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Xunzhong Zhang and Erik H. Ervin

Ultraviolet-B (280–320 nm) radiation is one of the major factors causing quality decline of transplanted turfgrass sod. Information on physiological parameters for assessing turfgrass tolerance to ultraviolet-B stress is lacking. The objective of this study was to evaluate ultraviolet-B tolerance of four cool-season turfgrass species and varieties using selected physiological parameters under artificial ultraviolet-B radiation stress. The physiological performance of 18 varieties of tall fescue (TF; Festuca arundinacea Schreb.), chewings fescue (CF; Festuca rubra L. ssp. commutata Gaud.), perennial ryegrass (PRG; Lolium perenne L.), and kentucky bluegrass (KBG; Poa pratensis L.) were subjected to continuous, artificial ultraviolet-B radiation (70 μmol·m−2·s−1) for 10 days. Visual quality ratings of TF, CF, PRG, and KBG measured at Day 10 were reduced by 49%, 18%, 51%, and 74%, respectively, relative to that at trial initiation. Ultraviolet-B tolerance in CF was the greatest, KBG was the least, and TF and PRG were intermediate. ‘Ebony’ was the most ultraviolet-B-tolerant TF variety, whereas ‘BlueTastic’, ‘BlueRiffic’, and ‘747’ had greater ultraviolet-B tolerance than the other four KBG varieties. No differences were observed in ultraviolet-B tolerance between varieties in either CF or PRG. The ultraviolet-B-tolerant species had less electrolyte leakage (EL), greater canopy photochemical efficiency (PEc), and relatively smaller and slower upregulation in antioxidant superoxide dismutase (SOD) activity relative to ultraviolet-B-sensitive ones. The results suggest that EL, PEc, and SOD may be used as physiological parameters in selecting ultraviolet-B-tolerant species and varieties for sod production and lawn establishment.

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Xunzhong Zhang, Kehua Wang and Erik H. Ervin

Recent advances in bermudagrass [Cynodon dactylon (L.) Pers. var. dactylon] breeding and cultural management practices have enabled its use as a sports surface in U.S. Department of Agriculture cold hardiness zones 5 and 6. Use of these more cold-hardy bermudagrass cultivars further into transition- and cool-season zones increases the probability of freezing injury and increases the need for an improved understanding of physiological responses to chilling and freezing temperatures. Abscisic acid (ABA) has been shown to increase during cold acclimation (CA) and play a role in dehydration tolerance. This study investigated changes in ABA metabolism and dehydrin expression during CA and their association with freezing tolerance in four bermudagrass cultivars. Two cold-tolerant (‘Patriot’ and ‘Riviera’) and two relatively cold-sensitive (‘Tifway’ and ‘Princess’) cultivars were either subjected to CA at 8 °C day/4 °C night with a light intensity of 250 μmol·m−2·s−1 over a 10-h photoperiod for 21 days or maintained at 28 °C day/24 °C night over a 12-h photoperiod. In a separate study, exogenous ABA at 0, 50, 100, and 150 μm was applied to ‘Patriot’ bermudagrass without CA. ABA content in leaf and stolon tissues increased substantially during the first week of CA and remained relatively stable thereafter. ‘Patriot’ and ‘Riviera’ had greater ABA content and less stolon electrolyte leakage (EL) relative to ‘Tifway’ and ‘Princess’. Expression of a 25 kDa dehydrin protein increased during CA in all four cultivars. A significant correlation was found between ABA content and freezing tolerance. Exogenously applying ABA to ‘Patriot’ at 50, 100, and 150 μm significantly increased endogenous ABA content and the 25 kDa dehydrin expression and reduced stolon EL. The results suggest that alteration of ABA metabolism during CA is closely associated with freezing tolerance. Selection and use of cultivars with substantial accumulation of ABA and certain dehydrins during CA or in response to exogenous ABA could improve bermudagrass persistence in transition zone climates.

