reliable selection criteria are fundamental for The objectives of this study was to evaluate growth developing salt-tolerant cultivars. and performance of Bermudagrass, a warm-season desert Grasses along with various other kinds of plants species, in terms of shoot and root length as well as shoot often have to endure environmental stresses, with salinity (clippings) and root fresh and dry weights under control issues being one of the most common stresses they treatment (no salt) and various levels of salinity (sodium encounter [2-4, 13-21]. Salinity stress can stunt growth, chloride) stress, characteristics of arid region soils, for use dehydrate, cause chemical imbalances and make the under such harsh and stressful desert conditions for plants susceptible to injury. sustainable agriculture and combating desertification Some grasses deal with salinity better than other processes. species [2-4, 13-21]. To help aid in salinity tolerance, cultivars are selected and bread for having increased MATERIALS AND METHODS tolerance. For a grass to be considered tolerant to the saline conditions, it must meet some basic criteria such as Bermudagrass, cv. Arizona Common was used in a having acceptable quality, reasonable growth and greenhouse experiment to evaluate its shoot and root persistence at various levels of salinity. With that been growth as well as its fresh and dry weights under control said, assessment of salt tolerance in grasses should be and salt (NaCl) stress conditions, using a hydroponics evaluated under control (no salt), low, moderate and high technique. salt levels. Factors to look for could be overall visual Plants were grown as vegetative propagules in cups, appeal, shoot length, root length and biomass production. 9 cm diameter and 7 cm height, followed the procedures Perennial vegetation coverage in desert regions must used by Marcum and Pessarakli [2, 13], Marcum et al. [14], maintain adequate growth and persistence under variable Pessarakli [15, 23], Pessarakli and Touchane [4, 16], levels of soil salinity or salinity-laden water over several Pessarakli and Kopec [17, 18, 24] and Pessarakli et al. years. Successful assessment of salinity tolerance of [19-21]. Silica sand was used as the plant anchor medium. perennial, halophytic plants, therefore, should be based Each cup was fitted into one of the 9 cm diameter holes on growth at no saline, intermediate and high salinity cut in a rectangular plywood sheet 52 cm x 40 cm x 2 cm levels. In addition to shoot evaluation, root and verdure dimensions. The plywood sheets served as lids for the parameters should be measured in tolerance assessment, hydroponics tubs, supported the cups above the solution especially for plant species exposed to combined biotic or to allow for root growth and were placed on 48 cm x 36 cm abiotic stresses [5, 6, 22]. x 18 cm Carb-X polyethylene tubs containing 1/2 strength Salt problems in agricultural sites, especially in desert Hoagland solution No. 1 [25]. Four replications of each regions, are continually becoming more common. treatment were used in a randomized complete block Therefore, halophytic plants such as Bermudagrass design (RCBD) in this study. (Cynodon dactylon L.) need to be evaluated at salinity Plants were allowed to grow in this nutrient solution regimes up to sea-water salinity level [4, 6, 16] to select for six weeks. During this period, the plant shoots the best genotype. (clippings) were harvested weekly in order to allow the 9

Bermudagrass to reach full maturity and develop uniform statistically, there was not any significant difference and equal size plants. The harvested plant materials found between the cumulative root lengths at different (clippings) were discarded. The culture solutions were salinity levels (Table 1). Table 2 shows the root and shoot changed biweekly to ensure adequate amount of plant lengths and dry weights calculated based on the percent essential nutrient elements for normal growth and of the control values. The adverse effect of salinity on the th development. At the last harvest (6 week), the roots were root length started to show from the third harvest also cut to 0.5 cm length ensuring that all plants had generally at the higher levels (225 and 300 mm) of salinity. uniform roots and shoots for the stress phase of the This decrease was more pronounced as the exposure time experiment. to salinity progressed. As shown in Table 3, salinity The salt treatments were initiated by adding 50 mm, stress did not have any significant effect on the weekly 2.922 g NaCl per liter of the culture solution per day. Five root lengths of the grasses. treatments were used, including