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Bermudagrass is a warm-season turfgrass species widely used for sports fields, home lawns, and golf courses. Ultradwarf bermudagrass has been used for golf course greens, but its quality declines with abiotic stresses. This 2-year study was designed to investigate if foliar applications of seaweed extract-based biostimulant Utilize® could improve ultradwarf bermudagrass photosynthetic function, nitrate reductase activity, root growth, and root function while under heat stress and drought stress conditions. Utilize® was applied to ultradwarf bermudagrass canopy at 0, 88, 117, 175, and 234 μL⋅m−2. Two weeks after the initial application of Utilize®, bermudagrass was subjected to heat (40/36 °C, day/night) and drought stress (40–50% evapotranspiration replacement) for up to 42 days. Heat stress and drought stress caused decline of the turf quality. Foliar application of Utilize® at 117, 175, and 234 μL⋅m−2 biweekly consistently improved turf quality and leaf color ratings and increased leaf chlorophyll and carotenoid concentrations, net photosynthetic rate, nitrate reductase activity, and root growth and viability. On average, Utilize® at 117, 175, and 234 μL⋅m−2 increased turf quality ratings by 9.1%, 12.1%, and 10.6%, respectively, net photosynthetic rates by 32.4%, 45.0%, and 35.0%, respectively, and nitrate reductase activity by 16.7%, 18.8%, and 14.6%, respectively, compared with the control. Utilize® at 117, 175, and 234 μL⋅m−2 increased the root biomass, root length, surface area, and root volume compared with the control. Utilize® at 88, 117, 175, and 234 μL⋅m−2 increased root viability by 46.2%, 73.1%, 88.5%, and 74.4%, respectively, relative to the control. The results of this study suggest that seaweed extract-based biostimulant Utilize® improves nitrogen metabolism, photosynthetic function, root growth, and root viability. Foliar application of Utilize® at rates between 117 and 175 μL⋅m−2 biweekly can be considered an effective approach to improving ultradwarf bermudagrass performance under heat stress and drought stress environments.

Plant-based pigments have been used as substances to improve crop yield and quality, but the mechanisms of their action on plant growth and stress tolerance are not well understood. The objective of this study was to investigate effects of two formulations of plant-based copper chlorophyllin (Cu-Chl) with and without synthetic paraffinic oil. These formulations, referred to as B18-0074 and B18-0075, were applied as a soil drench plus foliar or a foliar-only application. We investigated their impact on physiological responses of tomato plants under prolonged drought stress conditions. In addition, we examined photosynthetic impacts associated with the application of Cu-Chl formulations. B18-0074 increased leaf photosynthetic rate (Pn) by 8.8% with soil plus foliar application and 18.6% with foliar application relative to the control under drought stress at day 21. Similarly, B18-0075 increased Pn by 16.9% with soil plus foliar application and 24.6% with foliar application relative to the control under drought stress at day 21. The application of the two Cu-Chl–containing products increased leaf antioxidant enzyme catalase (CAT) and ascorbate peroxidase (APX) activity, as well as glutathione (GSH) content. The two products also increased leaf soluble sugars and proline content, indicating improvement of osmotic adjustment. Soil plus foliar and foliar application only of B18-0075 increased root biomass but did not consistently affect plant shoot growth. The results of this study suggest that application of Cu-Chl in combination with synthetic paraffinic oil may improve photosynthetic function, osmotic adjustment, antioxidant defense capacity, and root growth of tomato plant grown under drought stress conditions.

