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Communications in Soil Science and Plant Analysis
ISSN: 0010-3624 (Print) 1532-2416 (Online) Journal homepage: http://www.tandfonline.com/loi/lcss20
Changes in antioxidant enzyme activity in
turfgrass cultivars under various saline water
irrigation levels to suit landscapes under arid
regions
Shyam S. Kurup, Mohammed Abdul Mohsen Ali Salem, Abdul J. Cheruth,
Subramaniam Sreeramanan, Fayas Thayale Purayil, Abdel Wahed Al Amouri
& Mohammad Pessarakli
To cite this article: Shyam S. Kurup, Mohammed Abdul Mohsen Ali Salem, Abdul J. Cheruth,
Subramaniam Sreeramanan, Fayas Thayale Purayil, Abdel Wahed Al Amouri & Mohammad
Pessarakli (2017): Changes in antioxidant enzyme activity in turfgrass cultivars under various saline
water irrigation levels to suit landscapes under arid regions, Communications in Soil Science and
Plant Analysis, DOI: 10.1080/00103624.2017.1395451
To link to this article: http://dx.doi.org/10.1080/00103624.2017.1395451
Accepted author version posted online: 23
Oct 2017.
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Download by: [UAE University]
Date: 25 October 2017, At: 08:16
Changes in Antioxidant Enzyme Activity in Turfgrass Cultivars
under Various Saline Water Irrigation Levels to Suit Landscapes
under Arid Regions
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Shyam S. Kurup, 1 Mohammed Abdul Mohsen Ali Salem, 1 Abdul J. Cheruth, 1
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Department of Aridland Agriculture, College of Food and Agriculture, UAE University,
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P.O. Box 15551, Al-Ain, United Arab Emirates
School of Biological Sciences, Universiti Sains Malaysia (USM), 11800, Georgetown,
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Penang, Malaysia
School of Plant Sciences, Forbes Bldg., Room 303, The University of Arizona, Tucson,
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Arizona, 85721 USA.
Address Correspondence to Mohammad Pessarakli: pessarak@email.arizona.edu
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Mohammad Pessarakli3*
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Sreeramanan Subramaniam, 2 Fayas Thayale Purayil, 1 Abdel Wahed Al Amouri1 and
ABSTRACT
Turfgrass cover is an inevitable component in the urban landscapes of the United Arab
Emirates. Tolerance to abiotic stress like salt, drought and high temperature is a potential
factor to be considered in the selection of turfgrass for the landscapes in the arid regions.
Three Seashore paspalum genotypes, four Bermudagrass cultivars along with tall fescue as a
control were screened for enzymatic changes under four different saline irrigation levels of
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5,000, 10,000, 15,000 and 20,000 mg L-1 of salinity. Irrigation with potable water served as
the control. Turfgrasses were maintained in a field experiment under factorial randomized
block design for a period of two years under sustained saline conditions by working out the
leaching fraction to maintain the precise level of salinity in each treatment throughout the
experimental period. The activities of antioxidant enzymes such as superoxide dismutase
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(SOD), ascorbate peroxidase (APX), peroxidase (POD), polyphenol oxidase (PPO) and
catalase (CAT) were assayed in each of the saline water treated turfgrass types. The SOD
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paspalum group, Salam and Sea Dwarf showed the highest SOD activity under saline
condition. In Bermuda types, Yukon and SR 9554 showed the highest SOD activity
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compared to other bermuda types. Compared to other cultivars, Sea Isle 2000 (Paspalum
type), SR 9554 and Yukon (Bermuda types) exhibited more APX activity without any
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significant differences among themselves. There was an elevated activity of peroxidase
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initially, followed by gradual reduction upon increasing the salinity level. Salam cultivar
maintained stable POD activity even at the elevated salinity levels. The paspalum types
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showed relatively high level of PPO activity compared to other types under salinity. Yukon
cultivar showed a significantly higher activity of PPO when the salinity level was raised from
15,000 to 20,000 mg L-1. A significant increase with respect to CAT activity was exhibited in
Sea Dwarf under elevated level of salinity. Saline water irrigation brought about a significant
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activity was found to be high in Yukon (bermudagrass) and Sea Dwarf paspalum. In the
effect on the antioxidant enzyme systems to impart oxidative stress tolerance in turfgrass
species.