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Erik H. Ervin, Xunzhong Zhang and John H. Fike

High ultraviolet-B (UV-B; 290-320nm wavelength) may significantly contribute to kentucky bluegrass (Poa pratensis L.) sod death at harvest and transplanting. As terrestrial UV-B levels continue to increase due to a depletion of the stratospheric ozone layer this problem may worsen. Epidermal attenuation from pigments and detoxification of reactive oxygen species by antioxidant metabolites and enzymes are involved in plant defense against oxidative stress caused by UV-B. Our objective was to determine whether the attenuation and detoxification systems of kentucky bluegrass could be artificially boosted by exogenous applications of ascorbic acid (AA), alpha-tocopherol (AT), or a colorant before exposure to high levels of UV-B. Ascorbic acid, AT, and the colorant Green Lawnger (GL), were applied to plugs of mature kentucky bluegrass alone or in combination, and then subjected to artificial, continuous UV-B exposure (70 μmol·m-2·s-1); three greenhouse experiments were conducted. By 3 to 5 days after UV-B initiation, visual quality and photochemical efficiency, as measured by chlorophyll fluorescence were significantly reduced. However, in Expt. 1, AA alleviated decline of visual quality, delayed loss of photochemical efficiency, and increased recovery relative to the control. In Expt. 3, decreased endogenous AT and antioxidant enzyme activities were measured due to UV-B stress. Application of AA, AA + AT, or GL partially alleviated photochemical efficiency decline from 4 to 12 days after initiation of UV-B. In addition, application of the chemical treatments increased leaf tissue AT concentrations by 32% to 42%, increased SOD activity by 30% to 33% and increased catalase activity by 37% to 59%, relative to the control as measured 10 days after UV-B initiation. Greater AT concentration and SOD and catalase activities were associated with greater visual quality under UV-B stress. The results of these studies indicate that kentucky bluegrass UV-B tolerance may be increased by supplementing its pigment and antioxidant defense systems with foliar applications of AA, AT or GL.

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Erik H. Ervin, Xunzhong Zhang and John H. Fike

High ultraviolet-B (UV-B; 290-320 nm wavelength) radiation may significantly contribute to the quality decline and death of kentucky bluegrass (Poa pratensis L.) sod during summer transplanting. Antioxidants and protective pigments may be involved in plant defense against oxidative stress caused by UV-B. Selected exogenous hormones may alleviate UV-B damage by upregulating plant defense systems. The objectives of this study were to determine if exogenous hormone or hormone-like substances could alleviate UV-B damage to `Georgetown' kentucky bluegrass (Poa pratensis L.) under greenhouse conditions. The hormone salicylic acid at 150 mg·m-2 and the hormone-containing substances, humic acid (HA) at 150 mg·m-2 and seaweed extract (SWE) at 50 mg·m-2, were applied to plugs of kentucky bluegrass and then subjected to UV-B radiation (70 μmol·m-2·s-1). The UV-B irradiation stress reduced turf quality by 51% to 66% and photochemical efficiency by 63% to 68% when measured 10 or 12 days after initiation of UV-B. Endogenous alpha-tocopherol (AT) and antioxidant enzymes (superoxide dismutase (SOD) and catalase) were reduced by UV-B stress. Anthocyanin content was increased from day 1 to 5 and then decreased from day 5 to 10 of continuous UV-B irradiation. Application of SA and HA + SWE enhanced photochemical efficiency by 86% and 82%, respectively, when measured 10 or 12 days after UV-B initiation. In addition, application of the hormonal supplements increased AT concentration, SOD, catalase activity, and anthocyanin content when compared to the control at 10 days after UV-B initiation. Bluegrass with greater AT concentration and SOD and catalase activity exhibited better visual quality under UV-B stress. The results of this study suggest that foliar application of SA and HA + SWE may alleviate decline of photochemical efficiency and turf quality associated with increased UV-B light levels during summer.