Turfgrass performance under drought stress is impeded by plant water deficit and oxidative damage, which might be improved by the external application of osmoprotectants. Creeping bentgrass (Agrostis stolonifera L.) is a valuable species for low-cut golf surfaces as a result of its high density and fine texture. However, weak tolerance to drought stress is a primary shortcoming. In this study, the effect of exogenous glycinebetaine (GB) pretreatment on mitigating the damage from drought stress in creeping bentgrass cultivar ‘T-1’ was evaluated. Pieces of creeping bentgrass sod were subjected to four treatments: 1) well-watered control, 2) well watered and sprayed with 100 mm GB, 3) drought stress, and 4) drought stress and sprayed with 100 mm GB. Drought stress resulted in a remarkable decrease in turf quality (TQ), relative water content (RWC), and chlorophyll content, with significant increases in superoxide anion content (O2–), malondialdehyde (MDA) content, superoxide dismutase (SOD), catalase (CAT), and peroxidase (POD) activity. In contrast, pretreatment with 100 mm GB decreased the O2– and MDA content in water-stressed plants, and increased turf quality, chlorophyll content, SOD, CAT, and POD activity. Meanwhile, the expression level of the psbA, SAMS4, CMO, and ACS1 genes in leaf samples collected during the drought-stress stage was elevated in GB pretreatment. Notably, SAMS4 gene expression in GB pretreatment was significantly greater than in the untreated GB groups subjected to water stress. These results suggested that GB could mitigate the adverse effect of water stress on creeping bentgrass. The amelioration related strongly to the maintenance of the antioxidant enzyme system, accumulated endogenous compatible metabolites, and the elevation of gene expression levels. These findings lead us to conclude that GB pretreatment could be used as an ameliorative agent for creeping bentgrass against the deleterious effects of water stress.

Ethephon [ETH (2-chloroethylphosphonic acid, an ethylene-releasing compound)] has been used as a plant growth regulator in turfgrass management. The aim of the study was to assess the effects of ETH seed treatment on drought tolerance of kentucky bluegrass (Poa pratensis) seedlings. Seeds of two kentucky bluegrass cultivars, Midnight and Nuglade, were exposed to ETH treatment or untreated as controls. Seedlings were then exposed to two water regimes: well-watered conditions and polyethylene glycol (PEG)–induced drought conditions. ETH-treated plants exhibited better turf performance relative to the untreated control under PEG-stressed conditions illustrated by higher relative water content (RWC) and lower lipid peroxidation and lower electrolyte leakage (EL). In both cultivars, ETH treatment increased enzyme activity of ascorbate peroxidase (APX), peroxidase (POD), and catalase (CAT); proline content; and soluble protein content under PEG-induced drought conditions. The results suggest that ETH seed treatment can improve drought tolerance in kentucky bluegrass seedlings.

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 (gs) 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 gs. 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.

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.

Zoysiagrass (Zoysia spp.), a warm-season turfgrass species, experiences freezing damage in many regions. The mechanisms of its cold acclimation and freezing tolerance have not been well understood. This study was designed to investigate effects of cold acclimation treatment on leaf abscisic acid (ABA), cytokinin (transzeatin riboside (t-ZR), and antioxidant metabolism associated with freezing tolerance in zoysiagrass (Zoysia japonica). ‘Chinese Common’ zoysiagrass was subjected to either cold acclimation treatment with temperature at 8/2 °C (day/night) and a photosynthetically active radiation (PAR) of 250 µmol·m−2·s−1 over a 10-hour photoperiod or normal environments (temperature at 28/24 °C (day/night), PAR at 400 µmol·m−2·s−1 and 14-hour photoperiod) for 21 days in growth chambers. Cold treatment caused cell membrane injury as indicated by increased leaf cell membrane electrolyte leakage (EL) and malondialdehyde (MDA) content after 7 days of cold treatment. Cold treatment increased leaf ABA and hydrogen peroxide content and reduced t-ZR content. Leaf superoxide dismutase (SOD), ascorbate peroxidase (APX) activity, and proline content increased, whereas catalase (CAT) and peroxidase (POD) activity declined in response to cold treatment. Cold treatment increased freezing tolerance as LT50 declined from −4.8 to −12.5 °C. The results of this study indicated that cold acclimation treatment might result in increases in ABA and H2O2, which induce antioxidant metabolism responses and improved freezing tolerance in zoysiagrass.