Keywords: Turfgrass cultivar; Salinity; Antioxidant enzymes
INTRODUCTION
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Turfgrass is an important part of the landscaping industries that act as a vegetative ground
cover. Turfgrass is monocot classified under the family Poaceae, which includes more than
800 genera and over ten thousand species (Piperno and Sues, 2005). Turfgrass plays a major
role in the beautification of garden, parks, playing grounds and lawns. Based on the
ecological adaptations and geographical distribution, turfgrasses were categorized as tropical
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warm season turfgrasses and cool season turfs (Chapman and Peat, 1992). The scarcity of
water and salinity in the arid regions remains the major constraints for agricultural
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productivity in all over the world despite of its irrigation level. Majority of the crop plants are
under the glycophytes, which cannot survive under salt condition. Under salinity stress,
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plants experience various physiological and metabolic changes, ultimately leads to reduced
crop production (Munns, 2005; Rozema and Flowers, 2008).
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In arid regions, the limited fresh water availability and fresh water conservation
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efforts lead to the usage of non-potable water such as recycled and domestic water for the
turfgrass management (Marcum, 2004). The presence of salt in non-potable water and the
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poor soil conditions will lead to reduced plant growth. During salinity stress, turfgrasses
undergo various changes in the physical and biochemical processes viz, reduced water
uptake, reduced nutrient uptake, increased root biomass and reduction in plant growth by
hindering photosynthesis (Harivandi et al. 1992).
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developments. Salinity is one of the major stress conditions limiting the plant growth and
Studies revealed that different environmental stresses result in damages to plants
through the increased production of reactive oxygen species (ROS) which ultimately leads to
plant death (Thompson et al. 1987; Larson, 1988; Price and Hendry 1989; Smirnoff, 1993).
Salinity stress is one of the major stresses which can induce the accumulation of ROS in
plants. The ROS accumulation leads to various changes in the plant system such as damaging
the photosynthetic components, inactivating protein and enzymes, destroying cell membrane
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structure and permeability by causing lipid peroxidation (Price and Hendry 1987; Winston,
1990). The scavenging of ROS through enzymatic break down of ROS is considered as one
of the efficient mechanisms for stress tolerance. To combat against this oxidative stresses,
plants possess various antioxidant mechanisms which include Non-enzymatic antioxidants
(β-carotenes, ascorbic acid (AA), α-tocopherol (α-toc), reduced glutathione (GSH) and
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antioxidant enzymes (superoxide dismutase (SOD), guaiacol peroxidase (POD), ascorbate
peroxidase (APX), catalase (CAT), polyphenol oxidase (PPO) and glutathione reductase
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The effect of salinity on antioxidant enzyme system was reported in turfgrass earlier.
In kikuyugrass (Pennisetum clandestinumHochst. ex Chiov.), a salinity stress of 100 mM
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sodium chloride (NaCl) leads to reduced level of APX, CAT, and Dehydroascorbate
Reductase (DHAR) activity (Muscolo et al. 2003). Some of the cultivars or ecotypes of the
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ryegrass and bermudagrass showed higher activities of antioxidant enzymes such as SOD,
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CAT, APC, and DHAR when they were subjected to salinity treatments (Hu et al. 2011; Hu
et al. 2012). An increase in NaCl concentration from 20 mM to 80 mM caused a greater
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reduction in activities of SOD, APX, GR and CAT in Kentucky bluegrass compared with
non-salt control (Arghavani et al. 2012). The results suggest that maintenance of antioxidant
enzyme activity promote salinity tolerance.