control (no salt addition), 75, 150, 225 and 300 mm salinity levels. The culture Shoot Length: The shoot length was more severely solution levels in the tubs were marked at the 10 liter affected under salt stress compared to the root length volume level and maintained at this level by adding (Tables 1 and 2 for the cumulative values and Table 3 for water as needed. After the final salinity level was reached the weekly values). The effect of salinity stress on shoot (day 6 after the initiation of the salt treatment), the shoots length was shown from the first harvest at all salinity (clippings) were harvested and the harvested plant levels, including the lowest (75 mm) NaCl level. However, materials were discarded for the last time. Pessarakli et al. [19] found that the effect of salinity on After the completion of the salt treatments, plant Saltgrass (Distichlis spicata L.) shoot length was shoots were harvested weekly for the evaluation of the statistically significant at 200 mm or more salinity levels fresh and dry matter (DM) production. At each weekly compared to the control and the low (100 mm) level of harvest, both shoot and root lengths were measured and NaCl. This is an indication of the difference in salt recorded. The harvested plant materials were oven dried tolerance of these two grass species (Bermudagrass vs. o at 65 C and DM weights were measured and recorded. Saltgrass). The recorded data were considered the weekly plant DM production. The shoot succulence was also calculated by Shoot Fresh Weight: The cumulative shoot fresh weights the ratio of the shoot fresh weight to dry weight. At the presented in Table 1 show that, generally, for any harvest, termination of the experiment, the last harvest, plant roots only the high (300 mm) level of NaCl significantly reduced o were also harvested, oven dried at 65 C and DM weights the shoot fresh weight compared to the control or any were determined and recorded. other levels of salinity. However, Pessarakli et al. [19] Electrical conductivity (EC) of the culture solutions found that there was no significant difference detected were initially measured and recorded. At the end of the on the salt grass shoot fresh weights between the high experiment, the EC of the culture solutions were also (400 mm) salinity level compared with the low (100 mm) measured and recorded (Final EC). By subtracting the final NaCl stress and the control treatment. They also EC from the initial EC values, the quantity of the salts found that as the stress period progressed, the medium removed by the plants from the culture medium was (200 mm) sodium chloride stress enhanced the salt grass estimated. shoot fresh weight. Pessarakli [27] also reported enhancement of Cotton (Gossypium hirsutum L.), a salt Statistical Analysis: The data were subjected to Analysis tolerant plant, growth and protein synthesis under low of Variance, using SAS statistical package [26]. The level of NaCl stress. means were separated, using Duncan Multiple Range test. The decrease in plant biomass production due to high level of salinity found in the present study may be RESULTS AND DISCUSSIONS attributed to the low medium water potential, specific ion toxicity, or ion imbalance as reported by Greenway and Root Length: The effects of salinity stress on the Munns [28]. cumulative length of roots are shown in Tables 1 and 2. The weekly effects of salinity stress on roots are shown Shoot Dry Matter (DM) Weight: Shoot dry weights in Table 3. Table 1, cumulative root length, shows that essentially followed the same pattern as the shoot root length significantly decreased at all salinity levels fresh weights. Generally, for any harvest, only the high compared with the control treatment. However, (300 mm) level of NaCl significantly reduced the shoot 10

Table 1: Bermudagrass growth responses (cum.) to NaCl stress Length (cm) Fresh Wt. (g) Dry Wt. (g) Shoot/Root ------------------------------------- ----------------------------------- --------------------------------- ---------------- NaCl (mm) Root Shoot Root Shoot Root Shoot ratio Control 55.6a 83.9a 0.71a 16.64a 0.47a 7.6a 16.2c 75 47.4b 69.0b 0.51b 15.66a 0.34b 6.9a 20.3b 150 48.7b 69.6b 0.44b 17.22a 0.29b 7.0a 24.1a 225 43.8b 60.7c 0.48b 15.00a 0.32b 6.2a 19.4b 300 45.9b 54.8c 0.44b 11.82b 0.29b 4.6b 15.9c The values are the averages of 4 replications of each treatment. The values followed by the same letters in each column are not statistically different at the 0.05 probability level. Table 2: Bermudagrass growth responses (based on the% of the control) to NaCl stress Length (cm) Dry Wt. (g) ------------------------------------------------------ ---------------------------------------------------- NaCl (mm) Root Shoot Root Shoot Control 100 100 100 100 75 85 82 72 91 150 88 83 62 92 225 79 72 68 82 300 83 65 62 61 Table 3: Bermudagrass weekly growth responses to NaCl stress Length (cm) Dry Wt. (g) Shoot/Root ---------------------------------------- -------------------------------------------- ----------------- NaCl (mm) Root Shoot Root Shoot ratio Control 6.2a 9.3a 0.05a 0.84a 16.8c 75 5.3a 7.7b 0.04a 0.77a 19.3c 150 5.4a 7.7b 0.03a 0.78a 26.0a 225 4.9a 6.7bc 0.03a 0.69ab 23.0b 300 5.1a 6.1c 0.03a 0.51b 17.0c The values are the averages of 4 replications of each treatment. The values followed by the same letters in each column are not statistically different at the 0.05 probability level dry weight compared to the control or any other levels of (Table 1). However, there was no significant difference salinity (Tables 1 and 2 for the cumulative shoot dry between the average root fresh weights of the different weights and Table 3 for the weekly shoot dry weights). salinity treatments. Marcum and Pessarakli [2, 13], However, Pessarakli et al. [19] found that there was no Marcum et al. [14], Pessarakli [15], Pessarakli and Kopec significant difference detected on the salt grass shoot dry [17, 18] and Pessarakli et al. [20, 21] also found that the weights between the high (400 mm) salinity level adverse effects of salinity stress were more pronounced compared with the low (100 mm) NaCl stress and the on shoot than the root growth. control. They also found that as the stress period progressed, the medium (200 mm) sodium chloride Root Dry Matter (DM) Weight: The root dry weight stress enhanced the Saltgrass shoot dry weight. essentially followed the same pattern as the root length Pessarakli [27] also reported enhancement of Cotton and root fresh weight. It was reduced at all salinity levels (Gossypium hirsutum L.), a salt tolerant plant, growth and compared with the control treatment (Tables 1 and 2 for protein synthesis by this plant under low level of NaCl the cumulative root dry weights and Table 3 for the stress. In the present study, the difference between the weekly root dry weights). However, there was no average shoot (clippings) weights of the highest level significant difference detected between the average root (300 mm) of salinity stressed plants and the other dry weights of the different salinity treatments either for treatments was wider as the exposure time to stress the cumulative root dry weights or for the weekly root dry progressed. weights. Marcum and Pessarakli [2, 13], Marcum et al. [14], Pessarakli [15], Pessarakli and Kopec [17, 18] and Root Fresh Weight: The root fresh weight essentially Pessarakli et al. [20, 21] also reported that the adverse followed the same pattern as the root length. It was effects of salinity stress were more pronounced on shoot reduced at all salinity levels compared with the control than the root growth. 11

Table 4: Bermudagrass succulence (Fr.Wt./Dry Wt.) response to salinity Shoot succulence ----------------------------------------------------------------------------------------------------------- NaCl (mm) (F.W./D.W.) (Based on% of Cont.) Control 2.19 100 75 2.27 104 150 2.46 112 225 2.42 111 300 2.57 117 Table 5: Bermudagrass percentage of green cover under salinity stress Percentage of Green Cover ---------------------------------------------------------------------------------------------------------------------------------------------------------------------------- Week NaCl ---------------------------------------------------------------------------------------------------------------------------------------------------------------------------- (mm) 1 2 3 4 5 6 7 8 9 Control 100a 100a 100a 100a 100a 100a 100a 100a 100a 75 100a 100a 99a 99a 98a 97a 95a 92a 83b 150 98a 98a 96a 96a 95a 93a 92a 90a 78c 225 94a 93a 92a 90a 87b 83b 80b 76b 62d 300 81b 78b 74b 71b 68c 64c 59c 53c 42e The Values are means of 4 replications of each treatment. The values followed by the same letters in each column are not statistically different at the 0.05 probability level. Shoot to Root Ratio: The shoot to root ratio of the grasses control and the 75 and 150 mm NaCl treatments (Table 5). significantly increased under salinity stress levels up to Starting on week 5 to the end of the experiment, the 225 mm NaCl compared with that of the control plants percentage of the green cover of the grasses significantly (Table 1 for the cumulative data and Table 3 for the decreased at this high level of salinity (225 mm NaCl) weekly values). There was no difference between the compared to the control and the 75 and 150 mm NaCl shoot to