Soil water deficit impacts cold acclimation and freezing tolerance in creeping bentgrass (Agrostis stolonifera L.), but the mechanisms underlying have not been well understood. The objectives of this study were to investigate the effects of deficit irrigation before and during cold acclimation on osmoprotectants, antioxidant metabolism, and freezing tolerance in creeping bentgrass. The grass was subjected to three-soil moisture levels: well-watered [100% container capacity (CC)], deficit irrigation induced-mild drought stress (60% CC), and severe drought stress (30% CC) for 35 days including 14 days at 24/20 °C (day/night) and then 21 days under cold acclimation treatment (2 °C) in growth chambers. Leaf proline and total soluble sugar (TSS) levels were higher in the grass under mild drought stress relative to that under severe drought stress. Superoxide (O2−·), hydrogen peroxide (H2O2), and malondialdehyde (MDA) content were higher in the grass under severe drought relative to that under well-watered and mild drought stress at day 35. Mild drought stress increased catalase (CAT) and guaiacol peroxidase (POD) activity, induced new isoforms and increased band intensities of superoxide dismutase (SOD), CAT, and POD during cold acclimation (days 14 to 35). No differences in osmoprotectants, antioxidant metabolism, and freezing tolerance were found between mild drought and well-watered treatments. The results of this study suggest deficit irrigation-induced mild drought stress in late fall and winter could induce accumulation of osmoprotectants and improve antioxidant metabolism, and freezing tolerance, but severe drought stress could reduce freezing tolerance of creeping bentgrass in the region with limited precipitation.

Tall fescue is an important cool-season grass widely used for forage and turf, and its genotypic variation for morphological traits has not been well documented. One hundred and fifteen tall fescue accessions, including 25 commercial cultivars, were divided into five groups based on their origination. The morphological traits, including plant height, spike length, pulvinus distance, spikelet count, branch count per spike, spike count per plant, and spike weight in different accessions were determined under field conditions in 2013 and 2014. There was significant genotypic variation in morphological traits among the 115 tall fescue accessions. Wild accessions exhibited a greater variation in the morphological traits than commercial cultivars. Close correlations were found among plant height, spike length, pulvinus distance, and spikelet count. The results of this suggest plant height, spike length, pulvinus distance, and spikelet count could be used as key morphological traits for evaluating all fescue germplasm effectively.

Heat stress is a major limiting factor for growth of cool-season perennial grass species, and mechanisms of heat tolerance have not been well understood. This study was designed to investigate antioxidant enzyme and hormone metabolism responses to heat stress in two kentucky bluegrass (Poa pratensis L.) cultivars contrasting in heat tolerance. The plants were subjected to 20/20 °C [day/night (control)] or 38/30 °C [day/night (heat stress)] for 28 days in growth chambers. Heat stress increased leaf electrolyte leakage (EL) and malondialdehyde (MDA) with heat-tolerant cultivar EverGlade exhibiting lower levels of EL and MDA relative to heat-sensitive cultivar Kenblue under heat stress. Superoxide dismutase (SOD) and catalase (CAT) activity increased and then declined during 28 days of heat stress. Peroxidase (POD) and ascorbate peroxidase (APX) activity declined and then increased during heat stress. ‘EverGlade’ had greater activities of SOD, CAT, POD, and APX relative to ‘Kenblue’ under heat stress. In addition, ‘EverGlade’ had two additional SOD isozymes and three additional POD isozymes relative to ‘Kenblue’ under heat stress. Leaf abscisic acid (ABA) increased in response to heat stress. Leaf indole-3-acetic acid (IAA) increased and then declined during heat stress. ‘OverGlade’ had higher ABA and IAA content relative to ‘Kenblue’. At the end of heat stress, leaf IAA and ABA content were 27.8% and 73% higher in ‘EverGlade’ relative to ‘Kenblue’, respectively. The results indicated that antioxidant enzymes and the hormones (ABA and IAA) were associated with kentucky bluegrass heat tolerance. Selection and use of cultivars with higher IAA and ABA content and greater antioxidant enzyme activities may improve kentucky bluegrass growth and quality under heat stress.