The relationship between salt stress tolerance and antioxidant level has been studied
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(GR)) (Xu et al. 2008).
in different plant species (Gossett et al. 1994; Dionisio-Sese and Tobita, 1998; Hernandez et
al. 1999). Studies revealed that salt tolerant species have the ability to survive under salt
stress by increasing their antioxidant enzyme level compared to salt sensitive species (Shalata
et al. 2001; Demiral and Türkan 2005).
The objective of this study was to evaluate the antioxidant potential of turfgrass
species used for urban landscaping in the United Arab Emirates.
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MATERIALS AND METHODS
Experimental Site
The experiment was conducted at the Zayed International Agricultural Environment Program
(ZIAEP) of the UAE University located at Nashela, Al-Ain, Abu Dhabi, UAE. This region
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lies in the arid zone 24.2075° N, 55.7447° E under desert ecosystem.
Plant Materials
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The experiment was conducted with eight turfgrass cultivar/types consisting of Tall fescue
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and SR 9554, Tifgreen, Princess 77 and Yukon bermudagrass cultivars. The characters of
Bermudagrass (Cynodon dactylon L.)
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each turfgrass types are given below:
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Bermudagrass is a warm season grass which is relatively cold tolerant. The grass is sensitive
to shade and is suitable forlawns and sports turfs. It is medium to fine textured with a very
color.
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high shoot density. These grass types exhibit rapid growth rate with medium to dark green
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Yukon: It is cold tolerant compared to other types and has excellent salt, heat and drought
tolerance. This cultivar has high wear tolerance and color retention.
Princess 77: This cultivar is characterized by a high shoot density and is fine textured. It
produces dark green lawns and turf.
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(salt sensitive control), Salam, SeaIsle 2000, and Sea Dwarf Seashore paspalum genotypes,
SR 9554: It is fine textured with a high shoot density. It has shorter inter-nods compared to
the common Bermuda and is reported to be relatively salt tolerant.
Tifgreen: This grass exhibits fairly high tolerances to salinity with narrow soft leaf blades. It
has shorter inter-nods thus with standing intensive mowing.
Paspalum (Paspalum wagenitum Swartz)
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These are warm season perennial grass types. They tolerate very high salinity levels and their
morphological characters make them attractive turfgrass. They are rapidly spreading with
stolons and rhizomatous growth characterized by dark green color.
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Salam
Salam Seashore paspalum is a warm-season perennial grass type spreading by stolons.
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Regular mowing at ˂ 1inch height produces dense turf. It has outstanding salinity tolerance
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Sea Dwarf
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It is a warm season turfgrass with a fine texture characterized by bright green color. It has the
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potential to tolerate a wide range of mowing heights and is highly salt tolerant.
Sea Isle 2000
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This grass is characterized by a fine texture and playability, when mowed at 3mm height.
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With regular verti-cutting, light topdressing and periodic rolling, Sea Isle 2000 will be an
excellent golf turf with ability to tolerate high salinity.
Tall fescue (Festuca arundinacea)
Tall fescue is comparatively deep rooted, bunch type grass, adapted to cool, humid climates.
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under prolonged drought periods.
Tall fescues grow fairly well on soils with a pH of 5.5 to7.0 and low in fertility. This grass
was used as an absolute control for the experiment since it is saline sensitive.
Greenhouse propagation
The turfgrass types included in this research program were brought from the University of
Arizona and tested under identical climatic and soil conditions of the UAE. Plant materials
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were planted in a propagation program before they were transplanted in the experimental site.
The plant propagation was carried out under greenhouse condition to acclimatize the grass to
the changed climatic condition and to get the grass material established. Newly planted
turfgrass materials had produced stolons and rhizomes indicating a rapid spread of the grass
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under greenhouse condition.
Saline Irrigation Treatments
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Five (5) levels of saline water irrigation treatments viz., 5,000, 10,000, 15,000, and 20,000
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establishment under field conditions, saline water irrigation was carried out using micro
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sprinklers (Figures 1 & 2). Irrigation was applied twice during summer and once during
winter with half inchm-2. The leaching fraction was worked out to maintain the saline water
Depth of water leached below the root zone
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Leaching Fraction =
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irrigation level constant by applying the following formula.