root ratios of the grasses at the highest level of treatments (Table 5). salinity stress (300 mm NaCl) compared to the control plants (Table 1 for the cumulative data and Table 3 for the Salt Removal by the Plants from the Culture Medium: weekly values). The quantity of the salts removed by the plants from the culture medium was estimated by subtracting the final Shoot Succulence: Shoot succulence (Fresh Wt./Dry Wt.) electrical conductivity (EC) of the culture medium from was increased as salinity stress increased from control to the initial EC values of the salinized nutrient solutions 300 mm NaCl salinity (Table 4). These values (average of (Table 6). all the harvests) were 2.19, 2.27, 2.46, 2.42 and 2.57 for the As shown in Table 6, at each NaCl salinity level, control, 75,150, 225 and 300 mm NaCl treated plants, Bermudagrass removed substantial amount of salts from respectively. the culture medium. Therefore, this halophytic grass species would be effective in biologically reclaiming Percentage of Grass Green Cover: The grass percentage desert saline soils. There was no difference between the of green cover significantly decreased at the highest quantity of the salts removed from the culture media salinity level (300 mm NaCl) compared to the control or among the 75, 150 and 225 mm NaCl treatments, indicating any other salinity level at any evaluation week (harvest) that although the levels of salinity stress were increased, (Table 5). As shown in this Table (Table 5), at the low and the grass removed the same quantities of the salts from medium salinity levels (75 and 150 mm NaCl), the the culture medium. Only at the highest level of salinity difference between the grasses, except for the last treatment (300 mm NaCl), the amount of the salts removed evaluation week (harvest), was not significant compared from the culture medium was significantly lower than that to the control plants. At the higher level of salinity stress at the 150 and 225 mm NaCl treatments, while statistically (225 mm NaCl), the difference between the grasses the same as that at the lowest level of salinity (75 mm percentage of green cover was not significant at the first NaCl). Pessarakli and Touchane [4] also reported a 4 evaluation weeks (harvests 1-4) compared with the substantial amount of salts removed from the culture 12

Table 6: Salt removal by Bermudagrass from the culture medium under various levels of NaCl application rates Electrical Conductivity (EC) ---------------------------------------------------------- Salinity (NaCl) treatment Initial (mm) Final (g/l) Salt Removed from Culture Soln. --------------------(ds/m)------------------------- ---------------------(g/l)---------------------------- Control (0) Control (0) 0.87 0.82 ------ 75 4.383 7.60 5.33 1.453ab 150 8.766 14.45 11.86 1.658a 225 13.149 21.13 18.74 1.530a 300 17.532 27.92 26.80 1.222b The Values are means of 4 replications of each treatment. The values followed by the same letters in the last column are not statistically different at the 0.05 probability level. medium by two halophytic grass species (Bermudagrass 3. Pessarakli, M. and D.M. Kopec, 2008. Establishment and Seashore paspalume). Furthermore, Pessarakli and of three warm-season grasses under salinity stress. McMillan [29] also found a substantial amount of salt Acta HortScience, ISHS, 783: 29-37. removed from the culture medium by two cultivars of 4. Pessarakli, M. and H. Touchane, 2011. Biological Seashore paspalume (Aloha and Sea Dwarf). These technique in combating desertification processes results show that, these halophytic grass species could using a true halophytic plant. Intern. J. Water Res. & be effectively used in biologically reclaiming desert saline Arid Environ., 1(5): 360-365. soils and combating desertification processes, especially 5. Ashraf, M., 1994. Breeding for salinity tolerance in in arid regions. plants. Critical Review of Plant Sci., 13: 17-42. CONCLUSIONS 6. Duncan, R.R. and R.N. Carrow, 1999. Turfgrass molecular genetic improvement for biotic/edaphic stress resistance. Advances in Agron., 67: 233-305. Shoot length was decreased as NaCl levels increased. 