Biosolids are valued as a source of plant nutrients, soil organic matter, and biologically active substances. This greenhouse study was designed to examine if application of biosolids can improve plant drought tolerance by affecting nitrogen (N) and hormone metabolism as well as root growth in kentucky bluegrass (Poa pratensis L.; KBG). The three treatments, which provided N rates equivalent to 75 mg plant-available N/kg soil, included: 1) biosolids at 1× agronomic (Ag) N rate (75 mg N/kg soil completely provided with biosolids); 2) biosolids at 0.5× Ag N rate (37.5 mg N/kg soil provided with biosolids and 37.5 mg N/kg soil provided with NH4NO3); and 3) control (75 mg N/kg soil completely provided with NH4NO3). The treated KBG was grown under either well-watered (90% container capacity) or drought stress (≈25% container capacity) conditions. Biosolids application improved turf quality and delayed leaf wilting under drought stress. The grass treated with biosolids at 1× N rate had higher leaf proline and amino acid content and greater nitrate reductase activity than the control under drought stress. Biosolids treatments also increased leaf and soil indole-3-acetic acid (IAA) content. Moreover, biosolids at 1× N rate increased root length density by 23% compared with the control under drought stress. The results of this study suggest that biosolids may enhance plant drought tolerance by improving N and hormone metabolism and root growth in KBG.

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.

Drought stress is one of the major limiting factors for plant growth and development. The mechanism of drought tolerance has not been well understood. This study was designed to investigate proline and antioxidant metabolism associated with drought tolerance in transgenic tobacco (Nicotiana tabacum) plants overexpressing the OjERF gene relative to wild-type (WT) plants. The OjERF gene was isolated from mondo grass (Ophiopogon japonicus). The OjERF gene, driven by the CaMV35S promoter, was introduced into tobacco through agrobacterium (Agrobacterium tumefaciens)-mediated transformation. Five transgenic lines were regenerated, of which transgenic Line 5 (GT5) and Line 6 (GT6) were used to examine drought tolerance in comparison with WT plants in a growth chamber. Drought stress caused an increase in leaf malondialdehyde (MDA) and electrolyte leakage (EL), proline content, superoxide dismutase (SOD), and catalase (CAT) activity in both transgenic lines and WT plants. However, the transgenic lines had lower MDA content and EL and higher proline content, SOD and CAT activity relative to WT under drought stress. The activities of SOD and CAT were also greater in the transgenic lines relative to WT plants under well-watered conditions (Day 0). The OjERF activated the expression of stress-relative genes, including NtERD10B, NtERD10C, NtERF5, NtSOD, and NtCAT1 in tobacco plants. The results of this study suggest that the OjERF gene may confer drought stress tolerance through upregulating proline and antioxidant metabolism.

Salinity stress may involve the accumulation of glycine betaine (GB). The objective of this study was to examine whether exogenous GB would ameliorate the detrimental effect of salinity stress on perennial ryegrass (Lolium perenne). The grass was subjected to two salinity levels (0 and 250 mm NaCl) and three GB levels (0, 20, and 50 mm). Salinity resulted in a remarkable decrease in vertical shoot growth rate (VSGR), shoot and root fresh weight, relative water content (RWC), relative transpiration rate (Tr), and chlorophyll (Chl) content, superoxide dismutase (SOD), catalase (CAT), and ascorbate peroxidase (APX) activities. Plants subjected to salt exhibited an increase in leaf electrolyte leakage (EL), lipid peroxidation (MDA), and proline content. Application of GB reduced EL, MDA, and proline content in salt-stressed plants. Perennial ryegrass subjected to salt stress plus GB had a greater level of VSGR, RWC, relative Tr, Chl content, and activities of SOD, CAT, and APX when compared with salt-stressed without GB. Salt stress increased Na+ and decreased K+ content, which resulted in a higher Na+/K+ ratio in perennial ryegrass. Application of 20 mm GB suppressed Na+ accumulation, whereas the K+ content was significantly increased in shoot, which led to a higher K+/Na+ ratio under saline conditions. These results suggested that GB-enhanced salt tolerance in perennial ryegrass was mainly related to the elevated SOD, CAT, and APX activity and alleviation of cell membrane damage by reducing oxidation of membrane lipid and improving the ion homeostasis under salt stress.