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Depth of water applied at the surface
Percentage of Leaf Firing
Leaf firing percentage was estimated monthly and the data where pooled to get a final
percentage of leaf firing in each turfgrass under different salinity treatments.
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mgL-1with a control (potable water) were tested under field condition. Following initial plant
Antioxidant Enzyme Extractions and Assays
Superoxide Dismutase Assay
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Crude enzyme extract was prepared based on the method proposed by Hwang (1999). The
SOD activity was assayed as described by Beauchamp and Fridovich (1971) . The reaction
solution contained 1.17×10−6molL−1 riboflavin, 0.1 molL−1 methionine, 2×10−5molL−1 KCN
and 5.6×10−5molL−1 nitrobluetetrazolium (NBT) salt dissolved in 3 ml of 0.05 molL−1 sodium
phosphate buffer (pH 7.8). One (1) mL of the extracted enzyme solution was added to 3 mL
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of the reaction solution. The mixtures were illuminated under Philips 40W fluorescent tubes
at 30ºC for 1 hr. The absorbance was read at 560 nm in the spectrophotometer against the
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absorbance per hour per mg protein.
Ascorbate Peroxidase Assay
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Ascorbate peroxidase assay was performed as described by Asada and Takahashi (1987). For
extraction, 500 mg of fresh plant tissue was ground in a pestle and mortar under liquid
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nitrogen and 10 mL of 50 mM potassium phosphate buffer (pH 7.0) containing 1 mM
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ethylenediaminetetraacetic acid (EDTA), 1% PVP and 1 mM ascorbic acid. The homogenate
was filtered through adouble-layered cheese cloth and centrifuged at 15,000rpm for 20 min at
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48ºC. The supernatant was collected for further reaction. One (1) mL of reaction mixture
containing 50 mM potassium phosphate buffer (pH 7.0), 0.5 mM ascorbic acid, 0.1mM
hydrogen peroxide (H2O2) was mixed with 200 µl of enzyme extract. The absorbance was
read as a decrease at 290nm against the blank, correction was done for the low, nonenzymatic oxidation of ascorbic acid byH2O2 (extinction coefficient 2.9 mM-1 cm-1). The
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blank. SOD activity is expressed in Umg−1 protein. One (1) U is defined as the change in 0.1
enzyme activity was expressed in units mg-1 protein (U = change in 0.1 absorbance min-1 mg-
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protein).
Peroxidase (POX) Assay
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POX was assayed based on the method of Kumar and Khan (1982). The POX assay solution
mixtures contained 2 mL of 0.1 molL-1 phosphate buffer (pH 6.8), 1 mL of 0.01 molL-1
pyrogallol, 1 mLof 0.005 molL-1 H2O2 and 0.5 mL of enzyme extract. 5 min incubation at
25ºC was performed and the reaction was terminated by mixing 1 mL of 2.5molL-1 sulfuric
acid (H2SO4). The amount of purpurogallin formed was determined by measuring the
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absorbance at 420 nm. The blank was prepared by mixing the enzyme extract with 2.5 molL-1
H2SO4. The activity was expressed in unit/mg protein.
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The assay of PPO was carried out as described by Kumar and Khan (1982). The PPO Assay
mixture contained 2 mL of 0.1 M phosphate buffer (pH 6.0), 1 mL of 0.1 M catechol and 0.5
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mL of enzyme extract. This mixture was incubated at 25ºC for 5 min, immediately
thereaction was stopped by adding 1 mL of 2.5 N H2SO4.The absorbance of the purpurogallin
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formed was read at 495 nm. The blank was prepared by adding 2.5N H2SO4to the same assay
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mixture without incubation. PPO activity is expressed in U/mg protein.