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Erik H. Ervin, Xunzhong Zhang and John H. Fike

Plants possess various constitutive and inducible defense mechanisms such as pigment and antioxidant systems for protection against stresses such as ultraviolet-B (UV-B; 290 to 320 nm) radiation. Our previous research has indicated that higher chlorophyll, carotenoid, and anthocyanin concentrations were associated with greater tolerance of UV-B stress by `Georgetown' kentucky bluegrass (Poa pratensis L.). The objectives of this study were to determine if kentucky bluegrass cultivars with darker leaf color possessed greater pigment and antioxidant defense systems and if such increases were associated with greater resistance to UV-B. Eight cultivars exhibiting a range of green color intensity (`Apollo', `Brilliant', `Julius', Limerick', `Midnight', `Moonlight', `Nuglade', and `Total Eclipse') were selected and subjected to continuous, artificial UV-B radiation (70 μmol·m-2·s-1). UV-B irradiation reduced turf quality (55% to 62%) and photochemical efficiency (37% to 70%) when measured 5 days after initiation of UV-B exposure. Significant differences in turf color, photochemical efficiency, chlorophyll a, chlorophyll b, chlorophyll a+b, and carotenoids were found among the cultivars. `Moonlight' had greatest photochemical efficiency, chlorophyll, carotenoids, and turf quality. Positive correlations of pigment concentration with photochemical efficiency and turf color were observed under UV-B radiation stress, with correlation coefficients ranging from 0.49 to 0.62. The results of this study suggests that selecting cultivars with higher concentrations of chlorophyll and carotenoids and photochemical efficiency may be an effective way for turfgrass managers and sod producers to improve sod establishment and quality in environments with higher UV-B radiation.

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John H. Dunn, Erik H. Ervin and Brad S. Fresenburg

Various mixtures of tall fescue, Festuca arundinacea Schreb., Kentucky bluegrass, Poa pratensis L., and perennial ryegrass, Lolium perenne L., may be beneficial for turf culture because of genetic diversity and improved tolerance to environmental stresses compared with a single species. Turf-type tall fescue, dwarf tall fescue, Kentucky bluegrass, and perennial ryegrass were seeded as cultivar blends and in all possible combinations as species mixtures in two locations, irrigated and nonirrigated. Turf was mowed at 19 and 51 mm and subjected to an interval of brief, but intensive, simulated traffic. Perennial ryegrass was the dominant species in all mixtures with tall fescue, Kentucky bluegrass, or both. After 5 years, turf-type tall fescue comprised 62% of mixtures with Kentucky bluegrass when averaged over locations. Dwarf tall fescue comprised 48% of mixtures compared with Kentucky bluegrass at 44%. Kentucky bluegrass was more competitive with tall fescue in the irrigated vs. nonirrigated location. Mowing height effected small changes in populations year to year while simulated traffic had little effect on populations at 1 year following treatment. The advantage of mixing species compared with individual species to reduce disease occurrence was evident on several occasions. Our study supports earlier research reports that tall fescue will remain competitive in mixture with Kentucky bluegrass several years after seeding.

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Xunzhong Zhang, Wenli Wu, Erik H. Ervin, Chao Shang and Kim Harich

Plant hormones play an important role in plant adaptation to abiotic stress, but hormonal responses of cool-season turfgrass species to salt stress are not well documented. This study was carried out to investigate the responses of hormones to salt stress and examine if salt stress-induced injury was associated with hormonal alteration in kentucky bluegrass (KBG, Poa pratensis L.). The grass was grown in a growth chamber for 6 weeks and then subjected to salt stress (170 mm NaCl) for 28 days. Salt stress caused cell membrane damage, resulting in photosynthetic rate (Pn), chlorophyll (Chl), and turf quality decline in KBG. Salt stress increased leaf abscisic acid (ABA) and ABA/cytokinin (CK) ratio; reduced trans-zeatin riboside (ZR), isopentenyl adenosine (iPA), and indole-3-acetic acid (IAA), but did not affect gibberellin A4 (GA4). On average, salt stress reduced ZR by 67.4% and IAA by 58.6%, whereas it increased ABA by 398.5%. At the end of the experiment (day 28), turf quality, Pn, and stomatal conductance (g s) were negatively correlated with ABA and ABA/CK ratio, but positively correlated with ZR, iPA, and IAA. Electrolyte leakage (EL) was positively correlated with ABA and ABA/CK and negatively correlated with ZR, iPA, IAA, and GA4. GA4 was also positively correlated with turf quality and g s. The results of this study suggest that salt stress-induced injury of the cell membrane and photosynthetic function may be associated with hormonal alteration and imbalance in KBG.