Drought stress is a major factor in turfgrass management; however, the underlying mechanisms of turfgrass drought tolerance are not well understood. This greenhouse study was designed to investigate proline and hormone responses to drought stress in two tall fescue [Festuca arundinacea (Schreb.)] cultivars differing in drought tolerance. The two cultivars, Van Gogh (relatively drought-tolerant) and AST7002 (relatively drought-sensitive), were established and grown under either well-watered (maintaining 90% container capacity) or drought stress (≈26% container capacity) and then re-watered. Drought stress reduced turfgrass quality, relative leaf water content (LWC), leaf indole-3-acetic acid (IAA) and cytokinin zeatin riboside (ZR) content, and increased proline and abscisic acid (ABA) content. ‘Van Gogh’ had greater turfgrass quality rating, LWC, proline, ABA, and ZR content relative to ‘AST7002’ under drought stress conditions. At the end of drought stress, leaf proline, ZR, and ABA content were 32%, 43%, and 50% higher in ‘Van Gogh’ relative to ‘AST7002’, respectively. No cultivar difference was observed under well-watered conditions. The results of this study suggest that the proline, ABA, and ZR content are associated with drought tolerance. Selection and use of the cultivars with higher proline, ABA, and ZR content under drought stress may be a practical approach to improve tall fescue drought tolerance.

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.

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.

Traffic stress causes turfgrass injury and soil compaction but the underlying physiological mechanisms are not well documented. The objectives of this study were to investigate the physiological responses of kentucky bluegrass (Poa pratensis), tall fescue (Festuca arundinacea), and japanese zoysiagrass (Zoysia japonica) to three levels of traffic stress during the growing season under simulated soccer traffic conditions. Relative leaf water content (LWC), shoot density, leaf chlorophyll concentration (LCC), membrane permeability, and leaf antioxidant peroxidase (POD) activity were measured once per month. The traffic stress treatments caused a reduction in LWC, shoot density, LCC, and POD activity, and an increase in cell membrane permeability in all three species. Japanese zoysiagrass had less electrolyte leakage, and higher POD activity and shoot density than both kentucky bluegrass and tall fescue. The results suggest that turfgrass tolerance to traffic stress may be related to leaf antioxidant activity. Turfgrass species or cultivars with higher leaf antioxidant activity may be more tolerant to traffic stress than those with lower antioxidant activity.

Ultraviolet-B [UV-B (280-320 nm)] radiation is one of the major factors causing quality decline of transplanted sod. Pigments and antioxidants are associated with plant stress resistance, but their roles in turfgrass tolerance to UV-B damage are not well understood. The objectives of this study were to determine if kentucky bluegrass (Poa pratensis L.) cultivars with darker green genetic leaf color possessed greater pigment and antioxidant defense capacities and if such characteristics were associated with greater resistance to UV-B. Two cultivars, `Moonlight' (dark green) and `Limerick' (light green), were selected and subjected to continuous, artificial UV-B radiation (70 μmol·m-2·s-1). UV-B irradiation reduced turf quality by 58% (`Moonlight') and 77% (`Limerick') relative to day 1 when measured 10 days after initiation of UV-B exposure. Higher canopy photochemical efficiency (PEc) was found in `Moonlight' relative to `Limerick' under UV-B stress and during recovery. `Moonlight' contained greater levels of chlorophyll (1.5 to1.6-fold), carotenoids (1.3-fold), superoxide dismutase [SOD (1.0-fold)] and catalase [CAT (1.5-fold)] than `Limerick' when measured at 10 days after UV-B initiation. Turfgrass quality and PEc were positively correlated with pigments (chlorophyll and carotenoids) and antioxidant enzymes (SOD and CAT), and negatively correlated with lipid peroxidation. The results suggest that selecting dark-green cultivars with greater pigment content and antioxidant activity may be an effective approach for turfgrass breeders and sod producers to improve tolerance of newly transplanted sod to environments with higher UV-B radiation.