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Catalase (CAT) Assay
The activity of CAT was carried out based on method described by Chandlee and Scandalios
(1984) with small modification. Total of 500 mg of frozen material was washed and
homogenized in 5 mL of ice-cold 50 mM sodium phosphate buffer (pH 7.5) containing 1 mM
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Polyphenol oxidase (PPO) Assay
PMSF. The extract was centrifuged at 48ºC for 20 min at 12,500 rpm. The supernatant was
used for enzyme assay. The assay mixture contained 2.6 ml of 50 mmolL-1 potassium
phosphate buffer (pH 7.0), 0.4 mL of 15 mmolL-1 H2O2 and 0.04 ml of enzyme extract. The
decomposition of H2O2 was followed by the decline in absorbance at 240 nm. The enzyme
activity was expressed in Umg-1 protein.
Statistical Analysis
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The experimental design was Factorial random block design (RBD) consisting of 8 turfgrass
types and four levels of saline water irrigation and one control. The treatments were
replicated four times. Three samples were collected from each turf grass types under different
saline water irrigation levels and pooled. The mean data were compared using Duncan
Multiple Range Test (DMRT) with significance at P<0.05.
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RESULTS AND DISCUSSION
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Identification of salt tolerant variety is necessary in arid region in order to combat with the
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evaluate the antioxidant potential of different turfgrass varieties. The leaf firing percentage
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and the level of five different antioxidant enzymes viz, SOD, APX, POD, POP, and CAT
were assessed in paspalum and bermudagrass varieties under different saline conditions in
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order to identify the suitable variety that can tolerate high saline soils in the arid regions.
The percentage of leaf firing was considered as an assessment for salinity tolerance as
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reported in previous studies (Marcum, 1999; Lee et al. 2004). It is one of the easily
measurable parameters to be used as an assessment method for salinity tolerance. The
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percentage of leaf firing increased significantly in all the tested varieties when the salinity
level increased (Table 1). Similar result was reported earlier by various researchers (Uddin et
al. 2009; Uddin et al. 2011) Paspalam cultivars showed relatively low leaf firing compared to
Bermuda cultivars even at high salinity levels. In some of the cultivars, there was no injuries
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soil condition. In this study different salt conditions were applied to the soil in order to
recorded (0%) up to 5,000 mgL-1 of salinity. In paspalam genotypes, Salam showed relatively
low leaf firing (15.2%) at 20,000 mg L-1salinity compared to other genotypes. In
bermudagrass cultivars, SR9554 and Yukon varieties showed lower leaf firing percentage at
20,000mg L-1 salinity. Among all tested cultivars, tall fescue showed a potential increase in
leaf firing percentage (92.6%) at 20,000 mg L-1 salinity. In the previous reports, the
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percentage of leaf firing was low in paspalm genotypes compared to bermudagrass cultivars
(Uddin et al. 2012; Zulkaliph et al. 2013). Our results are in agreement with these findings.
Superoxide Dismutase
Superoxide dismutase acts as a major scavenger of reactive oxygen species (ROS) under
stress. SOD initiate the defense mechanism by converting the oxygen (O2−) radicals to
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H2O2and molecular oxygen O2 (Meloni et al. 2003). The bermudagrass type, Yukon showed
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the highest SOD level under salt stress (15,000 mg L-1) which is on par with the Sea Dwarf
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showed a gradual increase in SOD level by increasing salt concentrations and then decreased
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at 20,000 mg L-1. A significantly high increase was noticed at 15,000 mg L-1salinity level in
both cultivars. In SR 9554, SOD level was found to be high at 5,000 mg L-1 and a gradual
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decrease was found under high saline condition. In Sea Isle 2000, the SOD level was found to
be similar in all treatments. Compared to the control Tall fescue, the Bermudagrass cultivars
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(Yukon and SR 9554) and paspalum cultivars (Sea Dwarf and Salam) showed higher level of
SOD. Difference in SOD activity had also been reported in potato cultivars under NaCl
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salinity condition (Rahnama and Ebrahimzadeh 2005). Hu et al. (2012) reported a significant
increase in SOD level in salt tolerant Bermudagrass genotype C43when compared to salt
sensitive genotype C198.An increase in SOD activity under salt stress was reported in cotton
(Meloni et al. 2003). Increases in SOD activity in plants could protect against oxidative stress
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paspalum at 10,000 mg L-1 of salinity (Figure 3). Salam and Yukon bermudagrass cultivars
(Sen Gupta et al. 1993). In finger millet cultivars, 0.2 to 1.5fold increase in SOD activity was
observed under salt stress (Anjaneyulu et al. 2013). Enhanced tolerance to salt stress was
demonstrated by over-expression of mitochondrial Mn-SOD in Arabidopsis thaliana and
chloroplast related Cu/Zn SOD in Nicotiana tabacum (Wang et al. 2004; Badawi et al. 2004).