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Xunzhong Zhang, Damai Zhou, Erik H. Ervin, Greg K. Evanylo, Derik Cataldi and Jinling Li

The presence of biologically active substances (BAS) in biosolids may enhance plant stress tolerance and growth, but the underlying mechanisms are not well understood. This greenhouse study investigated the effects of two biosolids: Alexandria (anaerobically digested; Class A product from the Alexandria Sanitation Authority Wastewater Treatment Facility in Alexandria, VA) and Blue Plains (lime-stabilized; Class B product from Washington, DC, Water and Sewer Authority) on tall fescue [Lolium arundinaceum (Schreb.) S.J. Darbyshire] antioxidant enzyme activity associated with drought resistance. Treatments included a fertilizer control, Alexandria (11.9 g·kg−1 soil) and Blue Plains (17.6 g·kg−1 soil) biosolids to match the nitrogen in the control. Tall fescue physiological responses were measured under well-watered or drought-stressed conditions. Drought stress reduced turfgrass quality, photochemical efficiency (PE), and catalase (CAT) activity while increasing superoxide dismutase (SOD), ascorbate peroxidase (APX), and peroxidase (POD) activities. The two biosolids improved turfgrass quality and root mass under both soil moisture regimes and delayed leaf wilting during moisture stress. The biosolids also improved PE, SOD, and APX activities relative to the control under both soil moisture regimes. The data suggest that biosolids application may improve antioxidant enzyme activity and subsequent drought resistance.

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Xunzhong Zhang, Erik H. Ervin, Yiming Liu, Guofu Hu, Chao Shang, Takeshi Fukao and Jasper Alpuerto

Water deficit is a major limiting factor for grass culture in many regions with physiological mechanisms of tolerance not yet well understood. Antioxidant isozymes and hormones may play important roles in plant tolerance to water deficit. This study was designed to investigate antioxidant enzymes, isozymes, abscisic acid (ABA), and indole-3-acetic acid (IAA) responses to deficit irrigation in two perennial ryegrass (Lolium perenne L.) cultivars contrasting in drought tolerance. The plants were subjected to well-watered {100% container capacity, 34.4% ± 0.21% volumetric moisture content (VWC), or deficit irrigation [30% evapotranspiration (ET) replacement; 28.6% ± 0.15% to 7.5% ± 0.12% VWC]} conditions for up to 8 days and rewatering for 4 days for recovery in growth chambers. Deficit irrigation increased leaf malondialdehyde (MDA) content in both cultivars, but drought-tolerant Manhattan-5 exhibited lower levels relative to drought-sensitive Silver Dollar. Superoxide dismutase (SOD) activity declined and then increased during water-deficit treatment. ‘Manhattan-5’ had higher SOD activity and greater abundance of SOD1 isozyme than ‘Silver Dollar’ under water deficit. Deficit irrigation increased catalase (CAT) and ascorbate peroxidase (APX) activity in ‘Manhattan-5’, but not in ‘Silver Dollar’. ‘Manhattan-5’ had higher CAT, APX, and peroxidase (POD) activity than ‘Silver Dollar’ during water limitation. Deficit irrigation increased mRNA accumulation of cytosolic cupper/zinc SOD (Cyt Cu/Zn SOD), whereas gene expression of manganese SOD (Mn SOD) and peroxisome APX (pAPX) were not significantly altered in response to deficit irrigation. No differences in Cyt Cu/Zn SOD, Mn SOD, and pAPX gene expression were found between the two cultivars under deficit irrigation. Water limitation increased leaf ABA and IAA contents in both cultivars, with Silver Dollar having a higher ABA content than Manhattan-5. Change in ABA level may regulate stomatal opening and oxidative stress, which may trigger antioxidant defense responses. These results indicate that accumulation of antioxidant enzymes and ABA are associated with perennial ryegrass drought tolerance. Activity and isozyme assays of key antioxidant enzymes under soil moisture limitation can be a practical screening approach to improve perennial ryegrass drought tolerance and quality.