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.

Additional index words. alpha-tocopherol, antioxidant enzyme activity, ascorbic acid, ascorbate peroxidase, catalase, chlorophyll fl uorescence, photochemical effi ciency, superoxide dismutase, turf injury Abstract. High ultraviolet-B (UV-B; 290-320nm wavelength) may signifi cantly 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 detoxifi cation 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 detoxifi cation systems of kentucky bluegrass could be artifi cially 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 ap- plied to plugs of mature kentucky bluegrass alone or in combination, and then subjected to artifi cial, 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 effi ciency, as measured by chlorophyll fl uorescence were signifi cantly reduced. However, in Expt. 1, AA alleviated decline of visual quality, delayed loss of photochemical effi ciency, 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 effi ciency 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 toler- ance may be increased by supplementing its pigment and antioxidant defense systems with foliar applications of AA, AT or GL.

Bermudagrass [Cynodon dactylon (L.) Pers.] is widely used along its northern limit of adaptation. However, cold hardiness and winter survival are common concerns facing turfgrass managers in these areas. The objective of this study was to determine the effects of moderate salinity applications on bermudagrass cold hardiness. Two trials were conducted in Summer 2002. The cultivar Princess was seeded into pots in a glasshouse at a rate of 24 kg·ha-1. Pots received a weekly solution of 20-20-20 at a rate of 4.9 kg·ha-1 N. Bi-weekly salinity treatments began ≈2 months after germination and consisted of 0, 5, 20, and 40 dS·m-1 in the form of NaCl. These treatments continued for ≈8 weeks. Weekly quality ratings and chlorophyll fluorescence measurements showed similar results, with the high salinity treatments having the poorest quality. Soil electrical conductivity measurements showed a significant increase for the high salinity rates over the lower rates at the end of the trials. Proline concentrations increased with increasing salinity treatments in Trial 1 and were highest with the 20 dS·m-1 rate in Trial 2. Plants were acclimated in a growth chamber, and artificial freezing tests revealed that the 5 and 20 dS·m-1 treatments had the highest percentage of regrowth after freezing. These results indicate that moderate applications of salt or the use of effluent water prior to hardening may be an important way to increase bermudagrass cold hardiness.

Creeping bentgrass (Agrostis stolonifera) is used extensively on temperate zone golf course greens, tees, and fairways, but often performs poorly in shade. Previous research has indicated that sequential applications of gibberellic acid (GA) inhibiting plant growth regulators (PGRs) such as trinexapac-ethyl (TE) increase cool-season turfgrass performance in 70-90% shade. This research was conducted to: 1) confirm appropriate TE application rates and frequencies for maintaining `Penncross' creeping bentgrass in dense shade in the mid-Atlantic region of the U.S.; 2) determine the efficacy of other PGRs, biostimulants, and iron (Fe); and 3) assess whether the addition of a biostimulant with TE would have additive, synergistic, or negative effects. The other compounds tested against TE and the control were: propiconazole (PPC), iron sulfate, CPR (a seaweed and iron containing biostimulant), and a generic seaweed extract (SWE) (Ascophyllum nodosum) plus humic acid (HA) combination. These treatments were applied to 88% shaded bentgrass every 14 days from May or June through October in 2001 and 2002, with turf quality, leaf color, root strength, photochemical efficiency, and antioxidant enzyme superoxide dismutase (SOD) activity being determined. While the quality of control plots fell below a commercially acceptable level by the second month of the trial, repeated foliar TE application provided 33% to 44% better quality throughout the experiment. Propiconazole resulted in 13% to 17% better quality through September of each year. Trinexapac-ethyl and PPC resulted in darker leaf color and increased mid-trial root strength by 27% and 29%, respectively. Canopy photochemical efficiency and leaf SOD activity were also increased due to TE in August of both years. Treatment with Fe, CPR, or SWE+HA did not have an effect on quality, root strength, SOD, or photochemical efficiency, but periodic increases in color were observed. The addition of CPR to TE in 2002 provided results that were not different from those of TE-alone. This and previous studies indicate that restricting leaf elongation with anti-GA PGRs is of primary importance for improving shade tolerance, while treatments that increase leaf color or chlorophyll levels without restricting leaf elongation are relatively ineffective.