Increase in SOD activity means there is increased release of reactive oxygen species which
activates the gene responsible for SOD (Abedi and Pakniyat 2010).
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Ascorbate Peroxidase
The ascorbate peroxidase (APX) level was found to be higher in all the tested cultivars
compared to the control tall fescue (Figure 4). Sea Isle 2000 showed the highest APX level at
20,000 mg L-1 salinity, which is on par with the Bermudagrass types, SR 9554 (10,000 mg L1
) and Yukon (20,000 mg L-1). In Sea Isle 2000, a gradual increase in APX was observed and
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it rose significantly at 20,000 mg L-1 salinity. It shows Sea Isle 2000 cultivar has considerably
high antioxidant action even at severe saline condition. Next to Sea Isle 2000, SR 9554
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APX level gradually and it showed significantly high level at 20,000 mg L-1salinity level. The
response of tall fescue was found to be lower in all the treatments. Salam and Sea Dwarf
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showed an increase in APX level at 5,000 and 15,000 mg L-1, respectively.
In Bermudagrass, salt sensitive ‘C198’ genotype showed 37%increase in APX
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activity than salt tolerant ‘C43’ genotype under 400 mM NaCl (Hu et al. 2012). Yasara et al.
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(2008) reported a significant increase in APX activity of salt tolerant green bean ‘GS57’
compared to salt sensitive ‘cv. 4F-89’. Compared to other cultivars, Sea Isle 2000 and Yukon
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showed a significant change in the APX level under increased salt stress. Sai Kachout et al.
(2013) also reported elevated levels in APX activity due to salt stress in two Atriplex
varieties.
High SOD level is not enough to protect the plant against the damage caused by
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showed an increase in APX action at 10,000 mg L-1 salinity. Yukon displayed an increase in
superoxide radicals as it leads to the production of H2O2 which are harmful to the cells. These
generated H2O2 need to be scavenged quickly by antioxidant enzymes such as peroxidase and
catalase (Guo et al. 2006). H2O2 production resulted from SOD activity derived through
ascorbate–glutathione cycle (Willekens et al. 1994; Asada 1999; Mittler 2002) and thereby
oxidation re-reduction of ascorbate and glutathione by the ascorbate peroxidase (APX) and
glutathione reductase (GR) enzymes takes place (Foyer and Halliwell 1976; Noctor and
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Foyer 1998). Ascorbate which is used as electron donor by ascorbate peroxidase may
eliminate oxygen free radicals to its reduced form (Karyotuou and Donaldson 2005; Zhang
2013).
Peroxidase
Peroxidase enzymes play major role in scavenging the hydrogen peroxide produced under
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salt stress (Dionisio-Sese and Tobita 1998). The peroxidase activity significantly increased in
all the cultivars under salt stress and then gradually decreased upon increasing the salinity
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which is on par with the Yukon cultivar under same salinity condition.