7. Flowers, T. and A. Yeo, 1995. Breeding for salinity Root length was stimulated by NaCl at early harvests, but resistance in crop plants: where next? Aust. J. Plant as the stress period progressed, root length decreased, Physiol., 22: 875-884. especially at the higher NaCl levels. Shoot dry weights 8. Glenn, E.P., J.J. Brown and E. Blumwald, 1999. Salt generally decreased as NaCl levels increased, but shoot tolerance and crop potential of halophytes. Critical fresh weights significantly decreased only at the highest Review of Plant Sci., 18: 227-255. NaCl level. Considering these results, performance of 9. Munns, R., S. Husain, A.R. James, A.G. Condon, Bermudagrass at these relatively high levels of salinity M.P. Lindsay, E.S. Lagudah, D.P. Schachtman and stress indicate that Bermudagrass is a very high salt R.A. Hare, 2002. Increasing salt tolerance in crops tolerant halophytic plant species that can easily tolerate and the role of physiologically-based selection traits, high salinity levels of soils and waters in the arid regions Avenues for Assessment and Management. Ann and would be a suitable grass species for cultivation Arbor Press, Chelsea, MI. Plant & Soil, 247: 93-105. under harsh arid regions and effectively combat 10. Cushman, J.C. and H.J. Bohnert, 2000. Genomic desertification processes. approaches to plant stress tolerance. Current Opinion in Plant Biology, 3: 117-124. REFERENCES 11. Holmberg, N. and L. Bulow, 1998. Improving stress tolerance in plants by gene transfer. Trends in Plant 1. Heshmati, G.A. and M. Pessarakli, 2011. Threshold Sci., 3: 61-66. model in studies of ecological recovery in 12. Isla, R., R. Aragues and A. Royo, 1998. Validity of Bermudagrass (Cynodon dactylon L.) under nutrient various physiological traits as screening criteria stress conditions. J. Plant Nutr., 34(14): 2183-2192. for salt tolerance in barley. Field Crops Research, 2. Marcum, K.B. and M. Pessarakli, 2006. Salinity 58: 97-107. tolerance and salt gland excretion activity of 13. Marcum, K.B. and M. Pessarakli, 2010. Salinity Bermudagrass turf cultivars. Crop Sci. Soc. Amer. J., tolerance of Ryegrass turf cultivars. HortScience, 46(6): 2571-2574. 45: 1882-1884. 13

14. Marcum, K.B., M. Pessarakli and D.M. Kopec, 2005. 22. Lee, G.J., R.N. Carrow and R.R. Duncan, 2004. Salinity Relative salinity tolerance of 21 turf-type desert tolerance of selected Seashore paspalums and Saltgrasses compared to Bermudagrass. HortScience, Bermudagrasses: root and verdure responses and 40(3): 827-829. criteria. HortScience, 39(2): 1136-1142. 15. Pessarakli, M., 2011. Saltgrass, a high salt and 23. Pessarakli, M. 2007. Growth responses of drought tolerant species for sustainable agriculture in Bermudagrass to various levels of nutrients in culture desert regions. Intern. J. Water Res. & Arid Environ., medium. In: Handbook of Turfgrass Management 1(1): 55-64. and Physiology (M. Pessarakli, Ed.), pp: 57-63, CRC 16. Pessarakli, M. and H. Touchane, 2011. Biological Press, Florida. technique in combating desertification processes 24. Pessarakli, M. and D.M. Kopec, 2004. Growth using a true halophytic plant. Intern. J. Water Res. & responses of Bermudagrass and Seashore paspalum Arid Environ., 1(5): 360-365. to different levels of FerroGrow multi-nutrient 17. Pessarakli, M. and D.M. Kopec, 2005. Responses of fertilizer. J. Food, Agric. & Environ. (JFAE), 2(3&4): twelve inland Saltgrass accessions to salt stress. 284-286. USGA Turfgrass & Environ. Research Online 4(20): 1-25. Hoagland, D.R. and D.I. Arnon, 1950. The water- 5. http://turf.lib.msu.edu/tero/v02/n14.pdf culture method for growing plants without soil. 18. Pessarakli, M. and D.M. Kopec, 2009. Screening California Agric. Experi. Station, Circular, pp: 347. various Ryegrass cultivars for salt stress tolerance. J. 26. SAS Institute, Inc. 1991. SAS/STAT User s guide. Food, Agric., & Environ. (JFAE), 7(3&4): 739-243. SAS Inst., Inc., Cary, NC. 19. Pessarakli, M.K.B. Marcum and D.M. Kopec, 2005. 27. Pessarakli, M., 2014. Physiological responses of Growth responses and nitrogen-15 absorption of Cotton (Gossypium hirsutum L.) to salt stress. In: desert Saltgrass (Distichlis spicata) to salinity stress. rd Handbook of Plant and Crop Physiology, 3 J. Plant Nutr., 28(8): 1441-1452. Edition, Revised and Expanded (M. Pessarakli, Ed.) 20. Pessarakli, M., N. Gessler and D.M. Kopec, 2008. pp: 635-654, CRC Press, Florida. Growth responses of Saltgrass (Distichlis spicata) 28. Greenway, H. and R. Munns, 1980. Mechanisms of under sodium chloride (NaCl) salinity stress. USGA salt tolerance in non-halophytes. Annual Review of Turfgrass & Environ. Research Online, Oct. 15, 2008, Plant Physiol., 31: 149-190. 7(20): 1-7. http://turf.lib.msu.edu/tero/v02/n14.pdf 29. Pessarakli, M. and D.E. McMillan, 2014. Seashore 21. Pessarakli, M., D.M. Kopec and D.T. Ray, 2011. paspalum, a high salinity stress tolerant halophytic Growth responses of various Saltgrass (Distichlis plant species for sustainable agriculture in desert spicata) clones under salt stress conditions. J. Food, regions and combating desertification. Intern. J. Agric., & Environ. (JFAE), 9(3&4): 660-664. Water Res. & Arid Environ., 3(1): 35-42. 14