A variety of organic materials such as humic substances, seaweed extracts (SWE), organic matter, and amino acids are being used as fertilizer supplements in commercial turfgrass management. Among them, SWE and humic acid (HA) are widely used in various biostimulant product formulations. These compounds have been reported to contain phytohormones and osmoprotectants such as cytokinins, auxins, polyamines, and betaines. Manufacturer claims are that these products may supplement standard fertility programs by reducing mineral nutrient requirements while improving stress tolerance. There is a lack of season-long, fi eld-based evidence to support these claims. This study was conducted to investigate the infl uence of monthly fi eld applications of SWE, HA, and high and low seasonal fertilization regimes on the physiological health of fairway-height creeping bentgrass (Agrostis stolonifera L.). Plots were treated monthly with SWE at 16 mg·m -2 and HA (70% a.i.) at 38 mg·m -2 alone, or in combination, and were grown under low (20 kg·ha -1 /month) or high nitrogen (50 kg·ha -1 /month) fertilization regimes during 1996 and 1997. Endogenous anti- oxidant superoxide dismutase (SOD) activity, photochemical activity (PA), and turf quality were measured in July of each year. Superoxide dismutase activity was increased by 46% to 181%, accompanied by a PA increase of 9% to 18%, and improved visual quality of bentgrass in both years. There was no signifi cant fertilization × supplement interaction. Although not part of our original objectives, it was noted that signifi cantly less dollar spot (Sclerotinia homoeocarpa F.T. Bennett) disease incidence occurred in supplement-treated bentgrass. Our results indicate that increased SOD activity in July due to SWE and/or HA applications improved overall physiological health, irrespective of fertiliza- tion regime. This suggests that these compounds may be benefi cial supplements for reducing standard fertilizer and fungicide inputs, while maintaining adequate creeping bentgrass health. mic acid (HA) are two novel materials that have shown promise for protecting turfgrasses against oxidative stress. Much early work by Schmidt and collaborators focused on refi ning appropri- ate dilution rates of seaweed extracts and HA based on turfgrass responses such as leaf growth rate and senescence (Goatley and Schmidt, 1990), nutrient uptake (Yan, 1993), root mass (Nabati et al., 1994), and photochemical activity (Zhang, 1997). These studies have shown that certain rates of generic and commercial formulations of SWE and/or HA can improve turfgrass quality and resistance to environmental stresses such as drought (Zhang, 1997) and salinity (Nabati, et al., 1994). Further research indicated that improved stress resistance was associated with increases in anti- oxidant contents and activities (Zhang and Schmidt, 1997; Zhang and Schmidt, 1999, 2000b). All of these studies were conducted under uniform fertilization. Because the chemical components of SWE and HA have yet to be completely characterized and tested for specifi c activity, we do not currently understand the physiological or biochemical bases of SWE and/or HAʼs effects on antioxidants and other stress responses. However, information from other researchers has provided some clues as to constituents and possible modes of action. Fike et al. (2001) reported that SWE derived from A. no- dosum contains various compounds including amino acids and micronutrients; they also reported hormonal activity equivalent to 50 mg·L-1 kinetin. Additionally, auxin and cytokinins have been identifi ed and quantifi ed in SWE using GC-MS techniques (Crouch and Van Staden, 1993; Sanderson and Jameson, 1986; Sanderson et al., 1987; Senn, 1987). Lastly, three betaine forms have been quantifi ed in A. nodosum extracts (Blunden et al., 1986). Quarternary ammonium compounds such as the betaines are thought to play a pivotal role in plant cytoplasmic adjustment