A gradual decline in peroxidase level was observed in both cultivars by further
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increase in salt stress. In Sea Dwarf, a gradual increase in peroxidase activity was noticed and
it was found to be high at 10,000 mg L-1 salinity level. Salam cultivar confirmed an increase
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in peroxidase activity at 10,000 mg L-1and the level was maintained up to 20,000 mg L-1
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salinity level without any significant change, which shows Salam cultivar has a constant
expression of peroxidase action under increasing salt stress compared to all the other
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varieties. Both Tall fescue and Tifgreen maintained low levels of peroxidase compared to
other cultivars. Costa et al. (2005) reported that the peroxidase activity of salt the sensitive
sorghum genotype showed a significant decrease compared to that of the salt tolerant
genotype. In wheat, an increase in POD level was observed in the salt tolerant S-24 cultivar
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level (Figure 5). Sea Isle 2000 displayed an elevated peroxidase activity at 10,000 mg L-1,
compared to the salt sensitive DN-27 cultivar (Bhutta 2011). Cicek and Cakirlar (2008)
reported a decreased in POD activity under different NaCl treatments for different soy
cultivars, except for cultivar Nazlican which showed an increase in POD activity. An increase
in POD activity was observed in Cakile maritime with increasing concentrations of NaCl, but
the activity decreased when the concentration of NaCl increased beyond 40 mmolL-1. Jabeen
and Ahmad (2012) also reported increased POD activity due to salt stress in safflower and
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sunflower seedlings. Bor et al. (2003) also reported increased POD under salt stress in Beta
vulgaris and B. maritima. Foyer et al. (1994) gave an explanation that the ROS may have
played a role in the signal transduction which leads to decrease in the production of
antioxidant enzymes. Varying transcript levels and enzyme activities during salt stress may
attribute to the high turnover of these enzymes or H2O2 inactivation of these enzymes
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(Scandalios 1993).
Polyphenol oxidase (PPO)
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PPO has been studied in many plants in connection with wounding, and browning (Demir
and Kocaliskan 2001) and in alkaloid biosynthesis under biotic and abiotic stress conditions
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(Bilková et al. 2005). The PPO level was varied significantly in each cultivar of the turfgrass
tested (Figure 6). The paspalum cultivars showed relatively the same level of PPO activity,
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but under different saline conditions.
Both Salam and Sea Dwarf showed a significant increase in PPO activity at 10,000
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mg L-1 salinity level, whereas in Sea Isle 2000, the PPO level was found to be higher at
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15,000 mg L-1salinity level (Figure 5). In Tifgreen, a significant increase in PPO activity was
observed at 10,000 mg L-1salinity level and it showed a gradual reduction in PPO activity
under high salt condition. Weisany et al. (2012) reported a significant increase in PPO
activity in leaf and roots of soybean plants treated under different salt concentrations. Except
Yukon, most of the cultivars showed an elevation in PPO activity at 10,000 to 15,000 mg L-1
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Polyphenol oxidase enzymes are involved in oxidation of phenols to chinone. The role of
salinity levels and it was reduced significantly at 20,000 mg L-1 salinity level. Yukon showed
an increase in PPO activity at 20,000 mg L-1 salinity level. Tall fescue displayed a minimal
level of PPO activity compared to other cultivars. An increase in PPO activity under salt
stress was also reported in bean seedlings (Demir and Kocaliskan 2001). Enhanced PPO
14
under salt stress was also observed in Sesuvium portulacastrum (Rajaravindran and Natarajan
2012), and fenugreek (Pour et al. 2013).
Catalase
Catalase and peroxidase are two groups of enzymes mainly involved in detoxification of
H2O2. Compared to peroxidase, catalase has low affinity towards H2O2 and thus catalase is
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suggested to be involved in mass scavenging of H2O2, whereas peroxidase plays its fine
regulation (Willekens et al. 1994; Mittler 2002); Noctor and Foyer 1998).
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Bermudagrass types under salt stress (Figure 7). Sea Dwarf displayed a maximum catalase
activity at 20,000 mg L-1 salinity level. This showed Sea Dwarf has elevated antioxidant
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potential even at high salinity condition. Next to Sea Dwarf, Salam showed a significant
increase in catalase level at 5,000 mg L-1 salinity level and a gradual decrease was observed
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under high salinity. Similar trend was also observed in wheat cultivars (Alvand and Sardari)
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whereby salt stress induced similar CAT activity in both cultivars and showed increased
activity at 50 mM NaCl which further decreased at 200 mM NaCl (Esfandiari et al. 2007).