Decline of sod quality during the transportation, storage, and transplant stages of sale is a primary economic concern of sod producers. However, the mechanisms of extending sod quality during storage, transportation, and transplantation remain unclear. This study was conducted to investigate the influences of selected plant metabolic enhancers (PMEs) seaweed (Ascophyllum nodosum Jol.) extract (SWE), humic acid [93% a.i. (HA)], and propiconazole (PPC), on sod tolerance to stress during storage and posttransplant root growth of tall fescue (Festuca arundinacea Schreb.) sod. The SWE + HA, and PPC were applied alone, or in a combination, to tall fescue 2 weeks before harvest. Photochemical efficiency (PE) of photosystem II was measured immediately before harvest. The harvested sod was subjected to high temperature stress (40 °C) for 72 or 96 hours. The heated sod was replanted in the field and posttransplant injury and root strength were determined. On average over 1999 and 2000, application of SWE (50 mg·m-2) + HA (150 mg·m-2), PPC (0.30 mL·m-2), and a combination of SWE + HA with PPC (0.15 mL·m-2), enhanced PE of preharvest sod by 8.5%, 9.1%, and 11.2%, respectively, and increased posttransplant rooting by 20.6%, 34.6%, and 20.2%, respectively. All PME treatments reduced visual injury except SWE + HA and SWE + HA + PPC in 1999. Extension of heat duration from 72 to 96 hours caused significantly more injury to the sod and reduced posttransplant rooting by 22.9% averaged over 2 years. The data suggest that foliar application of SWE + HA, PPC alone, or in a combination with SWE + HA, may reduce shipment heat injury and improve posttransplant rooting and quality of tall fescue sod. Chemical name used: 1-(2-(2,4-dichloropheny)-4-propyl-1,3-dioxolan-2yl)methyl-1-H-1,2,4-triazole [propiconazole (PPC)].

Creeping bentgrass (Agrostis palustris Huds.) is an extensively used cool-season grass for fine turf areas such as golf course putting greens, but suffers from poor summer stress tolerance. These studies were conducted to investigate the influences of natural plant growth regulators (NPGR) and Fe on creeping bentgrass photochemical activity (PA), antioxidant superoxide dismutase (SOD) activity, root growth and leaf color under two fertilization regimes. The bentgrass was maintained in well-watered field conditions or water-stressed glasshouse conditions. A mature bentgrass was treated monthly during the field season with seaweed (Ascophyllum nodosum Jol.) extract (SWE) at 50 mg·m-2 or humic acid (HA) at 150 mg·m-2 or in combination with or without FeSO4 at 520 mg·m-2 and grown under a low or a high fertilization regime. Foliar application of SWE + Fe increased PA (14% to 15%), while applications of SWE + HA or SWE + HA + Fe increased SOD activity (49% to 114%) of creeping bentgrass in Summer 1997 and Summer 1998. There was no significant fertilization × NPGR interaction for PA and SOD activity. Bentgrass PA was increased by 13% to 46% when treated with NPGR with or without Fe compared to the control measured in May. The addition of Fe with each NPGR application improved fall and winter leaf color. All NPGR and Fe treatments increased root mass (17% to 29%) in Aug. 1997 and 1998, except HA alone in 1998. Under sustained low soil moisture (-0.5 MPa) conditions, application of NPGR with or without Fe increased PA and SOD activity. The data indicate that SWE and HA enhance the physiological function of `Southshore' creeping bentgrass, resulting in improved root growth regardless of low or high fertilization regime. However, addition of Fe to these NPGR served primarily to improve late season leaf color. The results suggest that, in addition to maintaining adequate plant-available nutrients, applications of natural PGRs, such as SWE and HA, prior to and during summer abiotic stresses would be beneficial.

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.