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CAT activity increased with increased concentration of NaCl in tobacco genotypes
(Hatamnia et al. 2013). In Yukon, a gradual increase in catalase activity was observed and it
reached the highest level at 15,000 mg L-1 salinity level. In two genotypes of Bermudagrass,
Hu et al. (2012) reported a significant increase in CAT activity in new leaves of ‘C198’ over
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The catalase activity found to be significantly high in paspalum cultivars compared to
‘C43’ genotype. Whereas in old leaves, ‘C43’ had higher CAT activity compared to ‘C198’
under moderate and high doses of salt, respectively. Tall fescue and Tifgreen maintained a
minimal catalase activity compared to the other cultivars.
15
CONCLUSIONS
The antioxidant enzyme activity significantly influenced different cultivars of turfgrass in
combating oxidative stress under different salinity regimes. These responses were expressed
in the morphological appearance of the grass in terms of expression of color, tolerance to leaf
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t
firing due to salinity stress and also in maintaining the density of the grass. The above factors
cr
are considered potential morphological traits in deciding quality turf for golf courses under
us
suitable for urban landscaping and golf courses in the UAE based on the different parameters
an
studied in the order of their priority are Salam, Sea Dwarf and Sea Isle 2000 Paspalum
genotypes for high saline areas. Some of the Bermuda varieties like SR 9554 and Yukon can
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also be considered for less challenging situations.
ACKNOWLEDGEMENT
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The financial support received from the Emirates Foundation to carry out this research is
greatly acknowledged. The greenhouse facilities and the field allotment received from
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pt
ZIAEP, Nashella and the laboratory facilities in the College of Food and Agriculture, UAEU
is acknowledged. The technical collaboration received from the University of Arizona, USA
is greatly appreciated. We extend our thanks to Dr. Ghaleb Ali Alhadrami the former Dean,
College of Food and Agriculture, currently Provost, the UAEU for providing research
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arid hostile environment, especially in challenging situations of salinity. The best varieties
infrastructure and support.
16
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CONFLICT OF INTEREST
“The authors (Shyam S. Kurup, Mohammed Abdul Mohsen Ali Salem, Abdul J. Cheruth,
Sreeramanan Subramaniam, Fayas Thayale Purayil, Abdel Wahed Al Amouri and
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Mohammad Pessarakli) declare that they have no conflict of interests.”
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Table 1. Effect of different salinity levels on leaf firing (%) of tested turfgrass cultivars
Salinity Concentration (mg L-1)
5,000 10,000 15,000 20,000
Salam
0
0
2.2c
5.1b
15.2a
SeaDwarf
0
0
3.1c
6.8b
18.5a
SeaIsle 2000
0
0
2.2c
6.4b
19.5a
SR 9554
0
1.1d
4.3c
12.5b
40.5a
Tall fescue
0
20d
30.2c
59.8b
92.6a
Tifgreen
0
20.4b
an
50.8a
0
1.2d
6.5c
13b
62.2a
0
0
6.5c
12.9b
42.2a
ed
Yukon
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Princess 77
10.6c
0
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Genotype/Cultivar 0
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Means represented with common letters are not significantly different at P ≤ 0.05 by LSD
test.
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Grass
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Figure 1. Experimental field
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Figure 2. Irrigation infrastructure
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Figure 3. Superoxide dismutase activity in the plant as influenced by the salinity
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treatments (U/mg)
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Figure 4. Ascorbate peroxidase activity in the plant as influenced by the salinity
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treatments (U/mg)
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Figure 5. Peroxidase activity in the plant as influenced by the salinity treatments
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(U/mg)
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Figure 6. Polyphenol oxidase activity in the plant as influenced by the salinity
cr
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treatments (U/mg)
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Figure 7. Catalase activity in the plant as influenced by the salinity treatments (U/mg)
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